n Encyclopedia of Invasive

Species
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n Encyclopedia
of Invasive Species
From Africanized Honey Bees
to Zebra Mussels
Volume 1: Animals

Susan L. Woodward and Joyce A. Quinn

Copyright 2011 by ABC-CLIO, LLC
All rights reserved. No part of this publication may be reproduced, stored in a retrieval
system, or transmitted, in any form or by any means, electronic, mechanical,
photocopying, recording, or otherwise, except for the inclusion of brief quotations in a
review, without prior permission in writing from the publisher.
Library of Congress Cataloging-in-Publication Data

Woodward, Susan L., 1944 Jan. 20–

Encyclopedia of invasive species : from africanized honey bees to zebra mussels / Susan L. Woodward and
Joyce A. Quinn.
p. cm.
Includes bibliographical references and index.
ISBN 978–0–313–38220–8 (cloth : alk. paper) — ISBN 978–0–313–38221–5 (ebook)
1. Introduced organisms—Encyclopedias. I. Quinn, Joyce Ann. II. Title.
QH353.W66 2011
578.60 2—dc23 2011026543

ISBN: 978–0–313–38220–8
EISBN: 978–0–313–38221–5
15 14 13 12 11 1 2 3 4 5
This book is also available on the World Wide Web as an eBook.
Visit www.abc-clio.com for details.
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An Imprint of ABC-CLIO, LLC
ABC-CLIO, LLC
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This book is printed on acid-free paper
Manufactured in the United States of America
n Contents
General Introduction: Invasive Species—Concepts and Issues n xiii

VOLUME 1: INVASIVE MICROORGANISMS, FUNGI,

AND ANIMALS
Preface n xxxv

Alphabetical List of Invasive Microorganisms, Fungi, and Animal

Entries n xxxix

Microorganisms
Avian Malaria (Plasmodium relictum capistranoae) n 1
Lyme Disease Bacterium (Borrelia burgdorferi) n 3
West Nile Virus (West Nile Virus) n 7

Fungi
Bat White-Nose Syndrome Fungus (Geomyces destructans) n 11
Chestnut Blight Fungus (Cryphonectria parasitica) n 14
Chytrid Frog Fungus (Batrachochytrium
dendrobatidis) n 18
Dutch Elm Disease Fungi (Ophiostoma novo-ulmi
and O. ulmi) n 21
Sudden Oak Death (Phytophthora ramorum) n 25
White Pine Blister Rust (Cronartium ribicola) n 29
Invertebrates
Bryozoan
Lacy Crust Bryozoan (Membranipora membranacea) n 36
Tunicates
Chain Tunicate (Botrylloides violaceus) n 39
Colonial Tunicate (Didemnum vexillum) n 42
Cnidarian
Australian Spotted Jellyfish (Phyllorhiza punctata) n 45
vi n CONTENTS

Annelid worms
European Earthworms (Lumbricus terrestris, L. rubellus,
Aporrectodea caliginosa, Dendrobaena octaedra, and
others) n 48
Mollusks
Asian Clam (Corbicula fluminea) n 53
Asian Green Mussel (Perna viridis) n 56
Chinese Mystery Snail (Cipangopaludina
chinensis malleata) n 58
Common Periwinkle (Littorina littorea) n 61
Giant African Snail (Achatina fulica) n 64
Golden Apple Snail (Pomacea canaliculata) n 67
Naval Shipworm (Teredo navalis) n 70
New Zealand Mud Snail (Potamopyrgus
antipodarum) n 73
Quagga Mussel (Dreissena rostriformis bugensis) n 76
Veined Rapa Whelk (Rapana venosa) n 79
Zebra Mussel (Dreissena polymorpha) n 82
Crustaceans
Chinese Mitten Crab (Eriocheir sinensis) n 86
Green Crab (Carcinus maenas) n 90
Rusty Crayfish (Orconectes rusticus) n 93
Spiny Water Flea (Bythotrephes longimanus) n 95
Arachnids
Honeybee Tracheal Mite (Acarapis woodi) n 99
Varroa Mite (Varroa destructor) n 102

Insects
Africanized Honey Bee (Apis mellifera scutellata) n 106
Argentine Ant (Linepithema humile) n 110
Asian Longhorned Beetle (Anoplophora
glabripennis) n 113
Asian Tiger Mosquito (Aedes albopictus) n 116
Brown Marmorated Stink Bug (Halyomorpha halys) n 120
Common Bed Bug (Cimex lectularius) n 123
Emerald Ash Borer (Agrilus planipennis) n 127
 CONTENTS n vii

Formosan Subterranean Termite (Coptotermes

formosanus) n 131
Glassy-Winged Sharpshooter (Homalodisca vitripennis) n 134
Gypsy Moth (Lymantria dispar) n 138
Hemlock Woolly Adelgid (Adelges tsugae) n 142
Japanese Beetle (Popillia japonica) n 146
Multicolored Asian Lady Beetle (Harmonia axyridis) n 148
Red Imported Fire Ant (Solenopsis invicta) n 152

Vertebrates
Fish
Alewife (Alosa pseudoharengus) n 157
Asian Swamp Eel (Monopterus albus) n 160
Bighead Carp (Hypophthalmichthys nobilis) n 163
Brown Trout (Salmo trutta) n 166
Gizzard Shad (Dorosoma cepedianum) n 168
Grass Carp (Ctenopharyngodon idella) n 172
Lionfish (Pterois volitans / P. miles) n 175
Mosquitofish (Gambusia affinis and G. holbrooki) n 178
Northern Snakehead (Channa argus) n 182
Rainbow Trout (Oncorhynchus mykiss) n 185
Round Goby (Neogobius melanostomus) n 187
Sea Lamprey (Petromyzon marinus) n 190
Silver Carp (Hypophthalmichthys molitrix) n 194
Spotted Tilapia (Tilapia mariae) n 196
Walking Catfish (Clarias batrachus) n 198
Amphibians
African Clawed Frog (Xenopus laevis) n 201
American Bullfrog (Lithobates catesbeianus) n 205
Coqui (Eleutherodactylus coqui) n 208
Cuban Treefrog (Osteopilus septentrionalis) n 211
Reptiles
Brown Anole (Norops [=Anolis] sagrei) n 214
Burmese Python (Python molurus bivittatus) n 217
Green Iguana (Iguana iguana) n 221
viii n CONTENTS

Nile Monitor (Varanus niloticus) n 225

Birds
Cattle Egret (Bubulcus ibis) n 228
Common Myna (Acridotheres tristis) n 232
Eurasian Collared-Dove (Streptopelia decaocto) n 234
European Starling (Sturnus vulgaris) n 237
House Finch (Carpodacus mexicanus) n 240
House Sparrow (Passer domesticus) n 243
Japanese White-Eye (Zosterops japonicus) n 246
Monk Parakeet (Myiopsitta monachus) n 248
Mute Swan (Cygnus olor) n 252
Rock Pigeon (Columba livia) n 256
Mammals
Black Rat (Rattus rattus) n 259
Feral Burro (Equus asinus) n 262
Feral Cat (Felis silvestris catus) n 265
Feral Goat (Capra hircus) n 268
Feral Horse (Equus caballus) n 271
Feral Pig (Sus scrofa) n 275
House Mouse (Mus musculus) n 281
Indian Mongoose (Herpestes javanicus) n 284
Norway Rat (Rattus norvegicus) n 287
Nutria (Myocastor coypus) n 290
State-by-State Occurrences of Invasive Microorganisms, Fungi, and
Animals n 295

Glossary n 311
Index n I-1

VOLUME 2: INVASIVE PLANT SPECIES

Preface n xiii

Invasive Plants in the United States: A Brief Overview n xvii

Alphabetical List of Invasive Plant Entries n xxiii
 CONTENTS n ix

Aquatic Plants
Eurasian Watermilfoil (Myriophyllum spicatum) n 321
Giant Salvinia (Salvinia molesta) n 326
Hydrilla (Hydrilla verticillata) n 331
Water Chestnut (Trapa natans) n 335
Waterhyacinth (Eichhornia crassipes) n 339

Forbs
Canada Thistle (Cirsium arvense) n 344
Chinese Lespedeza (Lespedeza cuneata) n 349
Common Mullein (Verbascum thapsus) n 353
Common St. Johnswort (Hypericum perforatum) n 358
Dyer’s Woad (Isatis tinctoria) n 362
Fig Buttercup (Ficaria verna) n 366
Garlic Mustard (Alliaria petiolata) n 369
Giant Hogweed (Heracleum mantegazzianum) n 372
Goutweed (Aegopodium podagraria) n 376
Halogeton (Halogeton glomeratus) n 379
Ice Plant and Crystalline Ice Plant (Carpobrotus edulis
and Mesembryanthemum crystallinum) n 383
Japanese Knotweed (Fallopia japonica) n 387
Kahili Ginger (Hedychium gardnerianum) n 391
Leafy Spurge (Euphorbia esula) n 395
Musk Thistle (Carduus nutans) n 399
Perennial Pepperweed and Hoary Cress (Lepidium
latifolium and Cardaria draba) n 404
Prickly Russian Thistle (Salsola tragus) n 409
Purple Loosestrife (Lythrum salicaria) n 414
Spotted Knapweed (Centaurea stoebe) n 417
Toadflax (Linaria dalmatica ssp. dalmatica and
Linaria vulgaris) n 421
Yellow Starthistle (Centaurea solstitialis) n 427
Graminoids
Asiatic Sand Sedge (Carex kobomugi) n 432
Buffelgrass (Pennisetum ciliare) n 435
x n CONTENTS

Cheatgrass (Bromus tectorum) n 439

Cogongrass (Imperata cylindrica) n 443
Common Reed (Phragmites australis ssp. australis) n 447
Cordgrasses and Their Hybrids (Spartina alterniflora, Spartina
densiflora, Spartina patens, Spartina anglica, and Spartina
alterniflora x foliosa n 452
Crimson Fountain Grass (Pennisetum setaceum) n 458
Giant Reed (Arundo donax) n 462
Japanese Stilt Grass (Microstegium vimineum) n 466
Johnsongrass (Sorghum halepense) n 469
Jubata Grass and Pampas Grass (Cortaderia jubata and Cortaderia
selloana) n 472
Kikuyugrass (Pennisetum clandestinum) n 478
Medusahead (Taeniatherum caput-medusae) n 481
Quackgrass (Elymus repens) n 485
West Indian Marsh Grass (Hymenachne
amplexicaulis) n 489

Shrubs
Asiatic Colubrina (Colubrina asiatica) n 493
Brooms (Cytisus scoparius, Spartium junceum, Genista
monspessulana, and Cytisus striatus) n 496
Exotic Bush Honeysuckles (Lonicera maackii, L. morrowii,
L. tatarica, and L. x bella) n 502
Gorse (Ulex europaeus) n 508
Japanese Barberry (Berberis thunbergii) n 512
Koster’s Curse (Clidemia hirta) n 515
Lantana (Lantana camara) n 518
Multiflora Rose (Rosa multiflora) n 522
Rattlebox (Sesbania punicea) n 527
Tropical Soda Apple (Solanum viarum) n 530
Yellow Himalayan Raspberry (Rubus ellipticus) n 535
Trees
Australian Pine (Casuarina equisetifolia) n 540
Brazilian Peppertree (Schinus terebinthifolius) n 544
 CONTENTS n xi

Carrotwood (Cupaniopsis anacardioides) n 548

Chinaberry (Melia azedarach) n 551
Fire Tree (Morella faya) n 554
Melaleuca (Melaleuca quinquenervia) n 557
Paper Mulberry (Broussonetia papyrifera) n 562
Princess Tree (Paulownia tomentosa) n 565
Russian Olive (Elaeagnus angustifolia) n 568
Silk Tree (Albizia julibrissin) n 572
Strawberry Guava (Psidium cattleianum) n 576
Tamarisk (Tamarix chinensis T. ramosissima,
T. parviflora, and T. gallica) n 579
Tree of Heaven (Ailanthus altissima) n 585
Velvet Tree (Miconia calvescens) n 589

Vines
Chocolate Vine (Akebia quinata) n 594
Climbing Ferns (Lygodium japonicum and Lygodium micro-
phyllum) n 597
English Ivy (Hedera helix) n 602
Field Bindweed (Convolvulus arvensis) n 606
Japanese Dodder (Cuscuta japonica) n 610
Japanese Honeysuckle (Lonicera japonica) n 614
Japanese Hops (Humulus japonicus) n 618
Kudzu (Pueraria montana) n 622
Mile-A-Minute (Persicaria perfoliata) n 626
Oriental Bittersweet (Celastrus orbiculatus) n 629
Porcelainberry (Ampelopsis glandulosa var. brevipedunculata) n
633
Swallow-Worts (Cynanchum rossicum and Cynanchum louiseae) n
636
Winter Creeper (Euonymus fortunei) n 640
Wisteria (Wisteria sinensis and Wisteria floribunda) n 644

Tables and Lists about Invasive Plants

Common Names and Scientific Names n 649
State-by-State Designations of Invasive or Noxious Weeds n 665
xii n CONTENTS

Pathways of Introduction for Plants n 673

Impacts of Invasive Plants n 678
Major Organizations and Publications Concerned about Invasive
Plants n 687
Plant Species Listed as Invasive or Noxious by Organizations and
State and Federal Governments n 689
Set Appendices

Appendix A: American Species That Are Invasive Abroad n 695

Appendix B: Major Federal Legislation and Agreements Pertaining to

Invasive Species n 699
Appendix C: Selected International Agreements and Conventions
Pertaining to Invasive Species n 707

Appendix D: ISSG’s 100 of the World’s Worst Invasive Alien

Species n 710
Glossary n 713
General Bibliography: Selected Classic and Contemporary Works and
Major Internet Data Sources n 723
Index n 727
n General Introduction: Invasive
Species—Concepts and Issues

Just What Is an Invasive Species?

Many different words are used to refer to those species that cause great concern among land
managers, ecologists, and ordinary folk dealing with the consequences of organisms that
have been transported from their places of origin and released to go wild in the waters, for-
ests, grasslands, and deserts of the United States. Some are synonyms, others are not. Some
terms may have subtle value-laden connotations, while others are attempts at scientific
objectivity. “Plant people” use different words than “animal people.” There is as yet no con-
sensus among scientists on what are the best terms to use, so the literature remains inconsis-
tent and definitions sometimes vague.
Several terms merely imply populations existing outside their native range and can
be used more or less interchangeably: alien, exotic, nonnative, and nonindigenous. The
term “alien” may be viewed as pejorative, causing unwarranted bias—and unnecessary
actions—against all foreign species. Nonetheless, the official legal definition of “alien” was
put forth by President Clinton’s Executive Order 13112 (February 3, 1999), which states
that an alien species is, “with respect to a particular ecosystem, any species, including
its seeds, eggs, spores, or other biological material capable of propagating that species, that
is not native to that ecosystem.” This means that a species native to one part of the United
States can be alien to another region of the country, as is the case with so-called native trans-
plants or “domestic exotics” such as the American bullfrog (Lithobates catesbeianus) in
western waterways, smooth cordgrass (Spartina alterniflora) in western salt marshes, or the
House Finch (Carpodacus mexicanus) in eastern states.
What constitutes a native species is open to question. Native species are generally consid-
ered “natural” and in their place of origin, i.e., the place where they belong. In the United
States, native species are often those considered to have been here when Europeans first
colonized the North American continent in the sixteenth century. This qualification, how-
ever, is based on two questionable concepts: (1) that North American environments were
pristine before the arrival of Europeans in the fifteenth century, and (2) that only habitats
not influenced by human activity are natural. Realizing that aboriginal peoples may have
had profound impacts on North American ecosystems through their use of fire, their hunt-
ing, their trade, and their agriculture, some argue that the date dividing pristine from dis-
turbed environments should be pushed back a few tens of thousands of years to when
humans—the most invasive species of all—first set foot on the continent.
Biogeographers tend to define native range as that geographic region where a species
evolved to its contemporary form and to which it was long restricted by natural barriers.
In the history of life, overcoming such barriers is a common occurrence, as is the removal
of barriers. Species constantly disperse from their place of origin as their populations grow,
as they evolve, as continents move and oceans shrink, as drainage systems pirate headwater
streams from other river systems, and so forth. Long-distance migrant species populated dis-
tant oceanic islands and led to the exchange of plants and animals among continental
xiv n GENERAL INTRODUCTION

About Range Maps

R ange maps are usually constructed from records of species occurrence compiled
in museum and herbaria collections. Each verified specimen is represented by a
dot on a map, and usually the outermost dots are connected to form the range
boundary. Adjustments are often made based on knowledge of the specific envi-
ronmental conditions of the area and of the habitat requirements and ecological
tolerances of the species in question. A given species thus does not occur every-
where in its mapped range, but only where conditions are suitable within the des-
ignated region.
Information on the distribution of many species is often a matter of presence or
absence in a particular area. Much of the information available for invasive plants,
especially, comes as records of occurrence (or absence) in a political unit such as an
entire state. Maps based on this information necessarily provide highly generalized
distribution patterns. Larger political units, such as Texas or China, mean even
more generalization. Absence of a species according to the maps may be because
it is not yet recognized or recorded at a particular location.
In order to present the best possible depiction of a species’ range the authors
have consulted a variety of sources and synthesized often-conflicting information.
Native ranges may be especially problematic because biological surveys are not
advanced in many parts of the world and reliable range maps did not exist. In many
cases the place of origin of a species that is invasive in the United States is simply
not known or at best uncertain. Such instances are indicated by question marks
on the maps. Plants, animals, fungi, and microorganisms have been carried around
the world with humans for millennia, obscuring the history of their origins. Some
have undergone subsequent evolution that complicates the picture further.
Increasingly, genetic studies are confirming earlier scientific conclusions or shed-
ding new light on the past travels of species and pinpointing their starting points
with increased accuracy.
Maps, like the species themselves, are dynamic. As new sightings and collections
are made, the range maps may change. The reader wanting the most up-to-date
information or more detailed information than can be derived from the small
maps in this work is encouraged to seek out recent publications and local experts
in the field.
When comparing maps from different sources, the reader is cautioned that the
appearance of distribution areas will vary when different map projections—meth-
ods of converting the spherical globe into a flat map—are used. Distortions in size,
shape, or cardinal direction (N, S, E, W) are inevitable. The projections chosen for
this volume (Eckert III for the world and Albers Equal-Area Conic for the United
States) display fairly accurately true area or size relationships with minimal shape
distortion. Areas closer to polar regions, as indicated by Greenland, are increasingly
more distorted in shape and direction.

landmasses many times in the geological past. Species are still on the move, actively and
passively, in the same manner as in the past. Those that arrive “on their own,” such as the
Cattle Egret (Bubulcus ibis), are referred to as adventive species. The vast majority of species
today, however, are transported, deliberately or accidentally, by humans. Such newcomers
to a region are known as introduced species.
 GENERAL INTRODUCTION n xv

Not all nonnative species are invasive; in fact, most are not. Legally, that term should be
reserved for “an alien species whose introduction does or is likely to cause economic or envi-
ronmental harm or harm to human health” (Executive Order 13112 of February 3, 1999). A
legally defined term that predates “invasive” is nuisance species, which appeared in
the Nonindigenous Aquatic Nuisance Prevention and Control Act of 1990 (Public Law
101-636) and continues in use. An aquatic nuisance species was defined as a nonindigenous
species that “threatens the diversity or abundance of native species or the ecological stability
of infested waters, or commercial, agricultural or recreational activities dependent upon
such waters.” It is the same thing as an invasive species.
Some scientists dislike having the definition of invasive species dependent on negative
impacts and prefer a more neutral definition that focuses on the population growth and
range expansion of certain nonindigenous species. They would argue that impacts can be
somewhat subjective and vague, and that range expansion or spread is the main process that
distinguishes some nonnative species from others. For example, many a nonindigenous
species arrives, reproduces, and establishes a population that is self-sustaining over several
generations and perhaps for a very long time in natural and seminatural habitats at or near
the point of entry; but it never spreads beyond the immediate areas of its introduction—
i.e., it never become “invasive.” Such alien plants and animals may be categorized as estab-
lished. Where they have become regular, functioning parts of a recipient ecosystem, even
if far from the point of entry, they are considered naturalized.
Some invasive plants are identified as “noxious” species, a category defined in the 1974
Federal Noxious Weed Act as “any living stage, such as seeds and reproductive parts, of
any parasitic or other kind of plant, which is of foreign origin, is new to or not widely preva-
lent in the United States, and can directly or indirectly injure crops, other useful plants, live-
stock, or poultry or other interests of agriculture, including irrigation, or navigation, or the
fish or wildlife resources of the United States or the public health.” Noxious plants are so
designated by federal and state laws. For animals, the term injurious is used to describe
those “species, including offspring and eggs, determined to be injurious to the health and
welfare of humans, the interests of agriculture, horticulture or forestry, and the welfare
and survival of wildlife resources of the U.S.” (Branch of Invasive Species, U.S. Fish and
Wildlife Service). Under the Lacey Act (see Appendix B), the designation of such species is
made by the U.S. Department of the Interior; and the importation and interstate transport
of wild mammals, wild birds, fish, mollusks, crustaceans, amphibians, and reptiles so desig-
nated are regulated by the secretary of the interior. (Currently, no amphibians are on the list
of injurious species, but in September 2009, the Defenders of Wildlife petitioned Secretary
Salazar to place all amphibians on the list unless certified free of the chytrid frog fungus
[Batrachochytrium dendrobatidis].)

The Size of the Problem

According to many resource managers and ecologists, invasive species and potential
invaders together constitute one of the major environmental threats facing the United
States in the twenty-first century. Preventing or controlling the establishment of nonindige-
nous species is complicated by a number of uncertainties, including how many and which of
the nonnative species will become abundant and widespread and inflict significant impacts
on natural ecosystems; national, regional, and/or state economies; or public health.
According to one estimate, some 50,000 species of plant, animal, fungi, and microbe have
xvi n GENERAL INTRODUCTION

been transported to the United States. Of these, perhaps 6,500 have established viable, self-
sustaining populations and 500 or so have become invasive in the sense both of demonstrat-
ing rapid population growth and range expansion and of causing harm or altering natural
ecosystems. Actual numbers are difficult to obtain and are more reliable for some classes
of organisms than others. Attention focused earlier and with greater federal funding on agri-
cultural pests and nonnative aquatic species than on others, so more data are available for
those groups. Inconsistent use of terms often further muddies the waters of how many spe-
cies are invasive as opposed to simply introduced or naturalized. The figures that follow
should only be considered indicators of the size of the problem and not definitive, accurate
measures.

Invasive Plants
Plants in the United States consist of approximately 18,000 native species and 5,000 non-
indigenous species. As is true for most if not all types of invasive species, the greatest num-
bers occur in Florida and Hawai’i. Florida is home to more than 900 naturalized plant
species; Hawai’i reports 946. However, only some are invasive. Most plants that have
become invasive were intentionally brought to the United States as ornamentals or for uses
as varied as erosion control or herbal medicine. Others were accidental introductions, pri-
marily as contaminants in crop seeds or hay or carried in ship ballast. Many nonnative plants
alter natural ecosystems, displacing native species, many of which are rare or endangered,
but impacts are far-reaching and varied. Nonnative plants are estimated to comprise
73 percent of the weeds of cultivated farmland and 45 percent of pasture weeds. Some
plants may be easily contained in drier western climates where limited water prevents their
spread, but may be unstoppable in the rainier eastern states. Others may be invasive in
grasslands or deserts, but not in ecosystems where competition from other plants keeps
them under control.
Overall, 138 trees and shrubs are considered invasive in the United States. Some of the
most notorious trees are salt cedar or tamarisk (Tamarix spp.), Australian melaleuca
(Melaleuca quinquenervia), and Brazilian peppertree (Schinus terebinthifolius). Tamarisk
depletes surface and groundwater supplies and makes soils so salty that other plants are
unable to grow. Melaleuca infestations are not only threatening the Everglades’ unique eco-
system, but the tree is also detrimental to human health and causes increased incidence of
fire. Brazilian peppertree is ranked with melaleuca as one of the worst invaders of the
Everglades. In Hawai’i, unless checked, velvet tree (Miconia calvescens) could devastate native
forests and alter hydrology as it has in Tahiti.
Shrubs can be equally devastating. Tropical soda apple (Solanum viarum), from South
America and now invasive in the southeastern United States, is a competitive plant in agri-
cultural fields, both pasture and cropland. It also harbors insects or pathogens that severely
damage and reduce yields of food crops. Gorse (Ulex europaeus), a large bushy shrub with
many attractive yellow flowers, forms thorny thickets that impede passage and degrade the
quality of recreational activities. The oily foliage and deep accumulation of dry matter cause
fires to occur more frequently and to burn hotter. The common garden plant lantana
(Lantana camara) is also a fire hazard in southeastern states, where it invades citrus groves.
Its leaves and unripe fruit are toxic to animals and can cause death.
More than 500 introduced forbs and graminoids are considered noxious weeds. Forbs
may be as small as the 12 in. (30 cm) tall fig buttercup (Ficaria verna) or grow as large
as giant hogweed (Heracleum mantegazzianum), which can be 25 ft. (7.6 m) tall with leaves
 GENERAL INTRODUCTION n xvii

The Problem with Common Names and Why Scientific

Names Are Surer Bets

T he common names of plants and animals are not always reliable indicators of
what species is being discussed. Different names may be used by people in dif-
ferent parts of the country. For example, in the eastern United States the leafy
weed known in most places as Japanese knotweed is called Japanese bamboo,
although it is not even a relative of true bamboos, which are grasses. The misno-
mer points to another difficulty: common names frequently do not reflect the true
identify of a plant or animal. Among the many tiny invasive animals causing prob-
lems in the Great Lakes is the spiny water flea, not an insect like real fleas but a
crustacean.
Scientific names may at first be strange tongue-twisters because they have been
latinized, but they have been approved by international committees and linked to
one species and one species only. Often the name provides insights into the nature
of the organism or, the place where it was first discovered. Scientific names are
revised from time to time as scientists come to new understandings about a spe-
cies’ evolution and relation to other species, but while in current use they leave
no doubt what plant, animal, fungus, or microorganism is being discussed. Each
scientific name has two parts. The first word refers to the genus into which biolo-
gists have classified the species and the second identifies the particular member
of the genus. Both words must be used to correctly designate a particular species.
In the Encyclopedia’s species accounts, common names appear as the title of
each entry, and entries are arranged within major categories alphabetically by
common name. More frequently used alternate names, especially those of regional
prominence, are given as appropriate. The scientific name—and some others that
have been used in older literature (synonyms)—follow. A reader seeking more
information should search according to the scientific name, or at least verify that
the scientific name is the same, to be sure he or she is gathering data about the
same organism as the one described in a particular entry.

as wide as 5 ft. (1.5 m). Fig buttercup displaces plants that are important sources of pollen
and nectar for pollinating insects, threatening their existence. Contact with the sap of giant
hogweed, a plant recently invasive in New England and the upper Midwest, causes a per-
son’s skin to be susceptible to severe sunburns. Several thistles and thistle-like knapweeds
degrade range and pasture land, and some, especially yellow star-thistle (Centaurea
solstitialis), are very poisonous to livestock. Prickly Russian thistle (Salsola tragus), more
commonly recognized as tumbleweed in the arid western states, looks more like a shrub
than a forb. As an annual, however, it lives only one year, leaving behind the woody skeleton
of its branches that rolls away in the wind, distributing seeds.
Many alien grasses alter the fire regime of rangeland or desert scrub, destroying native
plant species that are not fire adapted, and in turn displacing the animals dependent on that
ecosystem. These include the infamous cheatgrass (Bromus tectorum) that plagues western
grazing lands, lowering the nutrient value of rangeland and injuring livestock with their
sharp awns. The feathery plumes of fountain grasses (Pennisetum spp.) and Pampas grass
(Cortaderia selloana) that make them desirable as ornamental plants in the garden often make
it difficult to convince horticulturalists of the harm those plants can do when they escape
xviii n GENERAL INTRODUCTION

into the natural landscape, where they displace native plants and create fire hazards. Dense
stands of giant reed (Arundo donax), with vast intertwined root systems, clog stream
channels, altering hydrology and water flow.
Several vines grow so efficiently that they completely cover the ground and overtake
shrubs and even tall trees. Probably the best known is kudzu (Pueraria montana), which
blankets acres of land in southeastern states, not only smothering vegetation but breaking
trees and utility lines with the weight of the vines and foliage. Alien swallow-worts
(Cynanchum spp.) endanger monarch butterflies because eggs laid on the plants, in the
absence of the butterfly’s preferred host, do not survive. Japanese dodder (Cuscuta japonica),
a parasite just beginning to gain a foothold in California, poses such a significant threat, not
only to natural landscapes but also to crops, that it could affect international trade of crop
seeds.
Among the 160 or so aquatic plants that are nonindigenous, the best known and most
troublesome invasives are probably waterhyacinth (Eichhornia crassipes) and hydrilla
(Hydrilla verticillata). Both species grow into dense mats that alter natural ecosystems, down-
grade water quality, impede water transportation, and limit recreational activities.

Invasive Mammals
Introduced mammals include all domesticated livestock; some of their feral descendents
that run free in natural habitats are among the world’s worst invaders. In the continental
United States, feral pigs (Sus scrofa) are a particular scourge; on Hawai’i and islands in gen-
eral, feral goats (Capra hircus) are other voracious destroyers of native plants and the habitats
supporting endemic birds and other animals. Other mammals that were never domesticated
but nonetheless transported to the United States by humans, either deliberately or acciden-
tally, include such invasive species as the black or roof rat (Rattus rattus), Norway rat (Rattus
norvegicus), nutria (Myocastor coypus), and the Indian mongoose (Herpestes javanicus).
Introduced birds are some of the better-studied species as well as the animals perhaps
most familiar to the American public. The urban Rock Pigeon (Columba livia) is actually a
feral bird first brought to the country in colonial times as a food animal. Two other common
avian pests in cities and towns today were deliberately released to “enhance” the wild bird
fauna of the United States: the House Sparrow (Passer domesticus) and the European
Starling (Sturnus vulgaris). The most abundant bird in Hawai’i, the Japanese White-eye
(Zosterops japonicus), came from eastern Asia.
Reptiles introduced from other continents include the huge Burmese python (Python
molurus bivittatus)—now rapidly increasing its numbers in the Everglades and spreading
beyond the park boundaries to invade other parts of Florida—as well as the brown anole
(Norops sagrei), a small lizard that appears to be spreading out of Florida. Among invasive
amphibians is the annoying little Cuban treefrog (Osteopilus septentrionalis), which gets into
peoples’ houses, but the most ecologically damaging is a native frog from the eastern
United States introduced into western waters, the American bullfrog (Lithobates catesbeia-
nus). As further indication of how aquatic ecosystems have been affected by so-called native
transplants like the bullfrog, almost two-thirds of the fishes found in American drainages
were introduced by people from other U.S. waterways, sometimes in the same state and
sometimes from across the continent. Examples of such native invasive fishes are the alewife
(Alosa pseudoharangus) in the Great Lakes, the western mosquitofish (Gambusia affinis) in
states west of its native range limits in Texas, and rainbow trout (Oncorhynchus mykiss),
native to Pacific drainages but now stocked throughout the United States and indeed much
 GENERAL INTRODUCTION n xix

of the world. Alien invasive fish include various Asian carps and such strange creatures as
walking catfish (Clarius batrachus) and the northern snakehead (Channa argus).
Invertebrates, fungi, and microorganisms pose an even greater challenge when estimating
numbers of invasive species. We do not even know how many native species exist in most
taxonomic groups. According to one estimate, some 4,500 arthropods have been introduced
to the United States, more than 2,500 of them to Hawai’i. The hemlock woolly adelgid
(Adelges tsugae) has decimated hemlocks in natural forests and opened forest streams to sun-
light, while the Asian longhorned beetle (Anoplophora glabripennis) threatens shade and
ornamental trees in New York and Chicago but could become a forest pest on a par with
the invasive gypsy moth (Lymantria dispar) if it invades eastern broadleaf deciduous forests.
Some invading insects, such as the Formosan subterranean termite (Coptoptermes formosa-
nus), cause devastating structural damage to buildings; while others, such as the common
bed bug (Cimex lecturius) and brown marmorated stink bug (Halyomorpha halys), are simply
serious annoyances inside man-made structures. Among the more potentially damaging
insect invaders are those that could disrupt crop pollination by negatively affecting
European honey bees—themselves an introduced (domesticated) species. Africanized honey
bees (Apis mellifera scutella) hybridize with European honey bees and take over their hives.
Two introduced arachnids, the honeybee tracheal mite (Acarapis woodi) and the varroa mite
(Varroa destructor) parasitize bees and lead to the demise of bee colonies.
Snails, clams, and mussels are among other significant invertebrate invaders. Zebra mus-
sels (Dreissena polymorpha) can transform freshwater communities and clog water intake and
distribution pipes. Their rapid spread through Great Lakes and into the Mississippi River
drainage in the 1980s was a major stimulus to the development of interest in exotic species
in general in the United States.
Invasive fungi are largely associated with diseases affecting trees, such as Dutch elm dis-
ease and sudden oak death, but a fungus is also responsible for a disease infecting frogs
and toads in many parts of the United States. One of the newest invaders, the fungus
Geomyces destructans, is implicated in bat white-nose syndrome, a condition currently ravag-
ing bat colonies in the eastern United States.
The smallest of invasive species, the microorganisms, are represented by only three
entries in this encyclopedia. Humans have been transporting protists, bacteria, and viruses
as long as human migrations have taken place. The human diseases brought by early settlers
from overseas—smallpox, influenza, measles, to name a few—decimated Native American
populations. More recently, HIV and new strains of influenza have run rampant through the
U.S. population. Emerging infectious diseases such as dengue fever or ebola may be just
around the corner. These are more appropriately dealt with in a book on epidemiology than
one on invasive species. The three organisms chosen for inclusion here have close ties to
natural habitats and infect wild animals (as well as, in some cases, humans). Avian malaria
threatens rare, endemic birds in Hawai’i. The bacterium Borrelia burgdorferi has a compli-
cated life cycle involving two mammalian hosts and expresses itself as Lyme disease in
humans. The West Nile virus infects birds, horses, and humans. It is now controlled in
horses by vaccination; its long-term impact on wild bird populations remains to be seen.
To put the above information in some perspective, it should be noted that about 200,000
species are believed to inhabit the United States. About 91,000 have been described, leaving
a large number of plants, animals, fungi, and microbes yet to be discovered. As of the latest
count, plants account for nearly 19,000 of the native species, and vertebrates for about
3,000. (According to one count, there are 1,154 native fishes, 295 amphibians, 311, reptiles,
784 birds, and 428 mammals.)
 xx n GENERAL INTRODUCTION

Native aquatic species—mussels, fishes, salamanders, and turtles—are quite diverse by

global standards, as are habitats for both terrestrial and aquatic life. Nearly one-third of
our species are considered at risk, including almost 70 percent of freshwater mussels and
more than 50 percent of native crayfishes. Natural vegetation has been removed or greatly
altered in more than 60 percent of the land area in the lower 48 states. Experts generally
agree that destruction of habitat is the greatest threat to our native species, but that the intro-
duction of nonnative species is the second greatest cause of decline and disappearance of
native plants and animals.

The Invasion Process

For a nonnative species to gain a foothold and become an abundant and widespread
inhabitant in an area beyond its native range, several steps or stages are required. Each step
involves overcoming some sort of barrier. Species are normally held in their native ranges by
geographic barriers to dispersal, such as an ocean, a different and inhospitable (for them) cli-
mate region, a drainage divide, a mountain range, distance, wind or ocean current direction,
or some other natural feature of the planet. The first stage, the transport stage, requires get-
ting through or over the unfavorable conditions imposed by what is normally a geographic
barrier and entering a new site. Plants and animals have done this successfully on their
own over millions of years, either by chance or because changes to the environment weaken
or remove the barriers. Thus, species have colonized oceanic islands and moved from one
continent to another. When humans began to migrate and then to engage in trade, they
accelerated the process by either deliberately or unintentionally providing plants, animals,
and microorganisms safe transport to new areas in their provisions, packing materials, ships,
and other vehicles. Some organisms became desirable commodities in their own right as
exemplified today by the pet trade and horticultural industry.
Getting to a new site is only the first step. Once a species has arrived or “been intro-
duced,” it must be able to reproduce and establish a self-sustaining population before it
can be in a position to become invasive. Surmounting limitations imposed by small founding
population sizes and environmental barriers in the new location are the challenges of
the second or establishment stage. Typically, only a small number of individuals of a given
species occur in the new area, a factor that by itself makes them vulnerable to extinction.
The so-called Allee effect
depresses reproductive rates
since the few individuals present
may be sparsely distributed and
have difficulty finding mates, or
the sex ratio may be skewed
toward one gender or the other,
both limiting the number of off-
spring that can be produced
during the first few generations.
In many instances, a new spe-
cies persists in an area only
because it is repeatedly intro-
The three stages of the invasion process, showing population status and duced and not because the spe-
dispersal patterns in each. cies is reproducing at the site.
 GENERAL INTRODUCTION n xxi

Since the new environment will not be identical to that of the native range, the species
may need time to adapt to a new climate, substrate, ecological community, or other aspect
of the receiving habitat before its success is assured. If a population does not evolve, it
remains highly localized or goes extinct. Even when the demographic and environmental
barriers are overcome, it can take 10–25 years for a new species to become numerous
enough for people to detect its presence. Success in the establishment phase means repro-
duction occurs regularly and not only sustains the population, but allows for population
growth. The species nonetheless remains near the initial point of introduction at this stage.
When the number of individuals increases, the tendency is for a species to expand its dis-
tribution area—that is, to spread. Eventually, it will have to overcome local geographic bar-
riers to dispersal. As it spreads into new regions in its adopted home, it enters the invasive
stage and begins to produce viable offspring at some distance from the original place of
introduction. During the invasive stage, a species continues to increase in abundance and
expand its range. Typically, it will first occupy sites disturbed by human activities (such as
farmland and settlements) and then be found in seminatural and natural ecosystems. If the
new species is perceived to harm or change native ecosystems, it may be deemed officially
an “invasive species.” Some ecologists prefer that infliction of ecological and economic harm
on the recipient region be considered an additional, fourth stage in the invasion process.
Species can spread without causing harm, as has happened with the Cattle Egret.
It is frequently stated that, on average, about 10 percent of species actually move from
one stage to the next. This means that one in ten of those transported to a new area actually
survives and will be found outside of captivity or other controls (i.e., is introduced). One in
ten of those will establish self-replacing populations (i.e., become established), and one in
ten of established species will spread and become invasive. The result is that very few species
arriving in areas they previously have not occupied pass through all the barriers and become
invasive. Most species fail to gain even a temporary foothold, let alone invade new regions.
Studies have shown that this so-called “tens rule” describes the history of nonindigenous ter-
restrial vertebrates, fishes, insects, mollusks, and plant pathogens in the continental United
States fairly well, but fails to accurately describe nonnative birds in Hawai’i, where more
than half of all birds known to have been introduced have become established. It should
be noted that the original work that proposed the tens rule focused on plants introduced
into the United Kingdom; 10 percent was the average of a range running from 5 to 20 percent.
Research examining only those vertebrates coming into the United States from Europe
determined that 25 percent became invasive. At this point, it remains unknown how
many actually can be expected to succeed, or what makes a species a successful invader,
or what allows a native ecosystem to resist, accommodate, or succumb to invasive alien
species.
For species that do become “invasive,” the process proceeds in a typical pattern. Initially,
only a few individuals are spotted in a new area. There is a long lag time before, suddenly,
what seemed like an innocuous new member of an ecological community explodes in num-
bers (irrupts) and becomes a major nuisance or pest and may even begin to transform native
ecosystems. Very often, at least in non-island situations, after the initial irruption, the non-
indigenous population declines; and the native system accommodates the newcomer.
When this happens, the exotic becomes a naturalized member of the community.
Recognition of stages in the invasion process helps resource managers design control and
mitigation programs. The easiest and most cost-effective control scheme is to intercept arriv-
als and prevent establishment and/or spread in the first place. This involves detecting
incoming species through inspection and quarantines. Constant monitoring is necessary to
xxii n GENERAL INTRODUCTION

measure the success or failure of such measures. Once a population is growing and spread-
ing, management becomes problematic and eradication nearly impossible.

Pathways of Introduction
The means by which species are introduced into new areas are known as pathways. These
are generally separated into deliberate or intentional pathways and accidental or uninten-
tional pathways (Table 1 and Table 2). Escapes from confinement or captivity may
be thought of as a third, hybrid pathway in which the initial importation was intentional,
but the release of free-living individuals or populations was an accident. Generally speaking,
most plants and vertebrates have been introduced deliberately; most invertebrates and
microorganisms arrived by accident.
Plants naturally disperse by sending forth spores and seeds and, for a few, seedlings or
parts capable of sprouting vegetatively. These propagules are transported by wind, ocean
and stream currents, or animals to places beyond the parent plant and sometimes beyond
the range limits of the species. If they land in new territory, they may or may not survive
and establish a colony. Humans may have first changed the dispersal process for some plants
(and animals) by creating disturbed habitats around campsites and unintentionally carrying
seeds on their bodies or in their digestive tracts—just as other animals do. They also har-
vested and stored seeds and deliberately transported them as they moved from site to site.
Some plants adapted to the new dispersing agent and became “camp-followers,” showing
up unbidden at each new settlement. Cannabis sativa (marijuana) was such a plant in the
Old World. The precursors of many crops may have acted similarly. Mammals such as the
dog and pig may have also joined the retinue of people on the move along with arthropods
such as cockroaches, lice, silverfish, and other such species that came to live in close associ-
ation with human beings.
Once domesticated crops and livestock were available, people deliberately transported
them to far-off places. Seeds and cuttings were traded first among neighboring villages and
later to distant shores. With rafts and sailing ships, people helped terrestrial and freshwater
species overcome the geographic barrier of the sea. Plants and animals were carried inten-
tionally and accidentally around the world, especially by Europeans and Polynesians. As sea-
going technology improved, more distant parts of the planet became connected. Sailing
ships were pretty much confined to the natural routes determined by wind and current.
But sea-lanes were cut across these old routes when steamships came of age. Seaports were
the first point of entry for many species new to an area. Accidental travelers were hidden
in the dry or solid ballast of ships under sail. Others, such as shipworms, burrowed in the
wooden hulls. During the colonial period, the flow of goods ran largely from the Americas
to Europe. Ships from England, France, Spain, and Holland sailed without full loads and
took on needed ballast at their home ports. The ballast was offloaded at the port of destina-
tion, and if nonnative species had survived the journey, they might become established near
wharves and piers and at ballast dumping grounds. Botanists looking for “new” species
hunted in these locations. Seeds and insects also stowed away in the hay and straw carried
aboard ships as packing material or as fodder and bedding for livestock. Rats climbed the
mooring lines at one port to sail to and disembark at another. Everything was on the move.
From seaports, a few new arrivals were able to move inland along the canals built to tie
the port city to its hinterland. Purple loosestrife (Lythrum salicaria), for example, a plant that
came to dominate marshes in New England, was first collected from wetlands near the Erie
and Delaware-Ruritan canals. Railroads penetrated even farther inland; their right-of-ways

Table 1. Examples of Invasive Species That Have Been Introduced
by Intentional Pathwaysa
Aesthetic amenities, sentiment,
or nostalgia
Aquarium trade
Bait bucket releases
Biological controls
Bioterrorism (potential)
Botanical gardens
Domestic use (dyes, fish poisons)
Erosion control/bank stabilization
Food and beverage
Forage fish
Furbearers
Livestock abandonment
Livestock forage or fodder
Medicinal herbs or seasonings
GENERAL INTRODUCTION n xxiii
European Starling (Sturnus vulgaris)
House Sparrow (Passer domesticus)
Ornamentals, including but not limited toLantana
(Lantana camara)
Multiflora rose (Rosa multiflora)
Strawberry guava (Psidium cattleianum)
Wisterias (Wisteria sinensis, W. floribunda)
Common salvinia (Salvinia minima)
Hydrilla (Hydrilla verticillata)
Lionfish (Pterois volitans)
Nightcrawler (Lumbricus terrestris)
Rusty crayfish (Orconectes rusticus)
Common myna (Acridotheres tristis)
Grass carp (Ctenopharyngodon idella)
Mosquitofish (Gambusia affinis, G. holbrooki)
Multicolored Asian lady beetle (Harmonia axyridis)
Indian mongoose (Herpestes javanicus)
Infectious diseases
Bush honeysuckles (Lonicera spp.)
Japanese barberry (Berberis thunbergii)
Velvet tree (Miconia calvescens)
Common mullien (Verbascum thapsus)
Dyer’s woad (Isatis tinctoria)
Yellow toadflax (Linaria vulgaris)
Australian pine (Casuarina equisetifolia)
Giant reed (Arundo donax)
Ice plant (Carpobrotus edulis)
Japanese knotweed (Fallopia japonica)
Kudzu (Pueraria montana)
Asian clam (Corbicula fluminea)
Chinese mitten crab (Eriochor sinensis)b
Chinese mystery snail (Cipangopaludina chinensis
malleata)
Fire tree (Morella faya)
Golden apple snail (Pomacea canaliculata)
Himalayan blackberry (Rubus armeniacus)
Northern snakehead (Channa argus)
Alewife (Alosa pseudoharengus)
Nutria (Myocastor coypus)
Feral goat (Capra hircus)
Feral horse (Equus caballus)
Feral pig (Sus scrofa)
Buffelgrass (Pennisteum ciliare)
Cogon grass (Imperata cylindrica)
Gorse (Ulex europaeus)
Common St. Johnswort (Hypericum perforatum)
Garlic mustard (Alliara petiolata)
Giant hogweed (Heracleum mantegazzianum)
Japanese hops (Humulus japonicus)
(Continued )
xxiv n GENERAL INTRODUCTION

Table 1. (Continued)
Packing material Japanese stilt grass (Microstegium vimineum)
Princess tree (Paulownia tomentosa)
Smooth cordgrass (Spartina alterniflora)
Pet abandonment Burmese python (Python molurus bivittatus)
Feral cat (Felis silvestris catus)
Nile monitor (Varanus niloticus)
Pet trade Burmese python (Python molurus bivittatus)
Giant African snail (Achatina fulica)
Nile monitor (Varanus niloticus)
Research African clawed frog (Xenopus laevis)
Gypsy moth (Lymantria dispar)
Sport fishing Brown trout (Salmo trutta)
Rainbow trout (Oncorhynchus mykiss)
Timber/reforestation/firewood Carrotwood (Cupaniopsis anacardioides)
Melaleuca (Melaleuca quinquenervia)
Wildlife habitat or food Chinese lespedeza (Lespedeza cuneata)
Russian olive (Elaeagnus angustifolia)
Windbreaks/fencerows Melaleuca (Melaleuca quinquenervia)
Russian olive (Elaeagnus angustifolia)
Tamarisk (Tamarisk chinensis, T. ramosissima)
a
See table “Pathways of Introduction for Plants,” in Volume 2, for a more complete listing of plant pathways.
b
Probable means of introduction.

became avenues of expansion for weedy plants, while other organisms rode the rails as
hitchhikers in cargo and packing materials to almost all parts of the country. Airplanes
now reach the most isolated places, so nowhere is immune to the introduction of nonnative
species.
Sometimes military traffic is implicated in the transport of unwanted species to American
shores. Perhaps the most notorious example is the arrival of the brown tree snake (Boiga
irregularis) on the U.S. territory of Guam after World War II. But a noxious weed, Canada
thistle (Cirsium arvense), had also moved on Union military steamships during the
American Civil War, reaching its southern limits at the Virginia town of Remington. More
recently, the Argentine ant (Linepithema humile) arrived in Hawai’i on military ships during
World War II, and the Formosan subterranean termite (Coptotermes formosanus) reached
Houston, Texas, on ships returning from the Pacific theater.
Shipping remains an important pathway for the entry of new species. Fouling organisms
such as lacy crust bryozoan (Membraniphora membranacea), chain tunicates (Botrylloides vio-
laceus), and colonial tunicates (Didemnum vexillum) made their way to U.S. waters in this
manner. Today, seawater is used as ballast, and both larval and adult aquatic organisms,
plants as well as animals, have been transported from one port across an ocean to another
port, where the ballast water is discharged. The zebra mussel (Dreissena polymorpha) and
the spiny water flea (Bythotrephes longimanus) both presumably made their way to the
Great Lakes in this manner. Canal construction opened the Great Lakes to invasion by such
fish as sea lamprey (Petromyzon marinus) and round goby (Neogobius melanostomus). The
shipment of used tires on cargo ships provided shelter and breeding sites for Asian tiger
mosquitoes (Aedes albopictus), allowing their entry into the United States.

Table 2. Examples of Invasive Species That Have Been Introduced
by Unintentional Pathwaysa
Ballast and bilge water discharge
Crop seed/plant contaminants
Dry (solid) ballast
Escapes from aquaculture
Escapes from fur farms
Escapes from gardens
Escapes from pet owners
Escapes from research labs
Fish stock contaminants
Hull fouling
Ocean currents
Removal of natural barriers once
separating bodies of water
(e.g., canal construction)
Stowaways in cargo
Stowaways in clothing, bedding, and Common bed bug (Cimex lectularis)
luggage
Stowaways in equipment, including Canada thistle (Cirsium arvense)a
military
Stowaways in grain shipments
Stowaways in packing materials
GENERAL INTRODUCTION n xxv
Asian green mussel (Perna viridis)
Common reed (Phragmites australis spp. australis)
Eurasian watermilfoil (Myriophyllum spicatum)
Purple loosestrife (Lythrum salicara)
Quagga mussel (Dreissena rostriformes bugensis)b
Smooth cordgrass (Spartina alterniflora)
Spiny water flea (Bythotrephes longimanus)
Zebra mussel (Dreissena polymorpha)
Canadian thistle (Cirsium arvense)
Cheatgrass (Bromus tectorum)
Prickly Russian thistle (Salsola tragus)
Quackgrass (Elymus repens)
Yellow starthistle (Centaurea solsititialis)
Earthworms
Common periwinkle (Littorina littorea)
American bullfrog (Lithobates catesebianus)
Bighead carp (Hypophthalmichthys nobilis)
Giant salvinia (Salvinia molesta)
Grass carp (Ctenopharyngodon idella)
Water chestnut (Trapa natans)
Waterhyacinth (Eichhornia crassipes)
Nutria (Myocastor coypus)
Chocolate vine (Akebia quinata) and most others that
originated as ornamental plants
Feral cat (Felis silvestris catus)
Monk Parakeet (Myopsitta monachus)
Mute Swan (Cygnus olor)
Africanized honey bee (Apis mellifera scutellata)
Gypsy moth (Lymanatria dispar)
African clawed frog (Xenopus laevis)
New Zealand mud snail (Potamopygrus antipodarum)b
Chain tunicate (Botrylloides violaceus)
Colonial tunicate (Didemnum vexillum)
Lacy crust bryozoan (Membranipora membranacea)
Asiatic colubrina (Colubrina asiatica)
West Indian marsh grass (Hymenachne amplexicaullis)
Round goby (Neogobius melanostomus)
Sea lamprey (Petromyzon marinus)
Zebra mussel (Dreissena polymorpha)
Asian tiger mosquito (Aedes albopictus)
Formosan subterranean termite (Coptotermes
formosanus)
Argentine ant (Linepithema humile)
Argentine ant (Linepithema humile)
Asian longhorned beetle (Anoplophora glabripennis)
Brown marmorated stink bug (Halyomorpha halys)
(Continued )
xxvi n GENERAL INTRODUCTION

Table 2. (Continued)
Stowaways in wood products, Dutch elm disease fungus (Ophiostoma novo-ulmi)
including firewood Emerald ash-borer (Agrilus planipennis)
Stowaways on live plants, bulbs, or Glassy-winged sharpshooter (Homalodisca vitripennis)
root balls Hemlock wooly adelgid (Adelges tsugae)
Coqui (Eleutherodactylus coqui)
Chestnut blight fungus (Cryphonectria parasitica)
Earthworms
Japanese beetle (Popillia japonica)
Red imported fire ant (Solenopsis invicta)
Stowaways on ships Black rat (Rattus rattus)
Cuban treefrog (Osteopilus batrachus)
Norway rat (Rattus norvegicus)
a
See table “Pathways of Introduction for Plants,” in Volume 2, for a more complete listing of plant pathways.
b
Probable means of introduction.

Most fungal pathogens arrive with their natural host plants. In the case of the chestnut
blight fungus (Cryphonectria parasitica), live plant material was involved. Dutch elm disease
arrived on veneer logs. Fungi may also be inadvertently transported in soil and in root balls.
Insects, too, infest plants, including wood products, seed supplies, and grain shipments.
Terrestrial vertebrates are mostly imported and transported deliberately. Amphibians and
reptiles are often valued as bait and pets and used for biomedical research. They may also
hitchhike on plants; the coqui (Eleutherodactylus coqui), which has invaded Hawai’i, is a case
in point. Nonnative birds have been released as game animals, but most come in as pets and
either are released or escape captivity. Many of the more troublesome invasive mammals were
deliberately imported as livestock and then deliberately released to the wild (e.g., goats and
pigs). Some, such as the Indian mongoose (Herpestes javanicus), were deliberately released as
potential biological controls for agricultural pests. Terrestrial invertebrates have been intro-
duced as aquarium novelties, food, pets or ornamentals, and biological control agents.
Freshwater aquatic vertebrates (primarily fish) are often deliberately stocked in ponds
and streams for sport fishing or as future food sources. Some (e.g., alewives) were released
to serve as forage fish for larger game fishes. Invasive aquatic amphibians and reptiles tend
to result from pet releases, bait bucket releases, or biological control efforts. Contamin-
ation of fish stock by the African clawed frog (Xenopus laevis) led to the establishment of that
amphibian in parts of California, but American bullfrogs in western waters stem from
releases for food and escapes from aquaculture facilities.

Impacts
Invasive species are associated with a variety of ecological, economic, public health, and
aesthetic impacts. Sometimes, whether these are positive or negative lies in the eye of the
beholder. Usually “impact” is assumed to be negative, and known or potentially negative
impacts are the reasons why invasive species are receiving so much attention in the
twenty-first century. Indeed, the official definition of “invasive” in the United States includes
reference to the harm a species can or does do. Yet it should be remembered that many
 GENERAL INTRODUCTION n xxvii

Table 3. Potential Ecological Impacts of Invasive Speciesa

Genetic Change in genetic information in a native species
through introgression
Hybridizing with native species to produce offspring
(or new species) that outcompete parents
Individual Changes in foraging, pollination, or reproductive behavior
Population Reductions in population size; niche shifts; local
extinction
Community Changes in species composition and interactions
Ecosystem Changes in nutrient cycles and disturbance regimes
a
See table, “Impacts of Invasive Plants,” in Volume 2, for the complete listing of impacts of invasive plant
species.

species are initially introduced because someone foresees a benefit, be it a beautiful blossom,
a challenging game animal, a fascinating pet, or a way to control a pest or ameliorate an envi-
ronmental problem.

Ecological Impacts in Natural and Semi-natural Ecosystems

As new species spread into the wild and semi-wild habitats of the United States, they have
the potential to affect life in our forests, grasslands, and deserts at all biological/ecological
levels (Table 3). Native organisms may respond as individuals to a new predator or competi-
tor by altering their behavior. A case in point is the avoidance of some rodents to areas
infested by the red imported fire ant. As a result, the mice may forage in less protected areas
and become more vulnerable to predation by owls.
Individual organisms can also be affected by introduced pathogens, predators, or compet-
itors for limited resources to the extent that the impact becomes evident at the population
level, when increased mortality rates threaten the survival of the entire population of a given
organism. Avian malaria threatens to decimate several endemic honeycreepers in Hawai’i.
West Nile virus caused significant population declines in songbirds such as crows and chick-
adees when it first spread through eastern states. In the Great Lakes, the round goby
(Neogobius melanostomus) displaces native fish such as the mottled sculpin (Cottus bairdi) from
its customary spawning grounds and competes with it and other fish for food. Certainly the
most notorious recent invader is the predatory brown tree snake, which arrived in Guam
sometime between the end of World War II and 1952 and, in the next 20 years, caused the
extinction of 10 species of native forest birds and decimated lizards, causing the local extinc-
tion of 4 species. In addition, the brown tree snake is implicated in the loss of two species of
bat from Guam. While the example of the brown tree snake is unusually dramatic and illustra-
tive of what can happen when a new predator is introduced to an island previously lacking
predators, it does serve as a cautionary tale of how wrong things can go.
The zebra mussel (Dreissena polymorpha), through its rapid population growth and ability
to grow on the shells of native unionid mussels, can physically overwhelm the host and
reduce its access to nutrients. Among plants, spotted knapweed (Centaurea stoebe), a peren-
nial forb, is an introduced competitor species. It produces an allelopathic chemical that
depresses the germination or growth of native plants, such as the endemic Mt. Sapphire
rockcress (Arabis fecunda), and thereby preserves a greater share of light, water, and
nutrients for the invader.
xxviii n GENERAL INTRODUCTION

Native populations may also be affected at the genetic level through hybridization and
introgression. Hybridization involves the cross-breeding of members of two species. When
viable offspring are produced, they may exhibit hybrid vigor and grow faster or larger than
either parent and reproduce more quickly than either. If sterile offspring are produced, the
parent species have wasted their gametes, a practice that may be costly if their numbers are
already low. The rainbow trout (Oncorhynchus mykiss), a native transplant to western waters,
produces fertile offspring when it mates with the California golden trout (O. mykiss aguabonita)
and the threatened Paiute cutthroat trout (O. clarki seleniris). The hybrid offspring of rainbow
trout and golden trout can backcross with both parent species and contaminate the gene pool
of golden trout by introducing genes of rainbow trout, a process known as introgression.
Through this means the native genotype can disappear. Smooth cordgrass (Spartina alterni-
flora), another native transplant to California, readily hybridizes with the native California
cordgrass (Spartina foliosa). The first-generation hybrids have higher growth rates and greater
reproductive success than either parent. The hybrids also tolerate a broader range of salinity
and invade open mudflats, changing the physical environments in an estuary.
Loss of one or more species, as well as the addition of new species, has repercussions for
an entire ecological community. Mutualistic relationships such as predator/prey and pollina-
tor/host can be disrupted. Niche shifts can occur as new species are accommodated. Where
the brown anole is present, the native green anole (Anolis carolinensis), which customarily
seeks prey on the ground or lower regions of tree trunks, forages higher in the tree canopy.
Chestnut blight fungus (Cryphonectria parasitica) essentially eliminated the American chest-
nut from the tree layer of eastern forests. The relative abundance of other trees in the forests
changed as white oak (Quercus alba), chestnut oak (Q. prinus), and red oak (Q. rubra)
increased in the absence of chestnut. The red imported fire ant has changed the composition
of ant communities in areas it has invaded, which in turn may have reduced the dispersal of
seeds and thereby affected the plant community as well. Competition for nutrients, water,
shelter, or breeding grounds most affects those natives already threatened with extirpation
from other forces. Among the more than 950 species listed as endangered or threatened in
the United States, some 400, or about 42 percent, are believed to be at risk in part due to
the impacts of invasive species. Interestingly, despite the demise of some species, at the local
level, species richness usually increases, as invaders more than make up for the loss of
natives.
When abiotic as well as biotic elements are affected, impacts are occurring at the ecosys-
tem level. Two important ecosystem-level changes involve nutrient cycles and disturbance
regimes. By filtering such huge amounts of water, dense populations of zebra mussel
increase the amount of nitrogen and phosphorus in the water column and reduce the
amount of carbon, which migrates down to the bottom-dwellers in the mussels’ pseudofe-
ces. On Hawai’i, a plant alters the nitrogen cycle, with ecosystem-wide consequences. The
fire tree (Morella faya) fixes nitrogen from the atmosphere, which allows it—unlike any
native trees—to live on the nitrogen-poor volcanic substrates of the islands. Fire trees add
nitrogen to the soil and provide suitable conditions for a variety of other plants to colonize
the area, thereby giving rise to a whole new community of plants.
On western rangelands, the dominance of cheatgrass has altered the disturbance regime
from one where fires occurred on average once every 6–10 years to a burn cycle of 3–5 years.
Such frequent fires eliminate shrubs, forbs, and native grasses and produce a monoculture of
cheatgrass, an annual grass with low forage value.
At a regional or landscape scale, the mixing of species from all over the globe leads to the
homogenization of the world’s biota and a loss of global biodiversity. While locally, the total
 GENERAL INTRODUCTION n xxix

number of species may increase, the same species tend to be added everywhere. At the same
time, geographically restricted and unique species tend to be disappearing. This impoverish-
ment of the variety of life on earth is viewed with alarm, for homogenization is occurring at
all levels—genetic, species, community, and ecosystem—potentially interfering with ecosys-
tem functioning, ecosystem services, and the ability to evolve and adapt to changing envi-
ronmental conditions. Furthermore, the world becomes a less interesting place as a great
sameness spreads across not only our human-made townscapes and cityscapes, but the
natural world as well.
The impacts of new invasive species are often not as much of a problem as initially feared.
A case in point is the Monk Parakeet (Myiopsitta monachus), which had the potential of
becoming a major agricultural pest based on the habits of its extinct relative, the Carolina
Parakeet (Conuropsis carolinensis). However, the Monk Parakeet has not spread beyond
urban areas, where many people are pleased to have these colorful birds visit their feeders.
Unfortunately, it is still impossible to predict which introduced species will become invasive
and which invasive species will become major ecological problems.

Economic Impacts of Invasive Species

Estimates of the costs of invasive species to the United States are commonly reported at
well over $100 billion a year, but it is impossible to know the exact figure, which is likely
much higher (Pimental et al. 2000; Pimental et al. 2004). It is difficult to put a dollar value
on ecological damages or to separate the financial impacts of the combination of factors
affecting agriculture, forestry, fisheries, industry, land values, and human well-being, not
to mention the price of control and measures directed at preventing introductions, manag-
ing invasive species, and implementing remediation measures to repair the damage.
In agriculture, direct damage to crops and pastures affect yields and the quality of the
product, which in turn can have repercussions on market value. Weeds, insects, and patho-
gens cost agriculture about $25 billion each year in lost production and another $3 billion
for pesticides to control them (Pimental et al. 2000). Measures set up to protect plant and
animal life and human health in the United States can become trade barriers and violate
World Trade Organization (WTO) agreements, causing costly boycotts among international
buyers. The inspections, quarantines, monitoring, and response to introduced insects and
pathogens are all expensive but necessary to protect our food supply as well as our farmers.
Forestry suffers financial loss not only when trees sicken and die, but also when quarantines
prevent sales of wood products. Some 9 percent of lumber, pulpwood, and other forest
products are thought to be lost to insect damage at an annual cost of $7 billion. The “Slow
the Spread Program” for gypsy moth costs the federal government $8–10 million a year, with
additional funds provided by affected states (Tobin 2008). Similar programs are needed for
the emerald ash borer (Agrilus planipennis) and other invasive insects. The green crab
(Carcinus maenas) is assumed to have caused the demise of the soft shell clam (Mya arenaria)
fishery in New England in the 1950s and is also implicated in declines of the commercially
important northern quahog (Mercenaria mercenaria), a scallop (Argopecten irradians), and
other shellfish with annual harvests worth $44 million in 2000 (Perry 2000). Fouling dam-
age by Asian clams (Corbicula fluminea) in the United States reportedly causes $1 billion a
year. The damage by zebra mussels and control of those mollusks at raw water-using and
electricity-generating industries in the Great Lakes region are estimated to cost more than
$100 million a year (Wisconsin Department of Natural Resources, 2004). Other Great
Lakes invaders affect the recreation industry in the region, where losses from sport fishing
xxx n GENERAL INTRODUCTION

alone reach an estimated $200 million a year. In Florida, hydrilla clogs waterways and costs
$14.5 million a year to control, but there are still financial impacts on the recreation and
tourism industries (Pimental et al. 2000; Pimental et al. 2004).

Public Health and Well-Being Impacts

The most obvious impacts to human health are new pathogens. Global epidemics are
expected to become more common with ever-increasing international travel and the globali-
zation of world trade. Such pathogens, past and future, are not covered in this encyclopedia,
but they have followed or will follow the same pathways as other invasive species. For some,
their virulence depends on finding reservoirs and transmitters among both native and non-
native animals, as in the case of the West Nile virus and the bacterium that causes Lyme dis-
ease. A warming climate and invasive mosquitos likely mean the establishment of (currently)
tropical diseases like dengue fever in the near future; but the ubiquity and rapidity of air
travel opens the U.S. population to all sorts of emerging infectious diseases.
A host of simply annoying species have invaded the United States. Asian multicolored
lady beetles and brown marmorated stink bugs are two recent examples, while the common
bed bug represents a very old traveler now experiencing a resurgence. For folks living in
parts of Hawai’i, the noisy coqui can be added to the list, and for people in Florida, the
Cuban treefrog lurking in the toilet fits the bill. Such annoyances can be expensive. The
din of calling coquis can lower property values. Bug-bitten customers sue hotels and land-
lords, adding to the costs incurred in trying to eradicate bed bugs.

Invasion Science: A Brief History

The seminal work in the scientific study of biological invasions is generally considered to be
Charles S. Elton’s 1958 book, The Ecology of Invasions by Animals and Plants, which called atten-
tion to some of the most damaging of invaders in various parts of the world. Among featured
invasive organisms were the chestnut blight fungus, which had ravaged forests in the eastern
United States, and the sea lamprey, which had rapidly spread through the Great Lakes after
the opening of the Welland Canal and decimated native fisheries. He described the impact
of the American smooth cordgrass on the tidal flats in his native England, where it had hybrid-
ized with another cordgrass (Spartina maritima) to create a new species now known as
common cordgrass (S. anglica). The hybrid was replacing both parent species and spreading
quickly in wetlands on both sides of the English Channel. (Today, smooth cordgrass is causing
similar problems in California.) He also noted how the European Starling had expanded its
range across the United States and Canada in little more than 60 years after the first pairs
had begun to breed in New York’s Central Park. Elton’s compendium obviously drew on ear-
lier work by others, including classic papers by M. T. Cooke (1928) on the spread of the star-
ling between 1891 and 1926 and by J. C. Phillips (1928) on the history of the spread of
nonnative birds across the continent. Probably the earliest records of European plants growing
in North America were published by the English traveler John Josselyn in 1672, when he cata-
logued plants observed in New England. Nonetheless, Elton’s work was especially influential,
and he became the “Father of Invasion Ecology.” He had sounded the alarm about the disrup-
tions to native ecosystems that can or could occur from successful invaders and called for
renewed efforts in the conservation of native biodiversity, in studying the effects of invasive
species, and in learning how best to control them and increase the resistance of both natural
and human-dominated ecosystems to invasion.
 GENERAL INTRODUCTION n xxxi

For those with a more “objective,” less value-driven perspective on invasive species,
Joseph Grinnell’s 1919 paper, “The English House Sparrow Has Arrived in Death Valley:
An Experiment in Nature,” is a harbinger of the study of invasives as a way to understand
basic biogeographical, ecological, or evolutionary processes. Three papers by Richard
Johnston and Robert Selander (1964, 1973, 1973) demonstrated that evolution in nonnative
populations could be fast. They focused on the House Sparrow, which in a little more than
100 generations had diverged into distinct regional morphs based on size and plumage, dis-
playing well-known biogeographic patterns described by Bergmann’s Rule and Gloger’s
Rule, respectively. In 1964, H. G. Baker and G. L. Stebbins convened a conference on “The
Genetics of Colonizing Species” and published contributed papers the following year. The
1960s also saw publication of the paradigm-changing book The Theory of Island
Biogeography by Robert MacArthur and E. O. Wilson, which modeled how new species fit
into existing communities, envisioning an equilibrium number of species on an island that
meant that a new colonizer would cause the extinction of a previously established species.
Experimental work by Daniel Simberloff and others tested this hypothesis, which dominated
ecological research and thought for a couple of decades.
On the conservation/management side of the invasive species issue, George Laycock’s
1966 Alien Animals set the tone. Laycock condemned the negative consequences and the
human foibles associated with transplanting animals to new locations and strongly advo-
cated against future introductions. Laycock helped bring exotic animals in general to the
attention of the public and to resource managers, but may also have biased opinions against
all nonnative species. Nonetheless, he stimulated research on nonnatives at a time when sci-
entists were focusing their efforts on supposedly pristine or natural ecological systems. By
the 1970s, many scientific journals began to accept articles on invasive species, although
some range and wildlife management journals still refused manuscripts on nonnatives.
Invading plants and animals increasingly became important research subjects in efforts to
discover and understand ecological and evolutionary patterns and processes.
International interest in invading species as modifiers of natural communities with the
potential to drive or at least exacerbate system-wide changes was growing at the same time
research was developing in the United States. In the early 1980s, the Scientific Committee on
Problems of the Environment (SCOPE), part of the International Council of Scientific Unions
(ICSU), launched a program on the “Ecology of Biological Invasions.” The program sponsored
scientific meetings in Great Britain, South Africa, Australia, and the Netherlands, as well as a
major symposium in Asilomar, California, and a final convention to synthesize results in
Hawai’i in 1986. This global initiative resulted in 15 volumes, including Ecology of Biological
Invasions of North America and Hawaii, edited by H. A. Mooney and J. A. Drake (1986). The pro-
gram addressed two questions that remain at the core of invasion ecology today: what makes a
good invader, and what determines whether an ecosystem is prone to, or resistant to, invasion.
Invasion ecology had gained a separate space within ecology by the beginning of the
twenty-first century, as attested to by a spate of recent textbooks and other works devoted
to invasion ecology or invasion biology (e.g., Cadotte et al. 2006; Lockwood et al. 2007;
Ruiz et al. 2003; and Sax et al. 2005). Increasingly, invasive species were also being seen
as a component of global environmental change, expected to both affect and be affected by
climate change, altered nutrient cycles and disturbance regimes, and changing land-use pat-
terns. Another project of SCOPE, Invasive Species in a Changing World (Mooney and Hobbs
2000), highlighted some of these problems and also brought attention to the implications
of social views, monetary costs, and the global economy to the management and control of
invasive species.
xxxii n GENERAL INTRODUCTION

The Human Factor

Species that are invasive today depended on human choices, values, activities, and sys-
tems for their initial introduction as well as establishment and further spread. As outlined
above, almost all arrived in their receiving habitats as a consequence of their association with
humans. (The Cattle Egret is a major exception and, according to some definitions, may not
really be “invasive.”) They have followed people from place to place either surreptitiously or
as desired members of the human entourage. Most gained their first footholds in or near
human habitation or sites of commerce and then found suitable habitats in areas where
native plant and animal communities had been disturbed or destroyed by forest clearing,
cultivation, domestic livestock grazing, or urban development. In continental situations in
particular, their successful entry into wild and semi-wild ecological communities is more
likely when some native elements have already been removed or weakened by, for example,
predator control, fire suppression, selective grazing, trampling, or a changing climate.
Invasive species reflect us: our history of colonization, exploitation, and trade; our changing
technology; our human curiosity and aesthetics. Even our love of life and freedom come into
play when we release domesticated and pet animals to the wilds.
How one views invasive species and deals with them also reflects human values. Individual
rights, property rights, animal rights, the right to make a living, all influence whether or not
we take measures to prevent introductions, eradicate populations before they begin to spread,
or manage species that are beyond the stage where eradication is possible. Some of the largest
offenders today in terms of bringing new species into the country are the pet trade and horticul-
tural/nursery trade. Inspection and quarantine are expensive, and it is difficult to prove cost sav-
ings from species that do not make it through our ports of entry and to gain public support for
such measures. Conflicting viewpoints between conservationists and animal rights groups can
make management programs unwieldy at best. But who is right? It depends on what
Americans want and value as the natural heritage they pass to future generations. Informed deci-
sion making is paramount. People should at the very least know the risks and trade-offs. In the
entries contained in the two volumes of this encyclopedia, we try to present a sample of species,
pathways, and impacts—actual and potential—so the reader can be better able to make these
decisions. Many problems arise because we simply do not understand or are unaware of the con-
sequences of our actions. The species accounts that follow will help the reader identify invasive
species and learn some of the ways each of us can slow their spread or best manage them as fel-
low inhabitants of our land. The choice to do something or nothing is ours.

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n Preface
Invasive species have gained our attention in different ways. Susan Woodward, who wrote
about invasive microorganisms, fungi, and animals in Volume 1, had her interest in invasive
animals first sparked as a student of biogeography in the 1970s. Birds such as the European
Starling and House Sparrow were featured in textbooks to demonstrate how animals spread
in an environment that was new to them or how quickly they evolved adaptations to varying
local conditions across a whole continent. Graeme Caughley’s work on irruptions of red deer
in New Zealand was new, and the modeling of invasions and management of exotics in their
infancy. As a doctoral student at UCLA, she studied feral burros along the lower Colorado
River, viewing them as an example of humans “changing the face of the earth” (the buzz-
words of those days) by transporting domesticated and wild animals around the world.
Under contract to the Bureau of Land Management (BLM) at that time, she collected baseline
data on population dynamics, diet, home range size, and other aspects of burro behavior
and ecology that would help that agency devise policies and practices for the animals’ man-
agement. With an applied aspect to her work, she straddled what has become two main per-
spectives on invasive species in general: an academic interest in the science of invasions and
a management interest in preventing arrivals, eradicating or controlling the spread of those
species that were able to establish populations, and managing those whose numbers and dis-
tribution were for all practical purposes already beyond eradication.
Joyce Quinn, a biogeographer whose research dealt with distribution of plants and their
relationships with the natural environment, such as climate and soils, wrote the invasive
plants section in Volume 2. In spite of her experience, she found that researching and writ-
ing this book led her to learn more. Some plants that she had thought were an integral part
of the “natural” landscape, such as common mullein, are actually alien plants that had
become naturalized and are now widespread in the United States. She notes:

Several years ago, an uninvited plant sprouted unexpectedly in my yard. I tried for years
to get rid of it, dutifully pulling off the little sprouts as they emerged. After four or five
years, I gave up and decided to let the plant grow. In a couple of years, it became an
attractive tree about 10 ft. (3 m) tall. It had smooth speckled bark, long lacy compound
leaves, and clusters of small purple star-shaped flowers. As I was doing research for this
Encyclopedia, I discovered that my new plant was a chinaberry tree. While attractive, it
had few redeeming qualities so I decided to eliminate it. A friend helped me saw the
trunk, about 8 in. (20 cm) in diameter, slightly below ground level. I immediately poured
undiluted glyphosate on the freshly cut stump, thinking that was the end of it. I paid no
attention until four months later when I saw a 6 in. (15 cm) sprout! I sprayed it with her-
bicides, but another sprout soon emerged. I sprayed again, but at the time of this writing,
I still do not know if I have managed to kill the invader. I fear that it will be an on-going
process. If I have had such trouble with just one alien invasive plant, the challenges that
land managers, conservationists, and agriculturalists have in battling invasive species
seem insurmountable.
xxxvi n PREFACE

Scope
The purpose of the Encyclopedia of Invasive Species is to provide an introduction to the species,
issues, and management options involved with invasive animals, fungi, microorganisms, and
plants. The number of plants and animals introduced into the United States is staggering.
Only a relatively few establish self-sustaining populations, and very few of these actually become
invasive (in the scientific sense of greatly and rapidly expanding their range in the United States).
Still, there are hundreds of invasive species—too many to be included in a reference book of this
sort. For many species, much remains to be learned, and it is premature to develop full entries
for them, but this still leaves many to choose from. In selecting the 168 species for inclusion
in the Encyclopedia, we have tried to offer a wide spectrum of invasive species that includes some
present in the United States from colonial times, and some that have just been detected; some
that completed their spread across the country long ago, and others that are in the midst of rapid
population growth and range expansion. We also wanted to include some species that are found
throughout the country, and some that are limited to a region or single state.
For animals, we aimed to include representatives from all major classes of vertebrates and
a good variety of invertebrates. The reader will find common, well-known invaders and
others that may be a surprise. We also wanted to showcase a few fungi, especially those that
have been major transformers of urban, suburban, and natural forests, and at least acknowl-
edge the presence of invading microorganisms with a tiny sample of those threatening the
health of native animals and, in some cases, humans as well. Finally, we wished to have a
geographically broad selection of invasive animals, with all 50 states and Puerto Rico having
some members of their nonindigenous fauna represented. Florida, Hawai’i, and California
have the largest numbers of officially recognized invasive species. Residents of these states
will undoubtedly find nonnative organisms causing significant impact in natural and artifi-
cial ecosystems missing from our accounts. This was necessary in order to include some
organisms limited to other states.
For plants, we also tried to include a little bit of everything. Volume 2 addresses a variety
of growth forms, ranging from aquatic plants to trees and vines, and all regions of the United
States. Some plants are widespread throughout the country, while others are localized. Many
plants were deliberately brought to the United States as ornamentals or for some useful char-
acteristic, while others were accidentally introduced. The length of entries dedicated to each
species is variable. The taxonomic relationships of some plants and similar species are not
always clearly defined. Some plants, for example, hybridize so freely that it becomes difficult
to distinguish different species. A few accounts of invasive plants treat two or more related
species in the same entry because their effects and management are similar. As with animals,
we could always find “one more” species that should be included, but it was not possible to
include all. A wealth of information from various sources can be accessed by the reader
wanting to know more. The General Bibliography at the end of Volume 2 has a list of recom-
mended resources, including websites.
The Encyclopedia is specifically meant for high school and college students, but addresses
many of the informational needs of the curious naturalist, horticulturalist, or any home-
owner or environmentally concerned citizen who is interested in the origins and consequen-
ces of invasive plants and animals.
Although some invasive species have been part of the landscape of the United States for lit-
erally hundreds of years, the wide-reaching effects of most are only beginning to become real-
ized. Some invasive species are detrimental to native ecosystems and threaten biodiversity,
others are more economically damaging to crops and livestock, and a few pose a danger to
 PREFACE n xxxvii

human health. Invasive species are a major part of current global environmental change.
Experts consider them the second-greatest threat to native species after habitat destruction
and fragmentation. The control and interdiction of invasive species coupled with the damage
some incur on crops, pastures, livestock, native ecosystems, and human health and well-
being costs billions of dollars each year. The invasive species problem is dynamic—as those
in the Mid-Atlantic states weathering their first onslaught of the brown marmorated stink
bug know well—and endlessly fascinating.

How to Use the Encyclopedia

Volume 1 begins with an introduction to inform the reader of the nature and scope of issues
related to invasive species in the United States. Separate sections deal with the terminology
related to invasive species, the invasion process from an ecological point of view, the pathways
by which nonnative species have been and continue to be introduced to the United States,
some of the ecological and economic impacts of invasives, and a brief outline of the history of
modern invasion science. A final section of the introduction describes the human factors that
determine what species come in, where they succeed, and if and how they are managed.
The introduction is followed by 88 entries describing microorganisms, fungi, inverte-
brates, and vertebrates. Entries are arranged alphabetically within major taxonomic groups.
The species described represent the large number introduced and invasive in the continental
United States, Hawai’i, and Puerto Rico.
Each entry in both volumes includes the following elements, unless noted otherwise:

Native Range
Distribution in the United States
Description
Related or Similar Species
Introduction History
Habitat
Diet (animals only)
Life History (animals, fungi, and microorganisms only)
Reproduction and Dispersal (plants only)
Impacts
Management
Selected References

Additionally, each entry in both volumes is accompanied by at least one photograph and
maps that show the original and invasive range of the species in question according to the
best information available. Often range maps are, by necessity, approximate. This is espe-
cially true for organisms not native to the United States or Europe, where biological surveys
are more complete than on other continents.
In Volume 1, the entries are followed by a list of state-by-state occurrences of invasive
animals, fungi, and microorganisms; a glossary; and an index to both volumes.
Volume 2 begins with a brief overview of invasive plants in the United States, which
loosely follows the organization within species accounts, describing in general the scope of
xxxviii n PREFACE

the invasive plant problem, including the ways, both intentional and accidental, that plants
were brought into the country; and some of the effects invasive plants have on native plant
and animal species, natural ecosystems, agricultural or fishing industries, recreational activ-
ities, or human health. The ways in which invasive plants reproduce and expand their range
is summarized, as is information on management and prevention of invasive plants species.
A sidebar on herbicides accompanies the overview.
The 80 entries on invasive plants are arranged by growth form categories: aquatics, forbs,
graminoids, shrubs, trees, and vines. Photographs of each species show different parts of the
plant. Interesting facets of a plant’s use or history or of strategies attempted for its control are
related in sidebars.
Several supplementary lists follow the invasive plant entries to provide background infor-
mation, and various tables summarize plant data in different, easily accessible ways, includ-
ing a table of common and scientific names of both plants and animals briefly mentioned in
the text of Volume 2, and a list of organizations concerned with invasive plants in the United
States. Two tables of noxious or invasive plants, one organized by state and the other by spe-
cies, as well as a table of species listed by type of impact are also available.
Volume 2 concludes the set with these appendices to the Encyclopedia: a list of American
species that are invasive in other parts of the world; a list of federal laws related to the pre-
vention and management of invasive species; international agreements and conventions
dealing with invasive species; and the IUCN/SCC Invasive Species Specialist Group’s list of
100 of the “World’s Worst Invasive Alien Species,” with an indication of those covered in
the Encyclopedia. The glossary, a selected bibliography of classic and contemporary writings
and online information sources, and the index to the set complete Volume 2.
It is our hope that our efforts will stimulate thought and make the natural world more
accessible to the general public. Informed readers can help make the decisions that will cur-
tail the spread of species that have only recently arrived, prevent the arrival of yet others, and
manage those that are currently invasive

Acknowledgments
Both authors thank the photographers who graciously allowed their photos to be used,
often donating them, or sometimes providing them at a reduced fee. They deserve our spe-
cial thanks for giving life to the species descriptions. Bugwood.org and its associated person-
nel at the Center for Invasive Species and Ecosystem Health, University of Georgia, deserves
special mention as a clearinghouse for providing informational sources and photographs.
Joyce Quinn prepared the excellent maps for the species accounts.
Each author is most appreciative of the other’s contributions to the development of the
project, the overall organization of the volumes, and constructive critiques of text and illus-
trations throughout the manuscript preparation process. We complement each other and
work well as a team. We acknowledge Kevin Downing, originally of Greenwood Press and
now serving the broader ABC-CLIO community as editorial operations manager, who initi-
ated the proposal for the Encyclopedia set, and David Paige of ABC-CLIO, who guided us
through subsequent discussions and organizational details. Anne Thompson, development
editor, later offered guidance in the specifics of the manuscript, and Erin Ryan helped with
the specifications for the illustrations. We thank all four for creating a positive and flexible
working environment and offering valuable suggestions all along the way.
n Alphabetical List of Invasive
Microorganisms, Fungi, and Animal
Entries
Entries in the encyclopedia are arranged by categories. Following are the entries in Volume 1
in alphabetic order.

African Clawed Frog (Xenopus laevis)

Africanized Honey Bee (Apis mellifera scutellata)
Alewife (Alosa pseudoharengus)
American Bullfrog (Lithobates catesbeianus)
Argentine Ant (Linepithema humile)
Asian Clam (Corbicula fluminea)
Asian Green Mussel (Perna viridis)
Asian Longhorned Beetle (Anoplophora glabripennis)
Asian Swamp Eel (Monopterus albus)
Asian Tiger Mosquito (Aedes albopictus)
Australian Spotted Jellyfish (Phyllorhiza punctata)
Avian Malaria (Plasmodium relictum capistranoae)
Bat White-Nose Syndrome Fungus (Geomyces destructans)
Bighead Carp (Hypophthalmichthys nobilis)
Black Rat (Rattus rattus)
Brown Anole (Norops[=Anolis] sagrei)
Brown Marmorated Stink Bug (Halyomorpha halys)
Brown Trout (Salmo trutta)
Burmese Python (Python molurus bivittatus)
Cattle Egret (Bubulcus ibis)
Chain Tunicate (Botrylloides violaceus)
Chestnut Blight Fungus (Cryphonectria parasitica)
Chinese Mitten Crab (Eriocheir sinensis)
Chinese Mystery Snail (Cipangopaludina chinensis malleata)
Chytrid Frog Fungus (Batrachochytrium dendrobatidis)
Colonial Tunicate (Didemnum vexillum)
Common Bed Bug (Cimex lectularius)
Common Myna (Acridotheres tristis)
Common Periwinkle (Littorina littorea)
xl n ALPHABETICAL LIST OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMAL ENTRIES

Coqui (Eleutherodactylus coqui)

Cuban Treefrog (Osteopilus septentrionalis)
Dutch Elm Disease Fungi (Ophiostoma novo-ulmi and O. ulmi)
Emerald Ash Borer (Agrilus planipennis)
Eurasian Collared-Dove (Streptopelia decaocto)
European Earthworms (Lumbricus terrestris, L. rubellus, Aporrectodea caliginosa,
Dendrobaena octaedra, and others)
European Starling (Sturnus vulgaris)
Feral Burro (Equus asinus)
Feral Cat (Felis silvestris catus)
Feral Goat (Capra hircus)
Feral Horse (Equus caballus)
Feral Pig (Sus scrofa)
Formosan Subterranean Termite (Coptotermes formosanus)
Giant African Snail (Achatina fulica)
Gizzard Shad (Dorosoma cepedianum)
Glassy-Winged Sharpshooter (Homalodisca vitripennis)
Golden Apple Snail (Pomacea canaliculata)
Grass Carp (Ctenopharyngodon idella)
Green Crab (Carcinus maenas)
Green Iguana (Iguana iguana)
Gypsy Moth (Lymantria dispar)
Hemlock Woolly Adelgid (Adelges tsugae)
Honeybee Tracheal Mite (Acarapis woodi)
House Finch (Carpodacus mexicanus)
House Mouse (Mus musculus)
House Sparrow (Passer domesticus)
Indian Mongoose (Herpestes javanicus)
Japanese Beetle (Popillia japonica)
Japanese White-Eye (Zosterops japonicus)
Lacy Crust Bryozoan (Membranipora membranacea)
Lionfish (Pterois volitans /P. miles)
Lyme Disease Bacterium (Borrelia burgdorferi)
Monk Parakeet (Myiopsitta monachus)
Mosquitofish (Gambusia affinis and G. holbrooki)
Multicolored Asian Lady Beetle (Harmonia axyridis)
Mute Swan (Cygnus olor)
Naval Shipworm (Teredo navalis)
New Zealand Mud Snail (Potamopyrgus antipodarum)
Nile Monitor (Varanus niloticus)
ALPHABETICAL LIST OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMAL ENTRIES n xli

Northern Snakehead (Channa argus)

Norway Rat (Rattus norvegicus)
Nutria (Myocastor coypus)
Quagga Mussel (Dreissena rostriformis bugensis)
Rainbow Trout (Oncorhynchus mykiss)
Red Imported Fire Ant (Solenopsis invicta)
Rock Pigeon (Columba livia)
Round Goby (Neogobius melanostomus)
Rusty Crayfish (Orconectes rusticus)
Sea Lamprey (Petromyzon marinus)
Silver Carp (Hypophthalmichthys molitrix)
Spiny Water Flea (Bythotrephes longimanus)
Spotted Tilapia (Tilapia mariae)
Sudden Oak Death (Phytophthora ramorum)
Varroa Mite (Varroa destructor)
Veined Rapa Whelk (Rapana venosa)
Walking Catfish (Clarias batrachus)
West Nile Virus (West Nile Virus)
White Pine Blister Rust (Cronartium ribicola)
Zebra Mussel (Dreissena polymorpha)
This page intentionally left blank
n Microorganisms
n Avian Malaria
Scientific name: Plasmodium relictum capistranoae
Family: Plasmodiidae
Native Range. Eurasia. The exact place of origin is unknown, but genetic evidence
strongly suggests it arose somewhere in Eurasia, and it is known to infect Eurasian birds
such as House Sparrows (Passer domesticus) and Common Mynas (Acridotheres tristis), both
of which have been introduced to Hawai’i.
Distribution in the United States. Hawai’i. Avian malaria is most common at elevations of
3,000–5,000 ft. (900–1,500 m) on the moist windward sides of all the main islands.
Description. This parasitic protozoan requires microscopic examination of blood and tis-
sues for identification. The clinical symptoms of malaria in Hawai’i’s endemic honeycreepers
(Drepaniidae) include weight loss, lethargy, lack of appetite, and high death rates. Infected
birds will have a prominent sternum or breast bone (keel). Necropsies reveal enlarged,
chocolate-brown or black livers and spleens and thin, watery blood; up to 50 percent of
circulating red blood cells are infected by the microorganism.
Related or Similar Species. Other Plasmodium species are responsible for malaria in
reptiles, birds (including poultry), and mammals, including humans. Different subspecies
of P. relictum are known to infect birds in other parts of the world.
Introduction History. The introduction of avian malaria to Hawai’i required the arrival of
both the parasite and a vector. The mosquito vector was unintentionally brought to Hawai’i
in 1826. It seems to have reached Maui first and may have come in water barrels aboard a
ship sailing from Mexico. The hundreds of exotic birds released in the islands likely led to
the accidental introduction of Plasmodium relictum in the early twentieth century. Species
such as the House Sparrow and Common Myna, from Europe and India, respectively, are
the most probable sources. With no prior exposure to the parasite, native birds quickly fell
victim to avian malaria.
Habitat. Avian malaria occurs where its vector species live. In Hawai’i, the vector is
the southern house mosquito (Culex quinquefasciatus), itself an introduced species. This
mosquito is most common at elevations below 5,000 ft. (1,500 m), and avian malaria is
most prevalent among native birds in moist lowland forests. Cool temperatures prevent
the development of mosquito larvae, and temperatures below 55°F (13°C) restrict the devel-
opment of malarial parasites in adult mosquitoes. Recent studies, however, suggest that the
mosquito—and hence the disease—is beginning to occur at higher elevations in the islands.
This may be a product of warmer summer temperatures.
The disease is found in dry habitats if water is seasonally available. Mosquito breeding
sites occur in standing water, including ditches, stock ponds, the artificial containers
commonly found in human settlements, tree fern cavities, pools in intermittent streams,
and wallows made by feral pigs. Cavities in lava flows also trap water and create corridors
for mosquitoes to move between forest fragments.
 2 n MICROORGANISMS

Life History. P. relictum

undergoes both sexual and asex-
ual reproduction at different
stages of the life cycle in both its
bird and mosquito hosts. When
a mosquito bites an infected bird,
it will ingest gametocytes, the
male and female reproductive
cells of the protozoan. In the gut
of the mosquito, the gametocytes
produce encysted cells that
develop into motile spores (spo-
rozoites) which move to the sali-
vary gland and are injected into
a bird when the mosquito again
feeds on blood. Thus the sporo-
zoites are the infectious stage of
the protozoan. In the bird, sporo-
zoites attack red blood cells and
develop within them, replicating
their nuclei and other organelles.
The resulting multinucleated
cells, called schizonts, grow and
cause the red blood cell that they
have infected to burst. The schiz-
onts then break apart and release
single-nucleus daughter cells
(sporozoites) into the blood-
stream. (These daughter cells
produce toxins that, in human
malaria, cause the characteristic
Top: The place of origin of the protozoan that causes avian malaria is chills and fever.)
unknown, but somewhere in Eurasia is likely. Bottom: Avian malaria Impacts. Avian malaria in-
currently only affects birds in Hawai’i. fects passerine (perching) birds,
which in Hawai’i include the
Hawaiian Crow (Corvus hawaiiensis) and honeycreepers, a group of 57 birds (half of which are
now extinct), all believed to have evolved from a single finch-like ancestor and all endemic to
the islands. The introduction of avian malaria to the Hawaiian Islands has been implicated in a
modern wave of population reductions, range restrictions, and possibly extinction of a number
of Hawai’i’s rare birds. Only native bird species, having evolved during a long period of isolation,
seem to be deleteriously affected by the parasite. For some of them, such as the ‘I’iwi (Vestiaria
coccinea) and the Maui ‘Alauahio (Paroreomyza montana), infection by the malaria parasite is
almost always fatal. Other honeycreepers, such as the ‘Omao (Myadestes obscurus), show greater
resistance to the disease. Lowland populations of Hawai’i ‘Amakihi and O’ahu ‘Amakihi
(Hemiagnathus virens complex) in the Puna area of the Big Island appear to have evolved some
resistance to avian malaria since coming into contact with the parasite.
Most surviving honeycreepers today live at elevations above 5,000 ft. (1,500 m), where
both Culex mosquitoes and the malaria protozoan have difficulty reproducing. The
 LYME DISEASE BACTERIUM n 3

Akiapola’au (Hemignathus munroi), for example, once occupied forests as low as 1,600 ft.
(500 m) on the island of Hawai’i, but the remnant populations of this endangered species
are now found only in high-elevation forests. On Maui, the six remaining endemic passer-
ines (four are extinct) inhabit a narrow strip of high-elevation rainforest on the slopes of
Haleakalā Volcano.
Efforts are underway to reestablish some of Hawai’i’s threatened and endangered honey-
creepers in protected patches of high-elevation forest. If mosquitoes continue to expand
their distribution into higher and higher elevations as climate changes, these efforts could
be doomed. Development of resistance to Plasmodium relictum seems to be occurring in some
populations and may help some bird species survive; however, resistant birds still harbor the
parasite and can transmit it to mosquitoes. They serve as reservoirs for the disease, further
threatening vulnerable species.
Management. Control measures are directed at the vector, the southern house mosquito.
The most effective way of preventing the disease from spreading is to eliminate or reduce
populations of the mosquito by removing man-made habitats of standing water where the
mosquito breeds or applying Bt (Bacillus thuringiensis israelensis) to impoundments such as
horse watering troughs that cannot be removed. This attack on mosquitoes has also targeted
feral pigs (see Vertebrates, Mammals, Feral Pig), which dig in the forest floor in search of
food and leave behind depressions that catch rainwater and harbor mosquito larvae. The
pigs also push over tree ferns and eat the starchy pith, creating water-holding cavities for
mosquitoes to breed in.

Selected References
Atkinson, Carter T. “Ecology and Diagnosis of Introduced Avian Malaria in Hawaiian Forest Birds.”
USGS FS 2005-3151. Pacific Island Ecosystems Research Center, U.S. Geological Survey, 2005.
http://biology.usgs.gov/pierc/Native_Birds/Avian_malaria.pdf.
Atkinson, Carter T., and Dennis A. LaPointe. “Plasmodium relictum (Micro-organism).” ISSG Global
Invasive Species Database, 2005. http://www.invasivespecies.net/database/species/ecology.asp
?si=39&fr=1&sts.
Beadell, Jon S., Farah Ishtiaq, Rita Covas, Martim Melo, Ben H. Warren, Carter T. Atkinson, Staffan
Bensch, Gary R. Graves, Yadvendradev V. Jhala, Mike A. Peirce, Asad R. Rahmani, Dina M.
Fonseca, and Robert C. Fleischer. “Global Phylogeographic Limits of Hawaii’s Avian Malaria.”
Proceedings of the Royal Society B: Biological Science, 272(1504): 2935–44. Published online
August 22, 2006. http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1639517/. doi:10.1098/
rspb.2006.3671.
LaPointe, Dennis A. “Feral Pigs, Introduced Mosquitoes, and the Decline of Hawai’i’s Native Birds.”
USGS FS 2006-3029. Pacific Island Ecosystems Research Center, U.S. Geological Survey, 2006.
http://biology.usgs.gov/pierc/Fact_Sheets/Pigs_and_mosquitoes.pdf.

n Lyme Disease Bacterium

Scientific name: Borrelia burgdorferi
Phylum: Spirocheates
Native Range. North America and Europe. B. burgdorferi sensu stricto occurs in the United
States, while two other strains—B. burdorferi garinii and B. burgdorferi afzelli—are found in
Europe.
Distribution in the United States. Maine to Virginia; Great Lakes region; northern
California.
 4 n MICROORGANISMS

Description. B. burgdorferi
has the characteristic corkscrew
appearance of spirochetes in
general. Its clinical manifesta-
tion as Lyme disease in humans
is marked first by a diagnostic
bull’s-eye rash that usually sur-
rounds a tick bite and expands
with time (erythema migrans).
The rash may last for 2–3 weeks
and be accompanied by flu-like
symptoms. Untreated, stage 2
symptoms develop, including
intermittent inflammatory ar-
thritis, facial palsy, and extreme
fatigue and malaise. In stage 3
of the disease, inflammation of
the brain (encephalitis) and
spinal chord (myelitis) and
weakening of the lower limbs
(paraparesis) occur; symptoms
of fibromylagia can develop.
The microscopic spirochete
is transmitted to humans when
they are bitten by infected ticks.
In the eastern and midwestern
United States, the black-legged
or deer tick (Ioxodes scapularis)
is the vector; in the Pacific
Northwest, the western black-
legged tick (I. pacificus) is
Top: The spirochete responsible for Lyme disease is known from both involved. Adult deer ticks are
North America and Europe, but the disease was first reported in Europe, teardrop-shaped and 0.1 in.
making that continent the more probable place of origin. Bottom: (3 mm) long, about the size of
Borrelia burgdorferi infects ticks throughout the lower 48 states, but it is an apple seed. Females, the only
most prevalent in the Northeast, western Great Lakes region, and parts ones that seek blood meals, have
of northern California. (Adapted from American Lyme Disease
black heads and dorsal shields
Foundation, “U.S. Maps and Statistics,” http://www.aldf.com/usmap
.shtml.) and dark red abdomens. In
males, the hard shield or scutum
covers the entire back, and the head and whole body are black. The nymphs of these tiny arach-
nids are about the size of a poppy seed, have black heads and translucent bodies; they are most
apt to transmit Lyme disease to people because they are so small that they may not be noticed
and removed. Western black-legged ticks are very similar in appearance.
Introduction History. Lyme disease was first recognized in 1975 by Dr. Allen Steele of
Yale University, when about 50 children in the town of Lyme, Connecticut, developed
rashes and joint pain, swelling, and inflammation similar to arthritis. Indeed, the new dis-
ease was originally called “Lyme arthritis.” It was soon determined that the symptoms were
similar to a tick-borne infection that had been known in Europe since at least 1883, when
 LYME DISEASE BACTERIUM n 5

it was reported by the German physician Alfred Buchwald. It was not until 1982, however,
that Dr. Willy Burgdorfer discovered the causative agent of the disease in black-legged ticks.
The spirochete was named in his honor.
Since 1975, the incidence of the disease has increased and spread in the United States,
where it is now the most common tick-borne illness. More than 20,000 new cases are
reported each year. Ten states account for more than 90 percent of occurrences: Delaware,
Maryland, Massachusetts, Minnesota, New Jersey, New York, Pennsylvania, Rhode Island,
and Wisconsin. The increasing prevalence of the disease is due to a variety of factors, includ-
ing better identification and record keeping; encroachment of residential areas into tick hab-
itats; exploding deer and tick populations associated with suburban sprawl and forest
fragmentation, as well as reforestation in the northern United States; and the range expan-
sion of black-legged ticks into newly available habitats.
Habitat. The tick vectors of Lyme spirochetes inhabit the understory of moist, deciduous
forests and open grassy areas with tall vegetation. It is often found along paths and
roadsides.
Diet. Blacklegged ticks are external parasites on warm-blooded animals. They require a
blood meal in order to molt and develop to the next stage of life. Field mice, especially the
white-footed mouse (Peromyscus leucopus), serve as important hosts for larvae. Nymphs
and adults feed on deer, dogs, and humans.
Life History. The bacterium B. burgdorferi circulates between ticks and a variety of verte-
brates, each species affecting the survival of the spirochete because of varying competence
as host species and different rates of infestation by ticks. Ticks can obtain the spirochete as
larvae. Hatching during the summer, a larval tick waits on the ground for a small mammal
or bird to come into contact with it, whereupon it attaches to the passing animal and begins
feeding, sucking blood for a few days. If the host, most commonly a white-footed mouse,
is infected with the pathogen, the larval tick will likely also become infected. (Mice serve
as “reservoir hosts”: they easily acquire the spirochete and sustain viable bacteria in their
blood, allowing it to increase in numbers. They accept tick larvae again and again, and

A. The corkscrew shape of the Borrelia spirochete. (Centers for Disease Control and Prevention.)
B. Black-legged or deer tick (Ioxodes scapularis), the vector for Lyme disease in eastern and central
states. (Hardin MD/University of Iowa and CDC, http://www.lib.uiowa.edu/hardin%5Cmd/cdc/
1669.html.) C. Western black-legged tick (I. pacificus), adult female. This arachnid is the vector for
Lyme disease in western states. (Hardin MD/University of Iowa and CDC, http://www.lib.uiowa.edu/
hardin/md/cdc/ticks5.html.)
6 n MICROORGANISMS

thereby can transmit the bacteria to other larvae. Mice themselves show no signs of disease.)
Once the larva has gorged itself, it will not feed again, but drops off its host and, in the fall,
molts into a nymph. Nymphs remain inactive until early the following spring. When the
nymphs become active, they wait in ambush on vegetation for a new host to brush past
them. They attach to a deer, person, dog, or cat and feed for 4–5 days. If they were already
infected with the Lyme spirochete as larvae, they may transmit it to the host. If they were free
of infection, they may become infected during this time should the host already carry the
pathogen. Peak activity for nymphs is spring and summer, and this is when humans are
most apt to acquire Lyme disease. Once the nymph is fully engorged with blood, it drops
off the host and molts into an adult. Adults are active throughout the fall and wait on grass
or leaf tips about 3 ft. (1 m) above the ground to attach to deer or other larger mammals,
including people. Adult deer ticks are most active in late fall, but humans are less apt to
become infected at this time of year because the adults are large enough to be seen and
picked off before the spirochete can be transmitted. Adult ticks without hosts become inac-
tive when temperatures drop below 45°F (7°C) and typically find shelter in leaf litter on the
forest floor. With warming temperatures in early spring, the adults seek a final blood meal,
usually on deer, that will allow them to mate. Mating occurs on or off a host. The female lays
some 3,000 eggs under leaf litter and then dies, completing the two-year life cycle.
Infection of humans by B. burgdoferi requires that a tick be attached for at least 24 hours
and more likely for 2–3 days. Only small numbers of the spirochetes occur in a tick until it
feeds. With the intake of blood, the bacteria multiply in the tick’s gut. After 2–3 days, they
migrate to the salivary glands, where they are injected into the host as the tick completes
its feeding. This is why undetected nymphs usually spread the disease to humans. An esti-
mated 85 percent of infected persons received the bacteria from nymphs in the spring; the
other 15 percent obtained the infection from adults in autumn. Only about 1 percent of tick
bites in an area where Lyme disease is prevalent result in infection.
Impacts. Some people are able to clear the infection without developing any symptoms.
In others, the bacteria spread through the body and elicit inflammatory responses in the skin
(erythma migrans) and joints, tendons, and bursae, especially in the knees, ankles, and
wrists. These may be accompanied by fever and general malaise. Untreated, the conditions
last a week or more and often reoccur for up to 10 years. The most common neurological
condition associated with Lyme disease is facial paralysis, which may last for a couple of
months. More serious manifestations of the disease include inflammation of the brain, a
form of meningitis presenting as a headache, stiff neck, and sensitivity to light. It can be
quite debilitating for a long period of time. Most cases are successfully treated with
antibiotics.
Management. Control is aimed at preventing infection of humans. People should try to
avoid tick habitats, particularly in spring when the nymphs are active. Removal of vegetation
that is prime tick, mouse, and/or deer habitat—tall grass, brush, and dead leaves—from
housing and work areas is also advisable. Precautions such as wearing light-colored cloth-
ing, long-sleeved shirts, hats, and closed shoes, plus tucking pant legs into socks or boots,
can delay the attachment of ticks to skin and make them more visible. Insect repellent on
clothing and skin (except on the face) also helps. After outdoor activity, a careful body
inspection should be done, and any ticks should be removed with tweezers. Deer ticks
and western black-legged ticks seek body folds such as armpits, groin, back of the neck,
and back of the knee. Shower and wash clothes in hot water.
Vaccines to prevent Lyme disease are now available for dogs and cats.
 WEST NILE VIRUS n 7

Selected References
“Deer Tick Ecology.” American Lyme Disease Foundation, Inc., 2006. http://www.aldf.com/
deerTickEcology.shtml.
Meyerhoff, John O. “Lyme Disease.” Medscape, 2009. http://emedicine.medscape.com/article/330178
-overview.
Todar, Kenneth. “Borrelia burgdorferi and Lyme Disease.” Todar’s Online Textbook of Bacteriology,
2008. http://www.textbookofbacteriology.net/Lyme.html.

n West Nile Virus

Also known as: WNV
West Nile Virus
Family: Flaviviridae
Native Range. Uncertain. West Nile virus was first described in Uganda and then found
in other parts of Africa as well as in Europe, Southwest and Central Asia, and Australia.
Whether it was native or introduced to these regions has yet to be determined.
Distribution in the United States. West Nile virus has been reported in all of the 48 con-
tiguous states, although outbreaks do not occur in every state every year.
Description. This small flavivirus consists of a positive-sense single strand of RNA (ribo-
nucleic acid) containing between 11,000 and 12,000 nucleotides. The RNA is surrounded
by a protein coat (nucleocapsid) that is encased in a lipid membrane. The complete structure
or virion is spherical and measures 40–65 nm in diameter.
Related or Similar Species. Other flaviviruses cause human diseases, including St. Louis
encephalitis, yellow fever, dengue fever, and Hepatitis C.
Introduction History. West Nile virus was first described from Uganda in 1937.
Subsequently, it was discovered in many parts of the Old World and in Australia and found
to be one of the most widespread flaviviruses in the world. Confirmation of the virus in New
York City in 1999 marked its first appearance in the Americas. Early in the spring of 2000, it
showed up again in mosquitoes and birds and quickly spread to other parts of the eastern
United States. In 2000, it ranged from New Hampshire to North Carolina; by 2001, it had
crossed the Mississippi River; and by 2004, it was in every state except Alaska and Hawai’i.
The source of the initial introduction is unknown. Genetic studies point to origins in the
Mediterranean or Middle East. The strain that reached the United States seems to have
evolved into a more virulent form through the mutations of amino acids in a single gene.
How it reached the United States is a mystery. Once on the continent, the virus spread
geographically in migrating birds.
Habitat. West Nile virus cycles between two hosts, birds and mosquitoes. Other verte-
brates act as dead-end hosts; for though the virus may cause illness in them, it never reaches
high-enough levels in the blood to be transmitted to mosquitoes.
The virus thrives in environments where mosquitoes, especially of the genus Culex,
breed: wetlands, urban areas with artificial containers that collect rainwater, and wherever
else standing water persists for several weeks or more. Recent epidemics are correlated with
unusual hot, dry periods.
Life History. West Nile virus replicates best in certain bird species that become the major
amplification hosts. The virus has been isolated in well over 100 different species. Some are
extremely susceptible to infection and die, while others are very tolerant of infection and
show no signs of impairment. The West Nile virus is transmitted from bird to bird by
 8 n MICROORGANISMS

mosquitoes, the most compe-

tent vectors in the United
States being the southern house
mosquito (Culex quinquefascia-
tus); the northern house mos-
quito (C. pipiens); the white-
dotted mosquito (C. restuans);
C. salinarius, a mosquito of salt
marshes; and in the western
United States, the encephalitis
mosquito (C. tarsalis). These
mosquitoes feed preferentially
on birds, but also feed oppor-
tunistically on the blood of
other vertebrates and can trans-
mit the disease to humans,
horses, and other species. Other
mosquitoes such as Culex nigri-
palpus, a common mosquito
in Florida; the Asian tiger mos-
quito, Aedes albopictus; the
inland floodwater mosquito
(Aedes vexans); and the eastern
treehole mosquito (Ochlerotatus
triseriatus) may also spread
West Nile virus.
Some evidence suggests that
squirrels (Sciurus spp.), eastern
chipmunks (Tamias striatus),
eastern cottontails (Sylvilagus
floridanus), and alligators (Alli-
Top: The West Nile virus was first described in Uganda, but its origins gator mississippiensis) may build
remain unknown. Bottom: The West Nile virus has been reported in all up sufficiently high levels of
of the lower 48 states. the virus in their blood to serve
as reservoirs.
Impacts. During periods in 1999 and 2002–2003 when rates of human infection with
West Nile virus were epidemic, high mortality rates were experienced among several
common songbird species. Indeed, the widespread American Crow (Corvus brachyrhynchos)
is so susceptible to fatal infection that dead individuals serve as indicators of the virus’s pres-
ence in a given area. In some regions of the country, 45–100 percent of crows died during
past outbreaks. Among other birds studied in the eastern United States, sharp population
declines correlating with WNV epidemics occurred in the Blue Jay (Cyanocitta cristata), the
Tufted Titmouse (Baeolophus bicolor), the American Robin (Turdus migratorius), the House
Wren (Troglodytes aedon), the Black-capped and Carolina Chickadee (Poecile atricapillus
and P. carolinensis, respectively) and the Eastern Bluebird (Sialia sialis). The Common
Grackle (Quiscalus quiscula) was hard hit in Maryland. Others species showed great toler-
ance. These included Mourning Dove (Zenaida macroura), Northern Cardinal (Cardinalis
cardinalis), Baltimore Oriole (Icterus galbula), Chipping Sparrow (Spizella passerina), and
 WEST NILE VIRUS n 9

A. Image of West Nile virus particle produced by cryoelectron microscopy. (Purdue University
Department of Biological Sciences.) B. The southern house mosquito transmits the West Nile virus
from bird to bird. (U.S. Geological Survey.)

Catbird (Dumetella carolinensis). Populations of Blue Jays and House Wrens had rebounded
by 2005, after the epidemic in humans had subsided, but populations of other birds
remained low.
In the western states, Black-billed Magpies (Pica hudsonia), House Finches (Carpodacus
mexicanus), and Greater Sage Grouse (Centrocercus urophasianus) have been severely affected.
The ecological impacts of rapid changes in bird populations is unknown, but shifts in
species abundance and therefore possibly ecosystem functioning can be expected. Crows,
for example, are important scavengers and are also predators of the nestlings of other birds
and thereby control population sizes.
Many humans infected with the West Nile virus will develop no symptoms, but about
20 percent of infected people experience flu-like symptoms, including high fever, headache
and body aches, general fatigue, rash, vomiting, and diarrhea. For relatively few—fewer than
1 percent of those known to be infected—the disease is fatal. The virus invades the nervous
system and presents as encephalitis, meningitis, limb paralysis, or acute respiratory failure.
In horses, the symptoms of West Nile virus include weakness in the hindquarters, trem-
ors, muscle rigidity, and paralysis. Between 1999 and 2002, the disease affected some
20,000 equines in the United States and was fatal in an estimated 38–57 percent of cases.
Vaccination became available in 2002, but infection rates remained high.
Management. Control measures focus on eliminating breeding sites for the mosquitoes
that are the vectors for the virus. This is primarily an urban and suburban problem and
10 n MICROORGANISMS

means removing containers such as old tires, drums, bottles, and cans in which standing
water accumulates; repairing leaky pipes and outside faucets; unclogging gutters; and
replacing water in birdbaths, watering troughs, and the like several times a week. People
can minimize their risk of being bitten by mosquitoes by using insect repellent, wearing
clothing that covers arms and legs, and minimizing outdoor activity at dusk and dawn when
mosquitoes are most active. A vaccine is available for horses, but still under development for
humans.

Selected References
LaDeau, Shannon L., A. Marm Kilpatrick, and Peter P. Marra. “West Nile Virus Emergence and Large-
Scale Declines of North American Bird Populations.” Nature 447: 710–13, 2007. doi:10.1038/
nature05829.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “West Nile Virus (Micro-organism).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?si=304&fr=1&sts.
Weiss, Rick. “Bird Species Plummeted after West Nile: National Survey Finds Losses Among Many
Songbirds.” Washington Post, 2007. http://www.washingtonpost.com/wp-dyn/content/article/
2007/05/16/AR2007051601032_pf.html.
“West Nile Virus.” National Institute of Allergy and Infectious Diseases, 2009. http://www
.niaid.nih.gov/topics/westnile/understanding/pages/what.aspx.
“West Nile Virus Detected in Arizona.” Agency Directive, Arizona Game and Fish Department, 2003.
http://www.azgfd.gov/w_c/diseases_west_nile.shtml.
n Fungi
n Bat White-Nose Syndrome Fungus
Scientific name: Geomyces destructans
Phylum: Ascomycota
Family: Heliotiaceae
Native Range. Unknown. The fungus associated with bat white-nose syndrome may be
native to Europe or may represent a soil fungus that has recently mutated in the northeastern
United States to become a pathogen of hibernating bats.
Distribution in the United States. Geomyces destructans occurs from Vermont south to
Virginia and west into Tennessee and Missouri. It probably also occurs in northwest
Oklahoma.
Description. This fungus manifests itself as a white coating on the nose, ears, and wing
membranes of hibernating bats. The dense covering of fine hyphae can be removed through
grooming, but scars often remain. Under a microscope, the fungus is distinguished by asym-
metrically curved conidia, the asexually produced spores. These conidia and the fact that the
fungus has very low optimal temperatures for growth confirmed G. destructans was a new
and separate species in the genus Geomyces.
Introduction History. Bat white-nose syndrome was first reported from Howe’s Cave near
Albany, New York, during the winter of 2006–2007. Three years later, its presence was
confirmed in caves and mines in Connecticut, Massachusetts, New Jersey, Pennsylvania,
Vermont, Virginia, and West Virginia. In February 2010, the disease was reported in
Tennessee. It has also expanded into eastern Missouri, as well as eastern Canada. Its origins
and potential for further spread remain unknown.
Habitat. G. destructans proliferates on many organic surfaces in the dark, cool, high-
humidity environments of caves and underground mines, where it colonizes the skin of
hibernating bats. It fails to thrive at temperatures in excess of 68°F (20°C). Thus this cold-
loving fungus finds optimal growing conditions in the hibernacula used by nonmigratory
bats that must over winter in a dormant state at latitudes above 40° N or at higher elevations
in the southeastern United States.
Life History. Much is yet to be learned about this recently described fungus. It can grow
on a variety of organic substances and seems to persist year round in caves and mines. It
apparently becomes established in the skin tissue of bats when their body temperatures
are lowered to 35°–50°F (2°–10°C) during torpor in the winter. The hyphae penetrate tissue
by entering hair follicles and sebaceous glands. No inflammation or immune response seems
to be elicited in the bat.
Although G. destructans produces what is known as white-nose syndrome, penetration of the
wing membranes by the fungus may be the most detrimental aspect of the disease to
bats. Current high mortality rates are one sign that it is a new pathogen that has not achieved a
balance with its host. G. destructans is spreading rapidly from bat to bat and hibernacula
to hibernacula. Spores easily attach to skin, hair, ropes, and clothing; it is not known for how
long they remain viable outside their subterranean home or just how they disperse to other
 12 n FUNGI

caves. Migrating bats and cavers

likely disperse the fungus to new
caves and underground mines.
Impacts. White-nose syn-
drome has killed some 500,000
insect-eating bats since it was
first discovered in 2006. Dead
bats are usually emaciated, but
exactly how, or indeed if, the
fungus kills them is still to
be determined. In addition to
the tell-tale white encrustations
on ears and wing membranes
and around the nose, affected
bats display unusual winter
behaviors. They tend to come
out of torpor much more fre-
quently than normal and fly
around the cave or mine in
which they have been hibernat-
ing or leave the hibernacula
during the day to fly around out-
side. Bats are found closer to the
cave entrance than normal, and
large numbers of dead bats are
clustered near or just outside
the entrance. It may be that the
fungus is such a skin irritant that
it rouses the bats from hiberna-
tion. When they arouse from
torpor and fly around, they use
Top: It is not known if Geomyces destructans originated as a mutation in a vital body fat reserves, which
soil fungus native to the northeastern United States or if it is native to cannot be replenished because
Europe. Bottom: Areas in the United States where bat white-nose insects are not available for them
syndrome had been reported as of June 2010. (Adapted from map by to feed upon during the winter.
Cal Butchkoski, Pennsylvania Game Commission.) Ultimately, the bats may starve
to death.
Infection of the wing membranes may exacerbate the situation since the membranes play
important physiological roles in regulating body temperature, blood pressure, water bal-
ance, and gas exchange. It may also be that infected bats disturb other nonaffected bats in
the colony and rouse them from hibernation, too. Bats that have been sampled as they enter
hibernacula in the fall seem to be healthy, strongly suggesting they acquire the pathogen in
the winter cave.
Currently, six species of bats are known to be affected by white-nose syndrome. The most
dramatic losses have been suffered by the little brown bat (Myotis lucifugus), which has under-
gone population declines of 93 percent in some caves. The Indiana bat (Myotis sodalis), a
federally endangered species, has seen declines of 53 percent in certain hibernacula. Other
bats affected include northern long-eared bat (Myotis septentrionalis), tri-colored bat
 BAT WHITE-NOSE SYNDROME FUNGUS n 13

A. Little brown bat (Myotis lucifugus) affected by white-nose syndrome, Greeley Mine, Vermont.
(Marvin Moriarty, U.S. Fish and Wildlife Service.) B. Hibernating bats with white-nose syndrome,
New York. (Nancy Heaslip, New York Department of Environmental Conservation.) C. Geomyces
destructans infection on wing membrane of bat. (Ryan von Linden/New York Department of
Environmental Conservation.)

(Perimyotis subflavus; formerly known as Eastern pipistrelle [Pipistrellus subflavus]), eastern

small-footed myotis (Myotis leibii) and big brown bat (Eptesicus fuscus). The fungus has
recently been found in caves used by another endangered species, the Virginia big-eared
bat (Corynorhinus townsendii virginianus), although white-nose syndrome has not (yet) been
reported on this bat. In Tennessee, there is concern that it will affect the endangered gray
bat (Myotis grisescens). Most of these bats reproduce slowly, having only a single young each
year, so populations are not expected to rebound quickly from the devastating impacts of
white-nose syndrome.
Such sudden and widespread death among hibernating bats was previously unknown.
Lethal fungal skin infections are rare among mammals, but more common among “cold-
blooded” vertebrates whose body temperatures hibernating bats approximate.
Chytridiomycosis in amphibians (see Chytrid Frog Fungus below) may be an analogous
condition.
Bats have an enormous capacity for consuming flying insects, including moths, mosqui-
toes, and many plant pests. Their loss would represent a major change in natural, suburban,
and agricultural ecosystems in the eastern United States.
Management. While direct bat-to-bat transmission of the fungus is difficult if not impos-
sible to prevent, it is likely that humans transfer G. destructans spores from cave to cave, and
steps can be taken to reduce or halt that process. The U.S. Forest Service has closed caves
and mines to recreational cavers in national forests in 33 states, and many states, caving
clubs, and private owners have followed suit. The public should honor all cave closings in
affected states and adjoining areas to control the spread of this deadly pathogen. Those
entering caves for scientific purposes should carefully decontaminate all clothing and gear
14 n FUNGI

with household bleach or commercially available antibacterial cleaners. Ropes and harnesses
should not be used at all or, if essential, should be dedicated to use in only a single cave. The
U.S. Fish and Wildlife Service has issued containment and decontamination protocols and
provides an online listing of cave closures at http://www.fws.gov/northeast/wnscavers.html.

Selected References
Gargas, A., M. T. Trest, M. Christensen, T. J. Volk, and D. S. Blehert. “Geomyces destructans sp. nov.
Associated with Bat White-Nose Syndrome.” Mycotaxon 108: 147–54, 2009. Available online at
http://botit.botany.wisc.edu/toms_fungi/147gargas9-73.pdf.
“White Nose Syndrome: Could Cave Dwelling Bat Species in the Eastern US Become Endangered in
Our Lifetime?” Bat Conservation and Management, 2010. http://www.batmanagement.com/wns/
wns.html.
“White-Nose Syndrome Threatens the Survival of Hibernating Bats in North America.” USGS Fort
Collins Science Center, 2010. http://www.fort.usgs.gov/wns/.

n Chestnut Blight Fungus

Also known as: Chestnut bark disease fungi
Scientific name: Cryphonectria parasitica
Synonym: Endothia parasitica
Class: Pyrenomycetes
Order: Diaporthales
Family: Valsaceae
Native Range. Mountainous areas of China and Japan.
Distribution in the United States. Eastern states throughout the range of the American
chestnut (Castanea dentata), roughly from Maine west to Michigan and south to Georgia
and Mississippi. It also occurs outside the tree’s range wherever chestnuts have been
planted.
Description. This fungus causes cankers, localized areas of dead tissue, on the trunks of
American chestnut trees. On the surface of the tree, both swollen and sunken cankers form
above the infection; the sunken type expands to girdle the tree. Often orange or yellow fruit-
ing bodies the size of pin heads cover the canker. During moist weather, spores ooze out of
the fruiting bodies, looking like tiny curled orange horns.
Early symptoms of infection include reddish-brown patches on the bark that later
develop into cankers. The bark may crack, and yellow fruiting bodies appear on the surface.
Leaves on affected branches turn brown, but do not fall from the tree for months. The entire
tree above the infection will die as the canker grows, but the roots are not killed. An
American chestnut will continue to resprout from the root collar for decades after the
above-ground part of the tree has died.
Introduction History. The blight was first identified in the United States in 1904 at the
Bronx Zoo in New York City. It may have entered the country as early as 1876, when
Japanese chestnut trees (Castanea crenata) were first imported. By the end of the nineteenth
century, the Japanese species was being offered for sale by most U.S. mail-order nurseries. In
1904, the blight was actually widespread north of Virginia. By 1926, it occurred throughout
the natural range of the American chestnut. Its impact had been so devastating that in 1912,
the U.S. Congress passed the Plant Quarantine Act, the first legal action to stem the flow of
 CHESTNUT BLIGHT FUNGUS n 15

nonnative species into the

United States, in an attempt
to prevent future catastrophes.
The law gave the federal gover-
nment the authority to establish
inspection stations and quaran-
tine areas to intercept and pre-
vent the spread of exotic pests
and pathogens. Today, the
Animal and Plant Inspection
Service (APHIS), part of the
U.S. Department of Agriculture,
is in charge of these activities.
Habitat. In North America,
Cryphonectria parasitica occurs
almost exclusively in American
chestnut trees. It infected mature
trees in natural forests as well as
those planted as ornamentals,
and now attacks the saplings
that still sprout from old stumps
and root systems. The fungus
does also attack Allegheny chin-
kapin (Castanea pumila) and
bush chinkapin (C. alnifolia).
Several oaks (Quercus spp.) also
serve as hosts; but cankers usu-
ally remain small and superficial
on them and do not kill the tree.
Only the post oak (Q. stellata)
seems to be seriously damaged
by the fungus. Other trees on Top: The chestnut blight fungus originated in mountain forests of China
which chestnut blight has been and Japan. Bottom: The former range of the American chestnut
reported include shagbark (Castanea dentata), shown on the map, is a surrogate for the range of the
hickory (Carya ovata), red maple blight fungus; although the blight also affects chestnuts planted beyond
(Acer rubrum), and staghorn the tree’s native range. (Adapted from Saucier 1973.)
sumac (Rhus typhina).
Life History. The chestnut blight fungus produces small, sticky spores in the sexual fruit-
ing bodies visible on the bark’s surface. These so-called ascospores are forcibly ejected
during warm rains and dispersed by the wind or on the feet of birds and insects.
Nonmotile asexual spores called conidia are also produced. The fungal spores enter a tree
through wounds or cracks, often at a branch crotch. The junctions of limbs and the trunk
are particularly vulnerable because the movement of branches in the wind creates small rup-
tures in the bark in the crotches. The spores germinate in the inner bark and cambium,
where masses of thread-like hyphae (mycelia) form into small brown fans. The cushion-
like masses of solid mycelia (stromata) interfere with the flow of nutrients through the
phloem and with the growth of the cambium. On American chestnut, stromata can reach
densities of 8 per in2 (50 per cm2). First the leaves die, then the branches above the point
 16 n FUNGI

A. Developing cankers on young American chestnut tree. (USDA Forest Service-Region 8-Southern Archive,
USDA Forest Service, Bugwood.org.) B. Leaves of the American chestnut. The once-mighty tree is now repre-
sented by root sprouts in the understory. (Linda Haugen, USDA Forest Service, Bugwood.org.) C. A row of dead
American chestnuts after the blight struck. The split rail fence in the foreground is likely constructed of chestnut
timber. (USDA Forest Service-Northeastern Area Archive, USDA Forest Service, Bugwood.org.)

of infection. The sunken cankers can expand enough to girdle the tree within a single grow-
ing season and kill the entire above-ground portion of the tree, or it may take several years.
C. parasitica continues to live in dead wood, but is unable to enter the roots. Chestnuts con-
tinue to resprout for decades. While the sprouts may live for 5–10 years and attain heights of
25 ft. (8.0 m), they rarely live long enough to mature and flower or bear nuts before they,
too, succumb to the blight.
Impacts. The chestnut blight fungus eliminated the American chestnut as a major tree in
eastern forests. These trees were the largest in the broadleaf deciduous forests in the
southern Appalachians, where they commonly grew to heights of 60–90 ft. (18–27 m)
and diameters at breast height (DBH) of 3–5 ft. (1–1.5 m). Exceptional specimens were
120 ft. (35.6 m) tall and 7 ft. (2 m) in diameter. The crown of a single tree could be nearly
100 ft. (30 m) wide. Its sweet nuts were produced in great abundance every year and nour-
ished deer, turkeys, squirrels, and other wildlife as well as people. The fruits that fell to the
ground after the first frost were an important cash crop for many Appalachian families. The
wood was slow to decay and was therefore used for building barns, fences, log cabins, furni-
ture, and coffins. Split-rail fences made of chestnut still persist in woodlands grown up in
abandoned pastureland. The bark and wood was rich in tannic acids and preferred over
the bark of other trees for tanning leather. Large trees with their spreading branches and
 CHESTNUT BLIGHT FUNGUS n 17

reliable nut harvests had been favored shade trees in towns and on farms in and beyond the
native range of the chestnut.
Forest structure and composition changed dramatically with the loss of this major tree in
the forest canopy. Various oaks (Quercus spp.), red maple (Acer rubrum), and hickories
(Carya spp.) assumed dominance in the forest canopy.
By 1940, most large chestnut trees—an estimated 3.5 billion trees—had been killed.
Lumber could be salvaged from dead trees for 10 years after the blight, and bark and
wood was harvested for tannin until the late 1950s. Standing ghost trees became infested
with small boring insects that left pin-sized holes in the wood. This so-called “wormy
chestnut” is still valuable whenever it is discovered; it is used largely for making small
craft items.
IUCN has nominated the chestnut blight fungus as among 100 of the world’s worst
invasive species.
Management. Control of the chestnut blight fungus has followed two routes, one trying to
reduce the virulence of the fungus, and the other trying to increase the resistance of the tree.
A strain of the fungus first identified in Italy was discovered that had been weakened by the
presence of a virus. The spores of these so-called hypovirulent fungi can be inoculated into
cankers on American chestnuts to slow their growth or, for a time at least, have the fungus
produce only the swollen, nonlethal type of canker. A drawback of this procedure is that
hypovirulent fungi spread much more slowly in nature than the virulent strains.
Cross-breeding resistant Asian chestnut species with American chestnut can impart a
degree of resistance to the fungus in the hybrid generation. Several researchers are repeat-
edly back-breeding hybrids with American parents in an attempt to produce a tree with
good forest form, high-quality timber, abundant and good-tasting nuts, and resistance to
chestnut blight.
Some naturally resistant individual American chestnut trees survive both within and out-
side the original range of the tree. Attempts to graft scions from them to existing root stock
are also part of restoration efforts.

Selected References
Anagnostakis, Sandra L. “Revitalization of the Majestic Chestnut: Chestnut Blight Disease.” American
Phytopathological Society, 2000. http://www.apsnet.org/publications/apsnetfeatures/Pages/
ChestnutBlightDisease.aspx.
“Chestnut Blight.” Missouri Botanical Garden, 2001–2010. http://mobot.org/gardeninghelp/plant
finder/IPM.asp?code=283&group=39&level=s.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Cryphonectria parasitica (fungus).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?si=124&fr=1&sts.
Rellou, Julia. “Chestnut Blight Fungus (Cryphonectria parasitica).” Introduced Species Summary
Project, Columbia University, 2002. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/
inv_spp_summ/Cryphonectria_parasitica.htm.
Saucier, Joseph R. “American Chestnut . . . an American Wood. (Castanea dentata (Marsh.) Borkh.).”
FS-230, U.S. Department of Agriculture, Forest Service. Washington, DC: U.S. Government
Printing Office, 1973. Available online at http://www.fpl.fs.fed.us/documnts/usda/amwood/230
chest.pdf.
Treadwell, Judy C. “American Chestnut History.” NCNatural.com, 1996. http://www.appalachian
woods.com/appalachianwoods/history_of_the_american_chestnut.htm.
 18 n FUNGI

n Chytrid Frog Fungus

Also known as: Chytrid fungus, Bd
Scientific name: Batrachochytrium dendrobatidis
Order: Rhizophydiales
Family: Chytridiaceae
Native Range. Unknown. Africa may be the source of this fungus, because the earliest-
known case of infection is on a museum specimen of the African clawed frog (Xenopus laevis)
collected in 1938. The disease caused by this fungal agent, chytridiomycosis, was endemic to
sub-Saharan Africa 23 years prior to its being found elsewhere in the world. However, recent
genetic research suggests that
chytrid frog fungus may have
originated in Japan.
Distribution in the United
States. Throughout.
Description. Chytrid frog
fungus is an invisible external
parasite of amphibians. Positive
identification requires examina-
tion of tissue or water and sedi-
ment samples in a laboratory.
The several symptoms of chytri-
diomycosis are not unique to this
disease but can alert one to the
possibility of infection by Batra-
chochytrium dendrobatidis. These
include behavioral changes in
the infected animal such as leth-
argy, inability to right itself if
turned on its back, failure to flee
from humans, and failure to seek
shelter from the sun. Frogs and
toads may sit in the water all the
time. Redness can appear on the
belly skin, and wet skin may
slough off. The bodies of toads
and frogs can become bloated
from retention of fluids. Toe
curling, head held in a tucked
position, and other signs of mild
paralysis may occur. Sudden
mass die-offs occur in infected
populations. No single symptom
Top: Africa has been suspected as the place of origin for the chytrid frog
applies to all affected species,
fungus, but genetic evidence now points to Japan as the possible source making definitive field identifi-
area. Bottom: The chytrid frog fungus is found throughout the United States. cation impossible.
 CHYTRID FROG FUNGUS n 19

Related or Similar Species. Batrachochytrium dendrobatidis is the only chytrid fungus

known to infect living vertebrates. Other species decompose cellulose and chitin.
Introduction History. It is unknown when or how the chytrid frog fungus entered the
United States. Two hypotheses are being tested. The first poses that the fungus probably
originated in Africa and only began to spread rapidly around the world during the twentieth
century. The African clawed frog, a known host, was exported in large numbers for medical
research beginning in the 1940s and again in the 1960s and 1970s, when its use in human
pregnancy tests became popular. The first documented lethal outbreaks of chytridiomycosis
did not occur until 1998, when reports came simultaneously from Australia and Central
America. The fungus was not scientifically described and named until 1999. Today, it has
been identified on over 350 species of amphibians on all continents except Antarctica.
A second hypothesis states that the chytrid frog fungus has been widely distributed, but
not recognized, around the world for a very long time, and the sudden outbreaks that began
in the late 1990s in disparate locations have been triggered by recent global environmental
changes. The synchronous nature of the outbreaks supports this hypothesis, as does evi-
dence from preserved museum specimens and field analyses that show low-level infections
without lethal consequences on many amphibian species in the eastern United States.
Examination of preserved museum specimens of North American amphibians reveals that
the fungus was in the United States long before it gained the world’s attention in the late
1990s. The earliest record dates to 1974 and a leopard frog (Rana pipiens) collected in the
Rocky Mountains in Colorado. The fungus was also found on a specimen from the Sierra
Nevada in California dating to the same year. (It was found on specimens from Québec,
Canada, collected in the 1960s.)
Whatever its origins, chytrid can be dispersed on live amphibians sold for food, pets, or
research animals. The American bullfrog (Lithobates catesbeianus) could be an effective vector,
since it is a popular food internationally. Poison dart frogs (Dendrobates spp.) from South and
Central America could carry the fungus into zoo collections, where they are prized exhibits.
Contaminated habitat material such as water and sediment can transfer zoospores to new
bodies of water if lodged in tires or on boots or even possibly in the hooves of livestock.
Much research is still needed to unravel the mystery of the distributional history of this
fungus.
Habitat. This is an aquatic fungus, and it requires water as well as suitable hosts. It will
survive in any body of water from streams and lakes to artificial containers. Chytrid frog
fungus prefers cooler temperatures (in laboratory trials, it grows best in water temperatures
of 63–73°F [17–23°C] and dies at temperatures above 82°F [28°C]) and permanent flowing
water. It is more apt to be found in streams than ponds, because the former are cooler and
help to transfer spores over long distances.
Batrachochytrium dendrobatidis lives on the skin of adult frogs and treefrogs, toads, and lung-
less salamanders. It is most prevalent on the more heavily keritanized skin of the abdomen,
especially near the pelvis, on hind limbs, and on feet. It is also found on the oral discs (the
structures surrounding the mouths) of tadpoles. Chytrid frog fungus is found in wild popula-
tions as well as among captive amphibians held in zoos, aquaria, and aquaculture facilities.
Diet. It apparently feeds on keratin, since it is only found on the keratinized tissues of
amphibians. Some research suggests that it may also have non-amphibian hosts or live on
dead tissues as well.
Life History. This fungus has two life stages: an attached spherical zoosporangium that is
the reproductive stage, and a motile zoospore that disperses to new locations on the same host
or to a new host organism. No resting stage has yet been discovered. The zoospore has a single
20 n FUNGI

Chytrid Frog Fungus

T he global decline in amphibian species is an alarming trend with many possible

causes. Among them are habitat destruction, climate change, air pollution,
increased UV radiation, chemical contamination of water, and the introduction of
nonnative competitors and predators. The chytrid frog fungus may be a major
new player. Or it may have been there all along and only now reaches fatal levels
of infection when a frog or toad population is already severely stressed by other
environmental factors.

flagellum with which it moves through water. It attaches to the keratinized outer layers of skin
of its host, absorbs its tail, and burrows below the surface. In four days, it will mature into a
zoosporangium that has root-like rhizoids that both secrete enzymes to break down keratin
and absorb the digested organic products. The zoosporangium forms a single discharge tube
that protrudes out of the host’s skin and through which each zoosporangium will release as
many as 300 zoospores to start the cycle again. Reproduction is asexual or clonal; like many
other chytrid fungi, Batrachochytrium dendrobatidis seems to lack a sexual stage.
Impacts. Chytrid frog fungus has been implicated as one of several causes of declining
amphibian populations around the world. In the United States, regular low-level infections
without deleterious effects seem to have occurred since at least the 1960s. Yet particularly in
western states, severe chytrid infections have occurred as populations of several frog species
have been decreased dramatically. In Arizona, a rapid die-off in 1997 of the lowland leopard
frog (Rana yavapaiensis) and its subsequent extirpation at many of its past locations may have
been due in part to chytridiomycosis. In the Sierra Nevada, die-offs of the mountain yellow-
legged frog (Rana mucosa) correlated with the presence of Batrachochytrium dendrobatidis; but
in and around Pinnacles National Monument, California, few dead frogs were encountered
during surveys in 2006–2007 even though the fungus was discovered infecting Pacific tree
frogs (Pseudacris regilla), western toads (Bufo boreas), California red-legged frogs (Rana aurora
draytonii), and foothill yellow-legged frogs (Rana boylii). Some populations of the Oregon spot-
ted frog (Rana pretiosa), a species in decline in the Pacific Northwest, are persisting even though
heavily infected with the fungus. On the other hand, dead and dying boreal toads (Bufo boreas
boreas) in Rocky Mountain National Park, Colorado, were found to be infected with the chtyrid
fungus, a likely cause of their demise. Die-offs linked to chytrid also are reported in Wyoming
and Washington. Susceptibility to the disease is highly species specific and perhaps site specific.
It may be more virulent at higher elevations with cooler temperatures. Some more tolerant spe-
cies such as the American bullfrog and some salamanders may act as transmitters of the parasite.
It remains to be discovered how chytrid frog fungus kills its host. Various hypotheses
exist, including the possibility that respiration through the frogs’ skin may be affected or that
the fungus produces a toxin. Recent research showed that the skin of infected frogs was less
able to transport sodium and chloride ions and maintain proper balances of sodium and
potassium in the blood. Severe electrolyte imbalances lead to heart stoppage.
Much is yet unknown about the impacts of chytrid frog fungus and just how great a risk it
is to amphibian populations worldwide.
Management. Little can be done to eradicate the fungus once it invades a body of water.
To prevent the spread of this potentially lethal parasite, care should be taken to disinfect all
equipment, footwear, and clothing before entering a stream or pond. Do not move
 DUTCH ELM DISEASE FUNGI n 21

amphibians from one source area to another. Never release captive animals into wetlands,
ponds, lakes, or streams.

Selected References
Daugherty, Matt, and Kim Hung. “Chytrid Fungus, Batrachochytrium dendrobatidis.” Center for Invasive
Species Research, University of California, Riverside, 2009. http://cisr.ucr.edu/chytrid_fungus.html.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Batrachochytrium dendrobatidis (Fungus).” ISSG Global Invasive Species Database, 2006.
http://www.issg.org/database/species/ecology.asp?fr=1&si=123.
Ouellet, M., I. Mikaelian, B. D. Pauli, J. Rodriguez, and D. M. Green. “Historical Evidence of
Widespread Chytrid Infection in North American Amphibian Populations.” Conservation Biology
19(5): 1431–40, 2005.
Padgett-Flohr, G. E. “General Amphibian Disease Information.” California Center for Amphibian
Disease Control, 2002. http://ccadc.us/docs/AmphibianDiseasesPresentation.pdf.

n Dutch Elm Disease Fungi

Scientific name: Ophiostoma novo-ulmi and O. ulmi
Synonym: Ceratocystis ulmi
Class: Sordariomycetes
Order: Ophiostomatales
Family: Ophiostomataceae
Native Range. Uncertain; probably eastern Asia. The disease was first identified and scien-
tifically described in the Netherlands in 1921 and thus became known as Dutch elm disease.
Distribution in the United States. The disease occurs throughout the 48 contiguous
states except for the desert regions of the American Southwest.
Description. Dutch elm disease is caused by one of two closely related fungi. O. ulmi is
becoming less prevalent as it becomes replaced by the more aggressive O. novo-ulmi, believed
responsible for most elm mortality from the 1950s through the 1970s. The fungi grow inside
the xylem of living trees, where white fruiting bodies may be visible if the xylem is exposed
in cross-sectioning; they may be positively identified only in a laboratory.
Outward symptoms of this wilting disease of elm trees develop quickly, usually within a
month, and are obvious. Leaves on a branch of an apparently healthy tree yellow and wilt, a
process known as “flagging.” This is caused by the clogging of xylem tubes by the growing
fungi and prevention of water transport; flagging usually becomes evident in late spring,
when the tree’s leaves have reached full size. They will eventually turn brown and drop pre-
maturely. Wilting proceeds from the tips of branches downward through the crown, unless
the fungus has entered the tree from its roots. In that case, the signs of infection appear first
in the lower crown and quickly envelope the entire crown. Often it takes only one year for
the whole tree to be affected; but sometimes it takes two or more years.
Internal symptoms are the result of dead xylem tissue, revealed by a brown streaking of
infected sapwood. These streaks run with the grain and are evident when the bark is
stripped from a branch. In a cross-sectional cut of the branch, the vascular damage appears
as a ring of brown spots.
Two beetles are the vectors for Dutch elm disease, and management often targets these
species in order to slow the spread of the fungi. The native elm bark beetle (Hylurgopinus
rufipes) is a brownish-black coleopteran sparsely covered with stiff yellow hairs and about
 22 n FUNGI

0.08–0.12 in. (2–3.5 mm) long.

Its wing cases (elytra) are deep-
ly pitted. The larvae are small,
white grubs that produce dis-
tinctive galleries beneath the
bark of elms. Egg-laying gal-
leries of adults run across the
grain, while the smaller tunnels
made by larvae come off the
main gallery parallel to the
grain. H. rufipes burrows into
the bark on branches and
trunks of elm trees, so the infect-
ions it introduces start in major
branches.
The second vector is an
introduced insect, the smaller
European elm bark beetle
(Scolytus multistriatus). This
beetle is slightly larger than the
native elm bark beetle and red-
dish brown. A prominent spine
extends from the underside of
its concave abdomen. It con-
structs galleries in the opposite
manner of the native elm bark
beetle: the egg-laying gallery
runs along the grain of the
wood, while the larval galleries
are cut perpendicular to it
across the grain. Since the
Top: Eastern Asia is the likely place of origin for the fungi that cause smaller European elm bark bee-
Dutch elm disease. Bottom: The distribution of Ophiostoma novo-ulmi, tle feeds in the crotches of small
the chief fungus implicated in Dutch elm disease today. (Adapted from twigs, infections transmitted by
“Pest Distribution Map: Dutch Elm Disease, Ophiostoma novo-ulmi.” it occur first in twigs.
Alien Forest Pest Explorer, USDA Forest Service, Northern Research Related or Similar Diseases.
Station http://www.fs.fed.us/ne/morgantown/4557/AFPE).
Two other diseases of elms may
be mistaken for Dutch elm dis-
ease. Elm yellows (elm phloem necrosis) causes all the leaves of the crown to turn yellow,
usually between July and September. Leaves do not wilt or turn brown. No streaking occurs
in the sapwood, but the inner bark will be discolored and characteristically develops a
wintergreen scent. Bacterial leaf scorch, like Dutch elm disease, infects and clogs the xylem;
however, this disease results in a slow decline of the tree over many years. The older leaves
on a branch are first to show symptoms of stress. In summer and early fall, the margins of the
leaves turn brown, with a yellow area appearing between the green tissue and the scorched
outer edge of the leaf. No visible changes to sapwood or inner bark occur.
Introduction History. Dutch elm disease was first identified in the United States in Ohio
in 1931. O. ulmi had been introduced a few years earlier on diseased elm logs imported from
 DUTCH ELM DISEASE FUNGI n 23

France to be used as veneers in furniture making. The smaller European elm bark beetle had
arrived ahead of it. O. ulmi probably had been introduced to Europe from Asia around 1910.
It is believed responsible for the first epidemic of Dutch elm disease that swept across
eastern North America as well as Europe. The more aggressive O. novo-ulmi was likely intro-
duced to the southern Great Lakes region in the 1940s or 1950s; it probably has caused
most of the widespread loss of elms since then. The disease spread north and south through
the eastern United States and reached the West Coast in 1973. It continues to expand its dis-
tribution area and kill off urban and wild elms. However, many seedlings and saplings
escape infection long enough to reproduce, so elms are not threatened with extinction.
The fungi are spread short distances by elm bark beetles. These insects fly only a few hun-
dred feet away from the tree in which they hatched to feed on another elm. Dispersal dis-
tance may be 2 miles (3 km) or more to breeding sites. The rate of spread seldom exceeds
4–5 miles (6–8 km) a year.
When trees grow close together (within 25–50 ft. [7.5–15 m] of each other) in pure
stands, the usual pattern along city streets, spores from the fungi can be transferred from tree
to tree via the roots. Where roots from neighboring trees cross each other, they fuse to create
a root graft and allow the passage of the Dutch elm disease pathogen. The spores will carried
up the new host tree with the sap in the xylem.
Habitat. Urban forests, natural forests, and landscape trees along streets and in yards.
O. novo-ulmi and O. ulmi infect only elms (Ulmus spp.). American elm (U. americana) is
highly susceptible to the fungus, while other native elm species show a range of resistance;
but none is immune to Dutch elm disease.
Diet. The fungi that cause Dutch elm disease are both parasites feeding on living tissue of
elm trees and saprotrophs living off dead elm tissues. They produce enzymes that digest
plant cell walls and perhaps toxins that kill the parenchyma cells of xylem. It is the death
of xylem parenchyma that causes the diagnostic discoloration of the sapwood.
Life History. Dutch elm disease fungi are introduced into an elm tree when elm bark
beetles contaminated with spores bore into the bark to feed or to create egg-laying tunnels.
The spores become dislodged and germinate in the galleries to produce thread-like
cells, the hyphae, that penetrate into the xylem, where they grow and form the fruiting
bodies of the fungus. The fruiting bodies produce millions of white oval spores. These move
through the xylem, where they reproduce asexually by budding and spread the disease
through the tree. In dying and recently dead trees, a different type of asexual fruiting struc-
ture is produced asexually in the bark and in the galleries just beneath the bark: Round,
sticky white spores are attached to the top of dark stalks less than 0.1 in. (1–2 mm) tall.
The spores adhere to adult beetles and are carried to new elms, when the beetles exit the
tree. Sexually produced spores also occur. These accumulate in sticky droplets that can also
attach to elm bark beetles to be carried to new host trees. Newly emerged adult beetles first
feed on healthy elms. Later, they move to sick or dead elms to breed. Beetles usually have
two broods a year, the second of which overwinters and emerges in early summer. The sec-
ond brood is responsible for the majority of new infections.
Impacts. Dutch elm disease has destroyed many millions of elm trees in the United States.
The stately, vase-shaped American elm once lined the streets of towns and cities in the
eastern part of the country, their arching canopies providing welcome summer shade. The
elm was favored as a landscape and urban forest tree because it was beautiful, long-lived,
and tolerant of poor air quality and compacted soils. The urban landscape is totally altered
with the demise of these trees. The American elm was most susceptible to the disease, but
other species, including those in natural forests, are also vulnerable to varying degrees.
 24 n FUNGI

A. Smaller European elm beetle, the primary vector for this pathogen. (Gerald J. Lenhard, Louisiana State
Univeristy, Bugwood.org.) B. Fruiting bodies of the Dutch elm disease fungus. (William Jacobi, Colorado State
University, Bugwood.org.) C. American elms lined many city streets in days past. The vase-like structure of this
native species was one of its desirable characteristics. (Joseph O’Brien, USDA Forest Service, Bugwood.org.)
D. Galleries made by the smaller European elm bark beetle. (John A. Williams, USDA Forest Service, Bugwood.org.)

Trees are more resistant under drought conditions, and slower-growing individuals are less
susceptible than vigorously growing ones.
Management. Reducing losses of urban trees from Dutch elm disease requires a
community-wide campaign to remove and destroy diseased branches and trees. Root grafts
also must be destroyed. Application of systemic chemicals can prevent and treat infections
in individual specimen trees, but must be repeated every 2–3 years. Insecticides may control
beetle populations, but they must be synchronized with the beetles’ life cycles and can be
expensive, and there are potential hazards in handling these products.
American elms continue to be planted because of their desirable properties. If arborists
avoid large plantings of the same species and space elms well apart from other elms, infec-
tion rates and tree losses may be reduced significantly.
 SUDDEN OAK DEATH n 25

The best long-term solution is the breeding of resistant strains of the elm. Hybridization
between American elms and Asian species resistant to or tolerant of the fungus provided
resistance but failed to produce the elegant structure of the American tree. Scientists with
the Agricultural Research Service and the U.S. National Arboretum discovered and nurtured
some old surviving American elms with the necessary resistance and through careful breed-
ing produced varieties that preserved the tall vase shape of the original species; some are
now available commercially. They include such cultivars as American Liberty, Princeton,
Independence, Valley Forge, New Harmony, and, most recently, Jefferson elms.

Selected References
D’Arcy, C. J. “Dutch Elm Disease.” The Plant Health Instructor. American Phytopathological Society,
2000. Revised, 2005. http://www.apsnet.org/edcenter/intropp/lessons/fungi/ascomycetes/Pages/
DutchElm.aspx. doi:10.1094/PHI-I-2000-0721-02.
“Fact Sheet: Dutch Elm Disease (Ophiostoma novo-ulmi).” Cornell University, Plant Disease Diagnostic
Clinic, 2009. http://plantclinic.cornell.edu/FactSheets/dutchelmdisease/DED.htm.
Haugen, Linda. “How to Identify and Manage Dutch Elm Disease.” Northeastern Area State and Private
Forestry, USDA Forest Service, n.d. http://www.na.fs.fed.us/spfo/pubs/howtos/ht_ded/ht_ded.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Ophiostoma ulmi sensu lato (fungus).” ISSG Global Invasive Species Database, 2010. http://
www.issg.org/database/species/ecology.asp?fr=1&si=130.

n Sudden Oak Death

Also known as: Ramorum blight, SOD
Scientific name: Phytophthora ramorum
Class: Oomycetes
Order: Pythiales
Family: Pythiaceae
Native Range. Unknown. Two mating types exist, A1 in Europe and A2 in North
America. It is presumed that they have a common ancestor, perhaps from Asia. On both con-
tinents, the pathogen acts like a new or emerging disease, supporting the idea that it origi-
nated somewhere else. (Ramorum blight was first recorded in Europe, on ornamental
rhododendrons in Germany in 1993.)
Distribution in the United States. Sudden oak death occurs in 14 counties of coastal
California stretching 300 mi. (480 km) from Monterey County north to Humboldt
County. It is also in neighboring Curry County in southwestern Oregon.
Description. The fungus is best described according to the symptoms shown on host plants,
which manifest themselves in two groups according to the response to the parasite: bark canker
hosts and foliar hosts. Tanoaks (Lithocarpus densiflorus) fall into both categories. On the first
group, which includes coast live oak (Quercus agrifolia), California black oak (Q. kelloggii),
Shreve oak (Q. parvula var. shrevei), and madrone (Arbutus menziesii), the twigs show the first
signs of infection, with discolored patches of dead tissue beneath the bark separated from
healthy tissue by a black line or reaction zone. This zone is the advancing front of infection.
As the fungus grows in the twig, it spreads into larger branches and causes cankers that lead
to branch die-off. With continuing expansion, the infection reaches the stem, where cankers
may become more than 6 ft. (2 m) long and girdle the cambium, quickly killing the tree.
Characteristically, a red or black sap-like fluid oozes from the canker, staining the bark and
 26 n FUNGI

killing any lichen or mosses

growing on the trunk. The
canker attracts western oak
bark beetles (Pseudopityphthorus
pubipennis) and ambrosia bee-
tles (Monarthtum dentiger and
M. scutellare) in abnormally large
numbers. The abundance of
such beetles is one of the distinct
signs of sudden oak death. The
dead wood of the canker hosts a
saprophytic fungus, Hypoxylon
thouarsianum; its fruiting bodies
are flattened khaki-green domes
that turn black with age and are
another indication of infection
by sudden oak disease. On small
tanoaks, which are both bark
canker hosts and foliar hosts,
the first signs of the disease are
wilting of the branch tips. When
the branch dies, the response of
the tree is to resprout with multi-
ple new shoots. Tanoaks often
display several cankers at differ-
ent heights above the ground.
As with all bark canker hosts,
within a few weeks of girdling
of the trunk, the crown suddenly
browns. Death of evergreen trees
such as live oaks may take one or
Top: The place of origin of the fungus responsible for sudden oak death two years after infection, with the
remains a mystery, although somewhere in Asia is likely. Bottom: Areas brown leaves remaining on the
known to be affected by Phytophthora ramorum in 2009. (Adapted from tree during that period. One does
map by M. Kelly, University of California, Berkeley. http://www.sudden not find single isolated trees
oakdeath.org.) infected by the disease; instead,
many trees in close proximity
will be killed. Another field observation has been that oak and tanoak trees with bark cankers
are always growing near infected bay laurels, which are foliar hosts.
Foliar hosts such as coastal redwood (Sequoia sempervirens), Douglas fir (Pseudotsuga men-
ziesii), bay laurel (Umbellularia californica), rhododendrons, and huckleberries are not killed
by the fungus; and their infections are often referred to as ramorum blight. The main symp-
toms of the disease are brown or black spots on green leaves, browning leaves, and some-
times twig dieback.
Despite these several signs, sudden oak death cannot be reliably determined in the field,
but requires sophisticated laboratory analyses.
Related or Similar Species. The symptoms of several ailments may resemble those of
sudden oak death including canker rots, leaf scorch, freeze damage, and herbicide damage.
 SUDDEN OAK DEATH n 27

In the eastern United States, where sudden oak death is not as yet present outside the occa-
sional infections in nurseries, oak wilt presents without cankers or bleeding bark; it is
caused by a different fungus, Ceratocystis fagacearum. Oak decline, another disease of eastern
oaks, kills many trees, but slowly. It is believed to be brought on by the interactions of multi-
ple stresses such as drought, infections of root fungi such as Armillaria mellea, and infesta-
tions of inner bark borers such as two-lined chestnut borer (Agrilus bilineatus) and red oak
borer (Enaphalodes rufulus). The red oak borer by itself causes dark, wet stains, but stains
on the inner bark show no black zones and are accompanied by beetle burrows filled with
fine frass. All other Phytophthora species in temperate areas infect roots and root crowns
and spread as soilborne or waterborne spores.
Introduction History. The sudden death of large swaths of tanoaks was initially reported in
1995 in Marin and Santa Cruz counties, California. The cause was not identified until 2000,
when University of California scientists determined it was as a previously unknown species
of the fungus Phytophthora. Later that same year, a researcher in the United Kingdom deter-
mined that it was the same fungus that had been affecting rhododendrons in Germany and
the Netherlands since 1993. On both continents, the origins of the infections are unknown.
In 2001, P. ramorum was discovered on ornamental rhododendrons in a Santa Cruz nurs-
ery, and Oregon reported a 40-acre outbreak in Curry County. By 2002, the presence of the
fungus was confirmed in natural forests in 10 counties in California. Today, it occurs in 14.
In subsequent years, P. ramorum was detected in other nurseries in California and Oregon,
and many other states established outright bans on the import of plants from California, or
temporary quarantines, inspection, and treatment of imported nursery stock. A major scare
occurred in 2004 when infected plants from the huge Monrovia nursery showed up in
Colorado, Georgia, Louisiana, Maryland, North Carolina, Texas, and Virginia, as well as
Canada. P. ramorum continues to turn up in nurseries across the country. (It has also spread
across Europe.)
Habitat. The disease is most prevalent in the mild, moist habitats exemplified by the fog-
swept coasts of California. Temperatures of 64–68°F (18–20°C) and humidity close to
100 percent increase the chances of infection. The fungus that causes sudden oak death is
common in the understory of coastal redwood forests on tanoak, in Douglas fir–tanoak for-
ests with understories of huckleberry or other Vaccinium species, and in coastal broadleaf
forests on coast live oak, California black oak, Shreve oak, madrone, and bay laurel among
at least 100 species known to be susceptible to bark canker or—much more frequently—leaf
infections. It lives primarily in the phloem tissues of trees, shrubs, and perennial herbs in
natural forests, parklands, and residential areas.
Life History. This fungus shows little genetic variability throughout its range in the
United States, where it exists as a clone that reproduces only asexually. (Both mating types,
A1 and A2, are needed for sexual reproduction.) The fungus forms sporangia, the sacs that
produce zoospores, mostly during California’s rainy season between December and June.
The sporangia break off and may be carried by the wind some 15 ft. (5 m) away from the
original infection site. Sporangia apparently form only in infected leaves, and not on the
bark of oaks or tanoak. Zoospores are released when the sporangia land on a new host; they
are motile and can swim in water or a water film on the leaves and bark. The cankers on sus-
ceptible oaks are usually either close to the soil line, suggesting that rain splash may be a
means of dispersal; or adjacent to the leaves of infected bay laurels, suggesting airborne dis-
persal. No wound in the bark appears necessary to allow infection of those plants that are
bark canker hosts, nor can they reinfect themselves since the sporangia develop on fungi
infecting leaves.
 28 n FUNGI

A. Canker with zone lines caused by infection by Phytophthora ramorum. B. Canker bleeding. C. Oaks defoliated
as a consequence of sudden oak death. (Joseph O’Brien, USDA Forest Service, Bugwood.org.)

Spores can be found in streams all year, so sporulation must occur during the dry season.
P. ramorum also forms larger, thick-walled resting spores (chlamydospores) that survive
through unfavorable environmental conditions. Much remains to be learned about the life
history of P. ramorum, but in some other Phytophthora fungi, the resting spores can remain
dormant up to six years.
Impacts. In infested areas, high mortality is experienced by tanoaks; and major losses are
incurred by coast live oak, Shreve oak, and California black oak. These trees may be domi-
nants in the canopies of the forests in which they occur and many times make up pure
stands. When these trees die, the forest floor is no longer shaded, and soils become less per-
meable to water, changing hydrologic conditions. Gone too are food (acorns) and shelter for
wildlife. The dead trees and litter increase the fuel available for wildfires and may change the
 WHITE PINE BLISTER RUST n 29

fire regime. Possible changes in species composition in affected forests are yet to be
determined.
Foliar hosts, although not killed, serve as reservoirs for the disease and are necessary
components in the spread of the fungus. This is particularly a problem in ornamental rhodo-
dendrons and camellias in California’s nursery trade, which sells plants to other states and
countries, and also for companies exporting Christmas trees and redwood mulch. The eco-
nomic toll of import bans, quarantines, and inspections could be significant.
Experiments have shown that northern red oak (Quercus rubra), northern pin oak (Q. pal-
ustris), and mountain laurel (Kalmia latifolia) are highly susceptible to sudden oak disease.
Many Midwestern and southern forests would be threatened were the fungus to become
established in them. Most vulnerable may be forests in the Ozark-Ouachita highlands, live
oak stands in Florida, and pin oak sand flats in the Great Lakes area.
Management. No way exists to control the disease in the wild. The main management
strategy is to prevent its spread via the nursery trade. Since the resting spores survive in soil
and leaf litter, it is also important to clean the tires of vehicles and boots of hikers that have
been in affected areas and to prohibit the transport of plant debris, including bark, wood,
and mulch, from infected areas.
In nurseries and individual landscape plants, the infected plant is either treated with a
special fungicide or cut down and burned.

Selected References
Alexander, J. M., and S. V. Swain, Pest Notes: Sudden Oak Death UC ANR Publication 74151, UC
Statewide IPM Program, University of California, Davis, Ca, 2010. http://www.ipm.ucdavis.edu/
PMG/PESTNOTES/pn74151.html.
Garbelotto, Matteo. “Sudden Oak Death: A Tale of Two Continents.” Outlooks on Pest Management,
85–89. Research Information Ltd., 2004. http://www.ufei.org/ForesTree/files/collected/Pesticide
Outlook.pdf. doi:10.1564/15apl12.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Phytophthora ramorum (fungus).” ISSG Global Invasive Species Database, 2008. http://
www.issg.org/database/species/ecology.asp?si=563&fr=1&sts=sss.
O’Brien, Joseph G., Manfred E. Mielke, Steve Oak, and Bruce Moltzan. “Sudden Oak Death” Pest Alert.
U.S. Department of Agriculture, Forest Service, State and Private Forestry, Northeastern Area,
2002. http://na.fs.fed.us/spfo/pubs/pest_al/sodeast/sodeast.htm.

n White Pine Blister Rust

Scientific name: Cronartium ribicola
Division: Basidiomycota
Class: Teliomycetes
Order: Uredinales
Family: Cronartiaceae
Native Range. Asia. The natural range of this fungus is unknown but presumed to be
Asia, since the disease was unknown in Europe prior to the introduction of eastern white
pine (Pinus strobus) from North America in the 1600s. White pine blister rust was first dis-
covered in Europe in the Baltic provinces of Imperial Russia in 1854. From there, it spread
westward across Europe. It is supposed that the rust infected native pines in Asia without
significant symptoms but when a naı̈ve nonnative species became widely planted, it found
a new host and gained virulence.
 30 n FUNGI

Distribution in the United

States. This parasite occurs
throughout the range of white
(five-needled) pines in the
eastern United States, the Great
Lakes area, and the mountains
of the northwest. It can be
found in at least 33 states and
continues to expand its range
in the Southwest and in sou-
thern California.
Description. The white pine
blister rust fungus requires two
hosts, white pines and goose-
berries or currants (Ribes spp.;
but see below under Habitat)
to complete its life cycle; the
most visible signs of its pres-
ence appear on the pines. Early
symptoms of infection occur in
late summer or fall as small yel-
low spots on live pine needles.
The fungus spreads down the
needle into the vascular system
of the twig and, during the next
growing season, produces a
slight swelling and yellowing of
the bark as a diamond-shaped
canker forms on the branch.
Within a year or two of infec-
tion, yellow blisters (aecia) up
Top: The place of origin of white pine blister rust is unknown but to 0.25 in. (3 mm) wide erupt
presumed to be in the pine forests of Asia. Bottom: White pine blister through the bark in early
rust affects both western and eastern pines in the lower 48 states. spring. When the blisters rup-
(Adapted from map by the Global Invasive Species Team [GIST], the ture, they release vast quantities
Nature Conservancy. http://www.invasive.org/gist/photos/crori03.gif.) of yellow-orange spores (aecio-
spores) and then dry up, leav-
ing a rough patch of bark on the branch. In late spring and early summer, another form of
blister (pycnia) appears on the canker. It oozes a sticky yellow-orange fluid that contains
pycniospores. Both the blisters and the fluid harden and blacken, and remain on the tree
for several weeks. The blisters form again in following years around the margin of the can-
ker, creating a distinctive orange perimeter and expanding the canker until the stem is
invaded and girdled and the tree above the canker killed. Rust-infected bark is high in sugar
content and attracts rodents; their gnawing at the bark produces copious amounts of pitch
(resin) that run down the tree trunk and is another sign of rust infection.
The progression of the disease is visible from a distance: chloritic (yellowed) needles,
stunted or dead branches (flagging), and eventually dead crowns or dead trees. On goose-
berries and currants, the symptoms of rust infection are less obvious. The lower surfaces
 WHITE PINE BLISTER RUST n 31

of leaves lose their green color and within a few days become covered with minute orange
fruiting bodies (uredinia); these tiny bumps produce yellow-orange spores. In late summer,
a different type of fruiting body (telium) covers the underside of the leaves with short
yellow-brown hair-like structures.
Related or Similar Species. This fungus is the only stem rust of white pines in North
America, and it affects only members of the white pine group. Similar symptoms may appear
on lodgepole (Pinus contorta) and ponderosa (P. ponderosa) pines, but these are caused by the
western gall rust (Endocronartium harknessii). Pine bark adelgids (Pineus strobi), an aphid-like
insect introduced from Europe, cover themselves in white, woolly, and waxy secretions;
heavy infestations could be mistaken for the pitch-dripping cankers on white pines infected
with white pine blister rust. So could Armillaria root rot disease, caused by the fungus
Armillaria mellea. When the lower trunks of conifers host this fungus, large amounts of resin
are exuded.
Introduction History. White pine blister rust first entered the eastern United States on
eastern white pine seedlings imported from Germany sometime between 1898 and 1908.
The U.S. Forest Service had sent seeds to France and Germany, because American nurseries
could not meet the demand when replanting cutovers became the vogue, and then encour-
aged the planting of millions of imported seedlings across the northeastern states from
New England to Minnesota. The blister rust invaded the northwestern United States after
having been introduced accidentally in 1910 to Vancouver, British Columbia, Canada, on
eastern white pine seedlings from France; however, it was not diagnosed until 1921.
During this time period, Americans were beginning to understand that they could no longer
cut down forests and move ever westward, but had to practice forest management and make
efforts to reforest cutover areas. Europeans had faced these problems much earlier and
become accomplished nurserymen, producing seedlings of American white pine as well as
European species. Europe thus became a valuable source of seedlings for American foresters.
The first outbreak of the disease occurred in 1906 in Geneva, New York, where it was dis-
covered on currants. In 1909, it was found at several eastern locations on eastern white pine
seedlings from European nurseries and, by 1915, on native seedlings. By 1922, some areas
of New Hampshire saw 50 percent of white pines infected; the rust quickly spread through
the forests of the Appalachians and into the Great Lakes region, infecting most major pine
regions by 1950. In the west, the blister rust had spread east in western white pine as far
as Idaho as early as 1923, and south to Oregon—infecting sugar pine—by 1929. The disease
was observed in the northern Sierra Nevada in 1941 and had reached the southern Sierra by
1961. In 1970, it was discovered on southwestern white pine in New Mexico and limber
pine in southern Wyoming. It showed up in northern Colorado on limber pine in 1998.
The most recently documented range expansion is into central Colorado in 2003, where
for the first time it was observed on Rocky Mountain bristlecone pine. It continues to spread
through the Great Basin, the southern Rocky Mountains, and the Southwest.
Habitat. White pine blister rust infects pine trees that have five needles per bunch. It
attacks trees in natural forests as well as those grown in plantations or as ornamentals.
Cool, moist weather in summer and early fall is necessary for the infection of the pine host,
so it tends to be restricted to more northerly latitudes or high elevations. In the eastern
United States and the Great Lakes area, eastern white pine (Pinus strobus) is the sole host.
In western states, western white pine (P. monticola), sugar pine (P. lambertiana), whitebark
pine (P. albicaulis), limber pine (P. flexilis), and southwestern white pine (P. strobiformis)
are hosts. In 2002, the rust was detected on Rocky Mountain bristlecone pine (P. aristata)
for the first time.
 32 n FUNGI

All currants and gooseberries, both wild species and cultivars, are susceptible to some
degree to infection as the alternate hosts required by the rust. Most susceptible is the domes-
ticated European black currant (Ribes nigrum). Some red-currant varieties are highly resist-
ant, perhaps immune to the disease. Recently, two western members of the broomrape
family (Orobanchaceae), sickletop lousewort (Pedicularis racemosa) and giant red Indian
paintbrush (Castilleja miniata), have also been found to serve as alternate hosts.
Life History. The white pine blister rust has what is known as a macrocyclic, heter-
oecious life cycle. It is extremely complex and involves five spore types and requires two
alternate host species. Germ tubes from basidiospores released from the life stage of the rust

A. Earlier stage of infection with oozing blisters. (John W. Schwandt, USDA Forest Service, Bugwood.org.)
B. Fruiting bodies of Cronartium ribicola. (USDA Forest Service Archive, USDA Forest Service, Bugwood.org.)
C. Well-developed canker on eastern white pine. (Minnesota Department of Natural Resources Archive,
Minnesota Department of Natural Resources, Bugwood.org.) D. Uredinia on underside of currant leaf. (Petr
Kapitola, State Phytosanitary Administration, Bugwood.org.) E. Dead crowns are among the more conspicuous
symptoms of white pine blister rust. (Joseph O’Brien, USDA Forest Service, Bugwood.org.)
 WHITE PINE BLISTER RUST n 33

that lives on a member of the genus Ribes enter the stomata (pores through which a leaf
exchanges gases with the atmosphere) of a white pine needle. They grow down the length
of the needle and establish mycelia in the vascular tissue of the twig to which the needle is
attached, parasitizing living cells. Blister rust cankers expand with time to the branch and
eventually the main stem (trunk) of the tree. A year or two after infection, pycnia develop
at the margin of the canker and produce pycniospores. Pycniospores result in mycelia that
produce aecia (a cuplike fruiting body) the next year. Aecia break through the bark, covered
by white membranes that create blisters. When the membrane ruptures, quantities of orange
aeciospores are released. Pines cannot directly infect other pines; the aeciospores must find
an alternate host for the next phase of the rust’s life cycle. The wind can carry them over hun-
dreds of miles, and if conditions are optimal, they will survive for several months until they
land on and infect a wild or cultivated currant or gooseberry. Mycelia grow in the Ribes leaf,
and within a few weeks of the initial infection, the rust produces tiny pustules (uredinia) on
the underside of the Ribes leaf; from these fruiting bodies urediospores are released, spreading
the rust to other currants or gooseberries nearby. These spores cannot infect pines. The rust
has entered what is known as the repeating stage, when reinfection of the same alternate host
species occurs over and over. In late summer, after two weeks of cool, moist weather, hairlike
telial fruiting bodies emerge from the underside of the Ribes leaf. These release basidiospores
that are wind-dispersed a distance usually less than 1,000 ft. (300 m) and that, if successful,
land on a white pine. Their germ tubes enter the needles’ stomata to begin the cycle again.
The rust overwinters in the pine; it dies in Ribes or outside a living host.
The disease progresses from needle to twig, to branch, to the trunk of the tree as the mycelia
grow through the vascular system and absorb water and nutrients from the living cells. Cells of
the host pine die as they become undernourished or the flow of nutrients is blocked. Once a
branch or trunk is girdled by the canker, the tissues beyond the infection point are cut off from
water and nutrients, and they die. It may take 15 years for a tree to be killed. Ribes shrubs, on
the other hand, are not killed by the fungus, but severe infections may lead to defoliation.
Impacts. In the early twentieth century, when the white pine blister rust was introduced,
American commercial forestry revolved around the harvest of white pine. When the eastern
white pine was logged out of New England and other northeastern states, the industry had
shifted westward to the Great Lakes region. By 1900 it was beginning to cut western white
pine and sugar pine from the northern Rocky Mountains to the Cascades and Pacific
Northwest. Eastern white pine was being replanted in the eastern United States, often with
inexpensive seedlings from France and Germany. The rust is especially lethal to young trees
of sapling and pole size. Efforts to protect these valuable timberlands began almost immedi-
ately and influenced the impacts of the fungus on native forests.
White pine blister rust has caused more damage and had more money spent on its con-
trol than any other disease of conifers in the United States. Thousands of stands became
unsuited for lumber production or were lost entirely. The initial invasion in the Northeast
resulted in high rates of infection and mortality but also selected for existing resistance in
eastern white pine. In New Hampshire in the first half of the twentieth century, for example,
disease incidence ranged from 20 to 80 percent in sampled stands. By the end of the cen-
tury, the rate of infection was 7 percent or less. The maturing of surviving white pine prob-
ably affects the rate, since young trees are more susceptible to infection. Today, the rust is
widespread but not frequent, and not a major concern. Eradication of wild and cultivated
Ribes as a primary management strategy (see under Management), probably shifted the rela-
tive abundance of wild currants in natural forests, although increased forest cover also
reduces the abundance of these sun-loving shrubs.
34 n FUNGI

Sugar pine and western white pine are valuable timber trees that have been severely
affected by the rust in the western United States. The harvest of western white pines acceler-
ated when white pine blister rust was discovered in the Pacific Northwest. White pines were
selectively removed in salvage operations, leaving shade-tolerant conifers such as western
hemlock (Tsuga heterophylla) and grand fir (Abies grandis) to become dominants in the forest
canopy. Regrowth of white pines was impeded not only by the rust, but by fire suppression
policies, which eliminated the burns pines often require for germination.
High-elevation white pines in the west, especially whitebark pine and limber pine, provide
food and cover for a variety of wildlife and plant species. Several produce large seeds that are
important elements of the diets of Clark’s Nutcrackers (Nucifraga columbiana), and other birds,
as well as grizzly bear (Ursus arctos) and black bear (U. americanus). Squirrels could be nega-
tively affected if these forests decline, as would their predators. The demise of white pines that
occupy steep, dry slopes may turn these sites into treeless areas. The loss of trees will change
slope stability and hydrologic patterns and also alter fire and other disturbance regimes and
forest succession. These trees also have scenic value and contribute to the attractiveness of sev-
eral national parks as well as the tourist economy of the region. Stands of dead whitebark pine
occur in Crater Lake, North Cascades, and Glacier national parks. Some scientists believe that
all whitebark pines in Glacier National Park may be dead by 2015.
White pine blister rust is an important disease in terms of the history of forest pathology
and became one of the most famous tree diseases in the world. Since it struck valuable tim-
ber trees, and since it was discovered early on that something could be done to control its
spread (unlike the Chestnut blight fungus), it fostered new pest management agencies and
techniques including regulations against importing exotic species and establishment of
quarantine zones. The devastation wrought by white pine blister rust and chestnut blight
in the first decade of the twentieth century led to the passage of the Plant Quarantine Act
by the U.S. Congress in 1912. It was the first law aimed at preventing the introduction of for-
eign pests into the United States.
Management. The earliest strategies to control the spread of white pine blister rust
focused on the eradication of Ribes as an alternate host. This was somewhat successful in
the eastern United States, where cultivated currants and gooseberries were the chief hosts,
since only a few wild species in the genus occurred there. Planting, possessing, or importing
Ribes is still forbidden in some counties that remain in quarantine districts, although such
laws are rarely enforced. In the West, where many native species occur in the wild, the strat-
egy was totally impractical and soon abandoned.
Site management for new plantings grew in importance and included the hazard rating of
areas according to conditions that support basidiospore production. High-hazard sites usu-
ally are low areas where moist, cold air settles at night. Small openings in a forest also have
high potential for basidiospore production, because dew often persists on pine seedlings
in such areas. A thin overstory can be maintained to prevent dew formation and still allow
sufficient sunlight to reach pine seedlings. Pruning the lower branches of saplings up to
10 ft. (3 m) above the ground keeps fungal infections from extending into the trunks of
young trees, and it has the added commercial advantage of providing knot-free timber.
In natural forests, the greatest hope lies in the development of resistant trees. Natural
selection for such trees seems to have occurred in the Northeast, where the rust entered after
mature trees had been removed and many young trees were planted. Breeding programs
based on some naturally resistant individuals are progressing throughout the country.
Seedlings of resistant western white pine are available to the public; for other species, such
as eastern white pine and sugar pine, programs are still in experimental stages.
 WHITE PINE BLISTER RUST n 35

Selected References
Cox, Sam. “White Pine Blister Rust: The Story of White Pine, American Revolution, Lumberjacks, and
Grizzly Bears.” 2003. http://landscapeimagery.com/wphistory.html.
Geils, B. W. “Impacts of White Pine Blister Rust.” In Proceedings, U.S. Department of Agriculture inter-
agency research forum on gypsy moth and other invasive species 2001, ed. S. L. C. Fosbroke and
K. W. Gottschalk. January 16–19, 2001, Annapolis, MD. Gen Tech. Report NE-285. Newton
Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station:
61–64, 2001. Available online at http://www.rmrs.nau.edu/rust/Geils2001/text.html.
Laskowski, Michele. “White Pine Blister Rust.” High Elevation White Pines. http://www.fs.fed.us/rm/
highelevationwhitepines/Threats/blister-rust-threat.htm.
Maloy, O. C. “White Pine Blister Rust.” In The Plant Health Instructor. American Phytopathological
Society, 2003. Updated, 2008. http://www.apsnet.org/education/LessonsPlantPath/WhitePine/
symptom.htm. doi:10.1094/PHI-I-2003-0908-01.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Cronartium ribicola (Fungus).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=550&fr=1&sts=sss.
Nicholls, Thomas H., and Robert L. Anderson. “How to Identify White Pine Blister Rust and Remove
Cankers.” North Central Research Station, Forest Service, U.S. Department of Agriculture, 1977.
http://www.na.fs.fed.us/spfo/pubs/howtos/ht_wpblister/toc.htm.
“White Pine Blister Rust Factsheet.” Plant Disease Diagnostic Clinic, Cornell University, 2005. http://
plantclinic.cornell.edu/FactSheets/wpineblister/wpineblister.htm.
Worrall, James J. “White Pine Blister Rust.” Forestpathology.com, 2009. http://www.forestpathology
.org/dis_wpbr.html.
n Invertebrates
n Bryozoan
n Lacy Crust Bryozoan
Also known as: Sea moss, white lace bryozoan
Scientific name: Membranipora membranacea
Family: Membraniporidae
Native Range. Lacy crust bryozoans are native to the temperate, coastal waters of Europe,
where they are found in the northeast Atlantic Ocean, Baltic Sea, and Mediterranean Sea.
They also occur naturally along the Pacific Coast of North America from Alaska to Baja
California.
Distribution in the United States. The lacy crust bryozoan is a nonnative invasive in the
coastal bays of New England; it is established in Connecticut, Maine, Massachusetts, New
Hampshire, and Rhode Island.
Description. This epiphytic bryozoan forms white, circular colonies that are about 0.04 in.
(1 mm) thick and 0.5 in. (1 cm) in diameter. The colonies grow outward, with the oldest
member at the center and younger individuals, or zooids, radiating in rows away from it to
form lacy or net-like encrustations on seaweeds. Each zooid, about the size of a pin head, is
rectangular and box-like and has short knob-like spines at each corner that make the colony
feel like sandpaper. The side walls of the zooid are thin and only slightly calcified; the top is
transparent, allowing light to reach the seaweed that it encrusts.
Related or Similar Species. Native bryozoans of the Northwest Atlantic do not have rec-
tangular zooids. Sea mat or horn wrack (Electra pilosa), the encrusting bryozoan with which
lacy crust bryozoans might most easily be confused, has egg-shaped zooids surround by
spines (4–12; usually 9); a large central spine protrudes from the zooid, giving E. pilosa a
spiny appearance. Colonies of sea mat form snowflake- or star-shaped colonies.
Introduction History. This bryozoan first appeared on sugar kelp (Laminaria saccharina)
in the Gulf of Maine at the Isles of Shoals in 1987. It likely arrived in a ship’s ballast water
or as a fouling organism on the hull of a ship. It spread north and south from its point of
introduction and, by 1990, was the dominant epiphyte on kelp off the shores of Maine
and New Hampshire. It has since reached Cape Breton, Nova Scotia.
Habitat. Lacy crust bryozoan colonies encrust kelps and other macroalgae in the shallow
subtidal zone of temperate seas. They are most common on Laminaria kelps. These bryozo-
ans will also attach to other smooth, hard surfaces such as rocks, glass, and floats. They
flourish where there is fast-flowing water or a high tidal exchange.
Diet. These filter-feeders sieve phytoplankton from seawater with a ring of tentacles
called a lophophore.
Life History. After the initial zooid becomes attached to a kelp frond, new zooids bud off
(asexual reproduction); the colony grows several millimeters a day beginning in late spring.
Growth continues throughout the summer, and by fall, large crusts may be apparent as
 LACY CRUST BRYOZOAN n 37

several colonies merge together.

Unlike many other bryozoans,
in which different types of
zooids perform different func-
tions for the colony, only one
type of zooid occurs in the lacy
crust, and it is involved in feed-
ing, reproduction, and defense.
Sexual reproduction usually
occurs in spring and summer,
when zooids produce eggs and
release them into the water.
The individual animals may be
hermaphroditic in that eggs are
fertilized before they are shed.
The free-floating eggs quickly
develop into tiny triangular lar-
vae (cyphonautes) that become
part of the plankton. They settle
when they contact an algal
frond and grow toward its base,
the most stable part of the sea-
weed. Different colonies com-
pete for space and interact
both aggressively and coopera-
tively, and they can communi-
cate with each other by means
of electrical signals.
Sea slugs (nudibranchs)
such as Onchidorus muricata are
the primary predators of lacy
crust bryozoans. Top: The lacy crust bryozoan is native to the coastal waters of Europe and
Impacts. Lacy crust bryo- the Pacific coast of North America. Bottom: The lacy crust bryozoan has
zoan colonies weigh down the invaded coastal waters off New England. (Both maps adapted from
kelp fronds that they heavily “Membranipora membranacea,” USGS 2009.)
encrust and make the fronds
especially susceptible to breakage during storms. This has caused a reduction and, in some
cases, total loss of kelp beds in the waters off New England. They apparently also exclude
native encrusting animals from kelp and other suitable attachment substrates and interfere
with the host kelp’s spore production. Kelp beds are important habitat and nursery areas
for a variety of marine organisms, including sea urchins, lobsters, and finfish, all of which
are therefore threatened by the invasion of this bryozoan. The green or northern sea urchin
(Strongylocentrotus droebachiensis) in particular has declined in numbers since the introduc-
tion of lacy crust. Where kelp beds have been denuded in the Gulf of Maine, a nonnative
green algae called oyster thief or dead man’s fingers (Codium fragile tomentosoides) has
invaded and prevented the restoration of kelp beds.
Management. There is no effective way to prevent the spread of this organism.
 38 n INVERTEBRATES (BRYOZOAN)

A. Colony of lacy crust bryozoans, showing the box-like zooecia. B. Close-up of zooecia with knobby spines
clearly visible at each corner. C. Rings of tentacles or lophophores are extended when the zooecia are filtering
food particles from sea water. (Dave Cowles: http://rosario.wallawalla.edu/inverts.)

Selected References
Cowles, Dave, and Jonathan Cowles, “Membranipora membranacea (Linnaeus, 1767),” 2007. http://
www.wallawalla.edu/academics/departments/biology/rosario/inverts/Bryozoa/
Class_Gymnolaemata/Order_Cheilostomata/Membranipora_membranacea.html.
“Lacy Crust Bryozoan.” Project UFO (Unidentified Foreign Organisms), n.d. http://www.projectufo.ca/
drupal/Lacy_Crust_Bryozoan.
“Membranipora membranacea (Linnaeus 1767).” Nonindigenous Aquatic Species Program, U.S.
Geological Survey, 2009. http://nas3.er.usgs.gov/taxgroup/Bryozoans/.
“Membranipora membranacea.” Species Identification Card, Gulf of Maine Research Institute, n.d.
http://www.gmri.org/upload/files/VS_Membranipora_membranacea.pdf.
Telnack, Jennifer. “Membranipora membranacea: The LacyCrust Bryozoans.” Intertidal Marine
Invertebrates of South Puget Sound, n.d. http://www.nwmarinelife.com/htmlswimmers/m
_membranipora.html.
 CHAIN TUNICATE n 39

n Tunicates
n Chain Tunicate
Also known as: Chain sea squirt, orange or red sheath tunicate, violet tunicate
Scientific name: Botrylloides violaceus
Family: Styelidae
Native Range. East Asia, from southern Siberia to southern China and Japan. Those in
the United States are believed to have come from Japan.
Distribution in the United States. Chain tunicates have invaded both coasts of the
continental United States. Along
the Pacific Coast, they occur
in disjunct locations in Prince
William Sound and Sitka, Alaska;
Puget Sound and Willapa Bay,
Washington; Coos Bay, Oregon;
and Humboldt Bay, Bodega
Harbor, Tomales Bay, San
Francisco Bay, Half Moon Bay,
Monterey Bay, Elkhorn Slough,
Morro Bay, Santa Barbara,
Channel Islands Harbor, Port
Hueneme, Marina Del Rey, King
Harbor, Santa Catalina Island,
Alamitos Bay, Huntington
Harbor, Mission Bay, and San
Diego Bay, California. On the
Atlantic Coast, they are estab-
lished from Maine south to the
entrance to the Chesapeake Bay
and possibly to Florida.
Description. The chain tuni-
cate is a sessile, colonial sea
squirt consisting of many indi-
vidual small animals or zooids
arranged in elongated clusters
or double-rows and chains
called systems. Individual zooids
are about 0.1 in. (1–2 mm) long;
the largest colonies may be
about 1 ft. in diameter (0.3 m)
and sometimes develop lobes.
All the zooids in a given colony
Top: The chain tunicate Botrylloides violaceus comes from waters off East
are the same color: red, orange,
Asia. Bottom: Chain tunicates are invasive on both coasts of the
yellow, purple, or tan. continental United States. (Adapted from Fuller, P., “Botrylloides
Each zooid is vase-shaped violaceus.” USGS Nonindigenous Aquatic Species Database, Gainesville,
and sits upright on the substrate. FL. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=2418.)
40 n INVERTEBRATES (TUNICATES)

It separately draws water into its body through its own siphon but expels it into a common
space from which the waste water leaves the entire colony. The zooids are connected together
by a vascular network that ends in small sacs (ampullae) at the edge of the colony. These tiny
blobs are pigmented and the same color as the zooids. A clear, firm matrix forms between the
systems and surrounds the whole colony. This invertebrate is the only chordate known to be
able to regenerate its entire body from pieces of the colony’s vascular network.
Related or Similar Species. The golden star tunicate (Botryllus schlosseri) is another inva-
sive colonial tunicate found on both coasts of North America. As its common name suggests,
its zooids are arranged in star patterns and not in chains. Colonies display two colors rather
than just one. On the Pacific Coast, from Baja California to San Francisco Bay, one can find
Botrylloides diegensis, the individual zooids of which are dichromatic; the ring around the
oral siphon is a pale or bright white, yellow, or orange and contrasts with the darker color
of the body. A recently introduced colonial tunicate in San Diego and Mission bays,
Botrylloides perspicuum, is distinguished by a thicker and firmer matrix that forms ridges
between systems of zooids. There are also microscopic differences among these tunicates
and diagnostic characters in their larvae.
The chain tunicate might be mistaken for various red or orange sponges, but it is more
rigid than a sponge.
Introduction History. The earliest record of the chain tunicate in the United States is from
the Pacific Coast, where it was reported in 1973 in San Francisco Bay. It likely arrived as a
fouling organism on a ship from Japan, but could have been introduced with Japanese oys-
ters (Crassostrea gigas). It was first reported in Puget Sound in 1997 and in Prince William
Sound in 1999. However, the organism may have arrived earlier. Collection was limited
prior to the 1970s, and until 1997, the chain tunicate was not recognized as a form distinct
from the very similar B. diegensis. It was first reported on the Atlantic Coast in 1981,
although again, confusion with B. diegensis may have obscured an earlier arrival in the
1970s. In 1981, it was in Great Bay, New Hampshire, in the Gulf of Maine; by the mid-
1990s, it was in Penobscot Bay. Early records in the lower Chesapeake Bay date to 2000
and 2001. (In 2004, it had expanded northward and was reported on cultured mussels
being grown on Prince Edward Island, Canada, in the Gulf of St. Lawrence.)
Dispersal to new areas along the coasts of North America may occur in contaminated oys-
ter shell and spat associated with the oyster industry, as fouling organisms on the hulls of
ships, or by rafting on debris. Since the planktonic larval stage is so brief, it is unlikely that
it spreads far if at all in ballast water.
Habitat. Chain tunicates occur in sheltered areas of the shallow subtidal zone. Adults and
juveniles attach to submerged surfaces such as rocks, pilings, ropes, boat hulls, macroalgae,
eelgrass (Zostera marina), and the shells of mussels, oysters, and barnacles. They will also
grow over encrusting bryozoans and solitary tunicates. Chain tunicates can live in a range
of salinities (24–34 ppt) and temperatures (46–77°F [8–25°C]). It reportedly tolerates pol-
luted water.
Diet. Adults and juveniles are suspension feeders that filter phytoplankton and bacteria
from seawater.
Life History. Chain tunicates reproduce both sexually and asexually. The zooid is her-
maphroditic and viviparous. It has a brood pouch that extends out from the body wall into
the matrix. The egg is ovulated into the brood pouch, where it is fertilized. The embryo
develops into a relatively large (0.08–0.12 in. [2–3 mm] long), brightly colored, tadpole-
like larva. Gestation is longer than four weeks. Each pouch will hold one egg; each zooid
may have one or two pouches.
 CHAIN TUNICATE n 41

Upon release from the brood

pouch, the larva is free-
swimming for a very short time.
In just a few hours, metamor-
phosis takes place, and the
juvenile (oozoid) attaches head
first to a hard surface. The
newly attached oozoid meta-
morphoses into a zooid and
starts to reproduce asexually
by budding the next day. A
new colony is thus begun.
All the zooids of a given gen-
eration grow and die synchro-
nously. As the oldest generation
disintegrates, it is absorbed into Individual vase-shaped zooids of the chain tunicate are arranged in rows
the colony’s vascular system. and connected by a vascular system that ends in tiny sacs at the edge of
Impacts. Chain tunicates the colony. (Dann Blackwood, U.S. Geological Survey.)
may compete for settling space
and food with other native and introduced fouling organisms, but generally its ecological
impacts are unknown. It has become a nuisance species in mariculture operations produc-
ing mussels in Canada, where it overgrows and smothers the bivalve; so it could become a
pest in oyster operations in the Chesapeake Bay or wherever other shellfish are cultured.
As it becomes more abundant, it will become a problem, fouling fishing and boating gear
and requiring time and expense to remove.
Climate change may alter the story. Studies show that chain tunicates settle earlier and
grow faster than natives in warming waters. As natives decline in face of winter warming,
these tunicates may become dominant.
Management. Similar species have followed the typical pattern of a new invader: rapid
population growth followed by a period of decline and adjustment of the fouling community
without the loss of native species. It appears that the chain tunicate also will not replace
natives, and that management or eradication measures are unnecessary and would likely
be unsuccessful anyway.

Selected References
“Botrylloides violaceus.” Chesapeake Bay Introduced Species Database NEMESIS: National Exotic
Marine and Estuarine Species Information System, Smithsonian Environmental Research Center
(SERC), 2005, 2009. http://invasions.si.edu/nemesis/CH-INV.jsp?Species_name=Botrylloides
+violaceus.
Cohen, Andrew N. “Botrylloides violaceus Oka, 1927.” Guide to the Exotic Species of San Francisco Bay.
San Francisco Estuary Institute, Oakland, CA, 2005. http://www.exoticsguide.org/species_pages/b
_violaceus.html.
Pleus, Allen, and Pam Meacham. “Botrylloides violaceus (Chain tunicate),” Invasive Species Fact Sheets,
Aquatic Nuisance Species, Washington Department of Fish and Wildlife, n.d. http://wdfw.wa.gov/
fish/ans/identify/html/index.php?species=botrylloides_violaceus.
“Tunicate, Botrylloides violaceus.” Non-indigenous Aquatic Species of Concern for Alaska, Fact Sheet
15. Prince William Sound Regional Citizens’ Advisory Council, 2004. http://www.pwsrcac.org/
docs/d0016000.pdf.
 42 n INVERTEBRATES (TUNICATES)

n Colonial Tunicate
Also known as: Didemnid, colonial ascidian, colonial sea squirt, marine vomit
Scientific name: Didemnum vexillum
Synonym (in older literature): Didemnum sp., Didemnum sp. A.
Family: Didemnidae
Native Range. Uncertain; possibly Japan.
Distribution in the United States. East Coast, from Maine to New Jersey and offshore on
Georges Bank; West Coast, California and Washington.
Description. Didemnid colonies take various forms. Where the current is weak, they
hang from hard structures in long stringy lobes resembling ropes or beards. This is the usual
morphology on pilings, ship hulls, ropes, and the like. Where the current is strong, they
grow in dense flat mats with
low protruding lobes or tendrils
and encrust rock outcrops, peb-
bles, and cobbles. Colonies are
tan, cream-colored, white, or
yellow and may attain a diam-
eter of 3 ft. (1 m) or more.
Thousands of individual zo-
oids comprise each colony. Each
is about 0.1 in. (2.5 mm) long.
They are embedded in a clear,
firm matrix, the surface of which
is dotted with tiny white calcare-
ous balls, each covered with
spines. These may not be visible
to the naked eye. All zooids in a
given colony are the same color.
Related or Similar Species.
On the Pacific Coast, all three
related colonial tunicates (D. car-
nulentum, D. albidum, and Tridi-
demnum apacum) have spiny balls
throughout the matrix and not
limited to the surface as in D. vex-
illum. Their colonies are never
lobed, but always flat and en-
crusting; they are usually white
or gray. On the East Coast, the
colonial tunicate could be
confused with the chain tuni-
cate, Botrylloides violaceus (see
Tunicates, Chain Tunicate), but
Top: The colonial tunicate Didemnum vexillum may come from the waters the colonial tunicate is never the
off Japan. Bottom: D. vexillum has invaded both coasts of the continental red or orange color of chain
United States. (Adapted from “Species Didemnum vexillum.” USGS 2010.) tunicates.
 COLONIAL TUNICATE n 43

Colonial tunicates could also be mistaken for sponges, but they have a smoother texture
than a sponge.
Introduction History. The first documented occurrence of the colonial tunicate on the
East Coast was in Fort Island Narrows on the Damariscotta River in Maine in 1993. It was
likely present but unidentified prior to that time. Reports of a similar organism in
Walpole, Maine, date to 1988; and it is suspected that the tunicate was in New England
waters by the 1970s. Didemnum vexillum was first collected in Shinnecock Bay, Long
Island, New York, in late 2004. The first records on Georges Bank date to 2003 and are
the first evidence of invasion of offshore habitats.
On the West Coast, D. vexillum was collected from San Francisco Bay in 1993. It was
recovered from Elkhorn Slough, Monterey Bay, California, in 1998, but originally misiden-
tified as a native species, D. carnulentum. Since then, it has been collected in Bodega
Harbor, Humboldt Bay, Tomales Bay, Morro Bay, and Port San Luis. On the Washington
coast, it was first documented on a sunken wooden boat in Puget Sound in March 2004.
Colonial tunicates were most likely carried to the coasts of North America as fouling
organisms on transoceanic vessels. They also may have arrived as colony fragments in ballast
water. On the Pacific Coast, it is possible that they were accidentally introduced with oyster
or mussel stock or with equipment delivered to aquaculture facilities. Why this species has
suddenly become invasive is not known. However, fragments capable of starting new colo-
nies are easily broken off by bottom-fishing dredges, by divers, and by local boat traffic.
These pieces may then drift in currents over long distances to new sites.
Habitat. Colonial tunicates are usually associated with man-made structures in the subti-
dal zone of coastal waters. They have been found on docks, moorings, metal and wooden
pilings, ropes and steel chains, discarded automobile tires, and polythene plastic. They also
foul shellfish aquaculture gear and ships’ hulls. On the seabed, they encrust hard substrates
such as rock outcrops, cobbles, pebbles, and boulders and overgrow sessile benthic organ-
isms such as hydroids, solitary tunicates, sponges, barnacles, mussels, and oysters, as well
as seaweeds. They occur in shallow coastal waters and also on the continental shelf to depths
of about 200 ft. (65 m). These tunicates tolerate water temperatures ranging between 28°
and 75°F (−2°–24°C) and apparently require a salinity greater than 26 ppt.
Diet. These filter-feeders siphon seawater into their bodies and consume the phyto-
plankters and zooplankters, including the larvae of oysters and mussels, held in it.
Life History. The colony broods its larvae within its matrix and then releases them into
the water column. They are part of the plankton for only a few hours before they attach head
down to a firm surface. Larval settlement occurs in spring and fall. The settled larva quickly
metamorphoses into a zooid, which becomes the founder of a new colony. The colony
expands through budding (asexual reproduction). Young zooids reach sexual maturity in
just a few weeks.
A new colony can also form via fragmentation when a fragile lobe of the colony breaks off
and drifts to a new location, where it either settles to the bottom or becomes entangled in
marine structures. Such stray pieces may attach to a new substrate within six hours of arrival
and start overgrowing it.
Impacts. Colonial tunicates overgrow hard substrates and the native organisms attached
to them and thereby threaten to change benthic marine habitats. The high acidity of their
tunics may inhibit settling of the larvae of shellfish such as scallops. The overgrowth of grav-
elly bottoms could reduce habitat in which fish such as Atlantic cod (Gadus morhua) and
haddock (Melangrammus aeglefinus) deposit their eggs. The tunicates can smother bivalves,
 44 n INVERTEBRATES (TUNICATES)

Colonies of the colonial tunicate Didemnum vexillum take a variety of forms. A. Thousands of individual zooids,
each about 0.1 in. long, make up a colony. (Dann Blackwood, U.S. Geological Survey.) B. Stringy lobes occur
where currents are weak. Here Didemnum covers metal piling and a mussel shell. (Paul Barter, Cawthron
Institute, New Zealand.) C. Stiffer lobes protrude when currents are strong. Here the colonial tunicate has
encrusted a mussel shell. (Dann Blackwood, U.S. Geological Survey.) D. Mats overgrow the gravelly bottom of
Georges Bank. (Dann Blackwood, U.S. Geological Survey.)

so concern exists for oyster (Crassostrea spp.), bay scallop (Argopecten irradians irradians),
sea scallop (Placopecten magellanicus), and mussel fisheries, both natural and cultured.
Studies on Georges Bank show an increase in two polychaete species in areas infested
with the colonial tunicate, suggesting a change in native species composition on the seafloor.
Fouling of aquaculture gear, moorings, ships’ hulls, and so forth requires expensive mainte-
nance procedures and makes this a major nuisance species.
Management. Wrapping pilings in plastic may suffocate colonial tunicates, but this is
only of local significance as a method of control. Vacuuming colonies from the water has
been tried, but it did not succeed in eradicating the species. Prevention or slowing the
spread of the animal is difficult because of its fragile tunic and ability to rapidly reproduce
asexually. Boaters, divers, fishermen, and aquaculturalists could slow the invasion by keep-
ing equipment clean and not discarding bivalve shells in noninfested waters.

Selected References
Cohen, Andrew N. “Didemnum sp. A.” Guide to the Exotic Species of San Francisco Bay. San Francisco
Estuary Institute, Oakland, CA, 2005. http://www.exoticsguide.org/species_pages/didemnum
.html.
 AUSTRALIAN SPOTTED JELLYFISH n 45

Lengyel, Nicole L., Jeremy S. Collie, and Page C. Valentine. “The Invasive Colonial Ascidian Didemnum
vexillum on Georges Bank—Ecological Effects and Genetic Identification.” Aquatic Invasions (4):
143–52, 2009. doi:10.3391/ai.2009.4.1.15 http://www.aquaticinvasions.net/2009/AI_2009_4_1
_Lengyel_etal.pdf.
Morris, James A., Jr., Mary R. Carman, K. Elaine Hoagland, Emma R. M. Green-Beach, and Richard C.
Karney. “Impact of the Invasive Colonial Tunicate Didemnun vexillum on the Recruitment of the
Bay Scallop (Argopecten irrradians irradians) and Implications for Recruitment of the Sea
Scallop (Placopecten magellanicus) on Georges Bank.” Aquatic Invasions (4): 1: 207–11, 2009.
doi:10.3391/ai.2009.4.1.22 http://www.aquaticinvasions.net/2009/AI_2009_4_1_Morris_etal.pdf.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialty Group
(ISSG). “Didemnum spp. (Tunicate).” ISSG Global Invasive Species Database, 2007. http://
www.issg.org/database/species/ecology.asp?si=946&fr=1&sts=sss.
“Species Didemnum vexillum.” Marine Nuisance Species, USGS National Geologic Studies of Benthic
Habitats, Northeastern United States, 2010. http://woodshole.er.usgs.gov/project-pages/stell
wagen/didemnum/.

n Cnidarian
n Australian Spotted Jellyfish
Also known as: Spotted jellyfish, white-spotted jellyfish
Scientific name: Phyllorhiza punctata
Order: Rhizostomeae
Family: Mastigiidae
Native Range. Before the 1950s, the Australian spotted jellyfish was only known from the
Indo-Pacific region. Its native range is believed to have extended from the south-central
coast of eastern Australia northward into Southeast Asia and the Philippines.
Distribution in the United States. Populations are becoming established along the Gulf
Coast from Galveston Bay, Texas, to Orange Beach, Alabama, east of Mobile Bay. Another
permament population occurs on the Atlantic Coast in Indian River Lagoon, Florida. In
the summer of 2007, swarms of medusae were sighted off South Carolina, but these may
not represent a permanent population. In Hawai’i, populations occur in Pearl and
Honolulu harbors and in Kaneohe Bay, O’ahu. Australian spotted jellyfishes are also estab-
lished in Boqueron Bay, Puerto Rico.
Description. This is a large jellyfish with a somewhat flattened semi-globular gelatinous
bell or umbrella. In much of its natural and introduced range, it is bluish white or brown
due to symbiotic zooxanthellae; however, in the Gulf of Mexico, it is clear or white due to
the absence of these photosynthetic algae. Crystalline opaque white inclusions appear as
evenly spaced spots. Eight thick, branching oral arms surround the central mouth area; each
ends with a large brown bundle of stinging cells. Transparent ribbons of tissue hang from
each oral arm. The average bell diameter of an adult is 14 in. (35 cm), but in the Gulf of
Mexico, they are much larger than elsewhere, with an average diameter of nearly 18 in.
(45 cm). The largest reported was 24.4 in. (62 cm) in diameter. These large specimens can
weigh up to 25 lbs. (11 kg). They are only mildly venomous and not dangerous to humans.
Related or Similar Species. The native moon jelly (Aurelia aurita) is of similar size but
otherwise very different in appearance.
Introduction History. Australian spotted jellyfish likely entered the Atlantic Ocean and
Caribbean Sea on ships passing through the Panama Canal sometime in the early 1950s, if not
earlier. Polyps attached to the hulls of ships and/or young medusae held in ballast water could
 46 n INVERTEBRATES (CNIDARIAN)

have survived the journey from

the Pacific. The first colonies
appeared off northeast Brazil in
1995. Populations have been
established in Puerto Rico and in
other lagoons in the Caribbean
since at least the early 1970s.
The Australian spotted jellyfish
was first documented in the Gulf
of Mexico by a single specimen
collected in 1993. The natural
circulation pattern of ocean cur-
rents from the Caribbean into
the Gulf of Mexico could have
carried them northward to the
Gulf Coast of the United States,
where small populations such
as that in Terrebonne Bay,
Louisiana, became established in
the mid-1990s. They could also
have been introduced as attached
polyps on ships and towed struc-
tures. The proliferation of oil
platforms in the Gulf of Mexico
may have improved the habitat
for these jellyfish and facilitated
establishment of local popula-
tions by providing attachment
sites for the polyp stage and
allowing for natural dispersal
northward. Overfishing could
Top: The Australian spotted jellyfish was limited to the Indo-Pacific also have released resources pre-
region prior to the 1950s. Bottom: Australian spotted jellyfish are cur- viously tied up by native species.
rently established along the Gulf Coast of the United States and off The spectacular 2000 inva-
O’ahu, HI, and western Puerto Rico. (Adapted from Perry 2010.) sion of thousands of medusae
along the Gulf Coast, like the
later 2007 one, appeared to have come from the Caribbean though the Yucatán Straits
into the Gulf of Mexico, although recent genetic studies do not strongly support this
scenario and suggest that the Caribbean jellyfishes may be another species altogether.
Gulf Coast jellyfish are more similar to those from Australia or the West Coast, sug-
gesting an invasion via transoceanic shipping. Medusae were observed in Galveston
Bay in 2006.
The Australian spotted jellyfish was first identified on the Atlantic Coast in Indian River
Lagoon near Melbourne, Florida, in 2001. The 2007 sightings off the coast of South
Carolina represent the first known occurrence of the jellyfish north of Florida.
In Hawai’i, the jellyfish entered Pearl Harbor from the Philippines during World War II
(1941–1945). It appeared in Kaneohe Bay in 1953–1954. The first reports of Australian
 AUSTRALIAN SPOTTED JELLYFISH n 47

spotted jellyfish in coastal waters off the continental United States actually came from the
Pacific Coast, from southern California in 1981.
Habitat. Australian spotted jellyfishes usually inhabit warm coastal waters and lagoons
and may float into estuaries and bays on flood tides. They appear to prefer water tempera-
tures above 68°F (20°C) and salinities greater than 25 ppt. During the polyp stage of the life
cycle, hard substrates are needed for attachment.
Diet. These jellyfish use stinging cells to capture zooplankters, including the eggs and lar-
vae of many fish and shellfish. They feed almost constantly since they can digest the entire
content of their gut in two hours. They filter prodigious amounts of water each day, remov-
ing most of the small suspended particles. Except in the Gulf of Mexico, they also probably
benefit from the primary production of zooxanthellae living in their umbrella tissues.
Life History. Jellyfish under-
go alternate generations with
asexual and sexual stages in the
life cycle. The visible organism
is the adult medusa stage. Me-
dusae occur as separate sexes
that reproduce via external fer-
tilization and produce free-
swimming flat, ciliated larvae
(planula). The planulae settle
and attach to hard substrates
and begin the polyp stage of the
life cycle. Polyps develop into a
form (strobila) that asexually
produces free-swimming imma-
ture medusae (ephyrae). The
young medusae mature over sev-
eral weeks. Weak-swimmers,
they become part of the plankton
and are at the mercy of wind,
currents, and tides.
Australian spotted jellyfish in
the Gulf of Mexico experienced
huge population explosions
in 2000 and again in 2007.
During these episodes, the
medusae appeared in large num-
bers along the Gulf Coast of the
southeastern United States,
apparently carried in the Loop
Current and eddies of the Gulf
Stream. In contrast to these
irruptions, small populations
are maintained year after year in
places such as Terrebonne Bay, In the Gulf of Mexico, this large jellyfish is white because it lacks the
Louisiana, and Indian River symbiotic algae that make it brown or bluish white elsewhere in its
Lagoon, Florida. range. (Dwight Smith/Shutterstock.)
48 n INVERTEBRATES (ANNELID WORMS)

Impacts. The 2000 invasion was costly to the Gulf of Mexico shrimp industry. The large
gelatinous masses clogged nets and damaged gear and led to the closing of some productive
areas to commercial fishing. Concern exists that such large infestations of jellyfish that
coincide with the spawning season of many marine organisms could increase predation on
fish eggs and the larvae of mollusks and crustaceans to such a degree that forage fish such
as bay anchovy (Anchoa mitchilli) and oyster and crab populations in Mobile Bay and
Mississippi Sound could be affected. With their massive consumption of zooplankton, the
jellyfish potentially compete with native shrimp and fish for food and direct energy flow in
aquatic ecosystems to themselves, a dead end in the food chain since jellyfish have few
predators.
Management. There seem to be no strategies for control or eradication currently
available.

Selected References
“Australian Spotted Jellyfish.” Field Guide to the Indian River Lagoon. Smithsonian Marine Station at
Fort Pierce, n.d. http://www.sms.si.edu/IRLFieldGuide/Phyllo_punctat.htm.
Burkhard, Elizabeth. “A Survey of the Relationship of the Australian Spotted Jellyfish, Phyllorhiza punc
tata, and OCS Platforms.” GulfBase.org, n.d. http://www.gulfbase.org/project/view.php
?pid=asotrotasjppaop.
Masterson, J. “Species Report: Phyllorhiza punctata von Lendenfield, 1884.” Smithsonian Marine
Station at Fort Pierce, 2007. http://www.sms.si.edu/irlspec/Phyllorhiza_punctata.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Phyllorhiza punctata (Jellyfish)”. ISSG Global Invasive Species Database, 2006. http://
www.invasivespecies.net/database/species/ecology.asp?si=992&fr=1&sts.
Norris, Scott. “Australian Jellyfish Invade U.S. Waters.” National Geographic News, 2007. http://
news.nationalgeographic.com/news/2007/08/070827-jellyfish-invasion.html.
Perry, Harriet. “Phyllorhiza punctata.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL,
2010. Revised June 20, 2006. http://nas.er.usgs.gov/queries/FactSheet.aspx?speciesID=1192.
“Phyllorhiza punctata von Lendenfeld, 1884.” Hawaii Biological Survey, Bishop Museum, 2002. http://
www2.bishopmuseum.org/HBS/invertguide/species/phyllorhiza_punctata.htm.

n Annelid Worms
n European Earthworms
Also known as: Nightcrawlers, red worms, leaf worms, red wrigglers,
grey worms, angle worms
Scientific names: Lumbricus terrestris, L. rubellus, Aporrectodea caliginosa,
Dendrobaena octaedra, and others
Family: Lumbricidae
Native Range. The four species described below came from Europe. Of the known alien
earthworms in the United States, 25 are European and 14 are of Asian origin. Not all are invasive.
Distribution in the United States. Lumbricus terrestris and L. rubellus are found in two
disjunct areas. The eastern invasion region extends from the Atlantic coast of North
Carolina north through Maine and westward to the edge of the Great Plains. The western
part of the distribution area reaches along the Pacific coast of California north through
Washington and eastward to the foothills of the Rocky Mountains. Dendrobaena octaedra
 EUROPEAN EARTHWORMS n 49

has a continuous distribution

across the United States, avoid-
ing the Atlantic and Gulf
Coastal plains, Texas, the cen-
tral Great Plains, most of New
Mexico, and the southern parts
of California and Arizona.
Complete and accurate distri-
bution data for Aporrectodea cal-
iginosa are not available, but it is
widespread in northern forests.
Close relatives occur in grass-
lands, including the Palouse.
Other exotic earthworms,
including some from Asia such
as the Alabama jumper (Amyn-
thas agrestis), occur in south- Many of the alien earthworms in the United States, including the
ern forests. nightcrawler, red worm, angle worm, and octagonal-tail worm came from
Description. This entry per- Europe.
tains to several different annelid
or segmented worms from Europe, only a few of the 45 or more species introduced into the
United States. Basic earthworm identification can be based on gross characteristics such as
size, color, and the depth at which a species is found or the ecological group to which it
belongs (see under Habitat). A more precise and advanced analysis of the species that are
described in this entry relies on such anatomical details as the shape, size, and position of
the clitellum and the setae, tiny bristles on the body used for locomotion. All of the adult
worms are characterized by a smooth band or collar, the clitellum.
The nightcrawler (Lumbricus terrestris) is the largest of the common invasive earthworms,
attaining lengths of 5–8 in. (12.5–20 cm) or more. This reddish-brown anecic worm makes
permanent vertical burrows to depths of 6 ft. (2 m). Its presence may be evidenced on the
surface by a pile of casts (fecal material) and leaf stems some 2 in. (5 cm) across and 0.5–
0.75 in. (12.5–19 mm) high.
The red or leaf worm (L. rubellus) is a medium-sized annelid ranging from 1–4 in. (2.5–
10.5 cm) long. Its back is reddish brown and irridescent; the underside is pale. This is an
epi-endogeic species that occupies both the leaf litter and the uppermost few inches of the
mineral soil. This is the species commonly sold as fish bait.
The grey or angle worm (Aporrectodea caliginosa) is 2–6 in. (5–15 cm) long. It is unpig-
mented, but its internal organs and ingested food gives it a color ranging from gray or brown
to greenish. This species is endogeic and makes horizontal burrows in the topsoil layer.
The very small octagonal-tail earthworm (Dendrobaena octaedra) reaches lengths of only
0.6–2.5 in. (1.7–6 cm). They are reddish brown, darker on the back and head than the belly.
An epigeic species, this earthworm is only found in the leaf litter or duff of the forest floor.
Introduction History. European earthworms arrived in North America with early
European settlers beginning in the sixteenth century. They were likely carried in soils used
as ballast in ships and on the root balls of plants brought to the New World.
Reinforcements continue to arrive in contaminated shipments of ornamental plants and
intentional, permitted importations of live bait and composting worms from Canada, night-
crawlers, and red wrigglers (Eisenia veneta = E. hortensis), respectively.
 50 n INVERTEBRATES (ANNELID WORMS)

Northern forests developed

after the Ice Age without native
earthworms and are thus par-
ticularly vulnerable to invasion
by hardy species from similar
climate regions in Europe. On
their own, earthworms spread
very slowly, by an estimated
35 ft. (10 m) a year. In the
10,000 years that regions glaci-
ated during the Pleistocene
have been ice-free, native earth-
worms have been unable to
recolonize most of the area from
which they were eliminated by
continental ice sheets. How-
Top: Distribution of the octagonal-tail earthworm in the United States. ever, nonnative species have
Bottom: Distribution of the nightcrawler (L. terrestris) and the red worm been deliberately and widely
(L. rubellus) in the United States. (Both maps adapted from Proulx 2003.) introduced to agricultural areas
and garden plots, where they
are generally beneficial. Only recently have they invaded undisturbed forests and grasslands.
In glaciated regions, numerous lakes attract sport fishermen, and they seem to be respon-
sible for much of the spread of nightcrawlers and leaf worms into northern forests, as they
often dump unused live bait on the ground. The construction and logging industries also
move worms from place to place when cocoons and young worms become lodged in truck
tires. Earthworm concentrations tend to be highest near roads and recreation areas.
Homeowners bring composting worms and other garden worms to the forest edge as settle-
ment spreads into new areas. Earthworms only reached central Minnesota, for example, as
recently as the 1930s.
Habitat. Many of the European earthworms now found in the United States are closely
linked to disturbed and artificial habitats such as lawns, gardens, cultivated fields, and pas-
tures. They have recently been invading hardwood forests. Creatures of the underground,
they sort themselves into three main soil habitats. Anecic or deep-burrowing earthworms
excavate vertical burrows as deep as 6 ft. (2 m), but feed on the surface. Enogeic species bur-
row into the topsoil to depths up to 20 in. (50 cm) and come to the surface to feed. They are
responsible for much of the mixing of soil layers. Epigeic worms live in the litter and first
inch or so of forest soils. They do not excavate burrows.
A succession of earthworm species characterizes invasions of previously undisturbed for-
ests. Epigeic species such as Dendrobaena are the pioneers; they are followed by endogenic
types such as L. rubellus that prepare the soil for latecomers, the anecic nightcrawlers.
Diet. Earthworms eat and decompose leaf litter and also consume decaying organic mat-
ter and microorganisms. Nightcrawlers pull leaf debris down into their burrows; stems and
leaf fragments accumulate at the burrow entrance along with casts deposited by the burrow’s
inhabitant and form what are called middens. The leaf worm and grey worm feed on surface
litter, on organic material in humus-rich topsoil, and probably on fungi and bacteria in the
 EUROPEAN EARTHWORMS n 51

rhizosphere, that nutrient-enriched area surrounding the roots of plants. The octagonal-tail
earthworm feeds mainly on the bacteria and fungi of the litter layer.
Life History. Earthworms are hermaphroditic, each individual possessing both testes and
ovaries. They usually mate sexually with another individual, however, lining up “head to
toe” so that the clitellum of one worm lies against the segments of the other that contain
the male reproductive organs. The clitellum secretes a large amount of slime that encases
the two worms in a slime tube. The sperm are released into the slime and carried in special
grooves back to the sperm receptacles of the other individual. The worms then separate.
A second mucous ring is secreted from the clitellum, and it slides forward over the body.
As it passes the openings to the female organs, several ripe eggs are ejected, sticking to the
ring. The ring continues to move forward and brings the eggs into contact with the sperm,
which fertilize the eggs. The entire ring then passes over the head and away from the worm.
Its ends seal to make a cocoon. The cocoon lies in the soil, into which tiny young worms
emerge when the eggs hatch.
Impacts. Earthworms are well known as beneficial additions to garden and agricultural
soils, where they help decompose organic matter and accelerate nutrient cycles, aerate the
soil and increase water infiltration with their burrowing, and mix soil layers to return
nutrients to the root zone. They are the focus of a multimillion-dollar vermiculture industry
that produces worms for composting and for live bait. It is therefore surprising to many peo-
ple that they become a major ecological disaster when introduced to northern hardwood
forests.
In deciduous hardwood forests, invasive earthworms are “ecosystem engineers” that alter
the habitat. They increase decomposition rates of the leaf litter to the degree that they
destroy the duff and humus. These organic soil layers not only store nutrients until the
spring growing season, but provide insulation and protection from predation for seeds and
seedlings. In their absence, nutrients such as nitrates are leached, and seeds do not
germinate; bare spots open
to erosion develop. Exotic
weeds such as garlic mustard
(See Volume 2, Forbs, Garlic
Mustard) (Alliaria petiolata)
and Japanese barberry (See
Volume 2, Shrubs, Japanese
Barberry) (Berberis thunbergii)
that evolved in the presence
of earthworms can invade.
The loss of the herb layer of
native wildflowers such as tril-
liums (Trillium spp.), blood-
root (Sanguinaria canadensis),
trout lily (Erythonium ameri-
canum) and blue cohosh
(Caulophyllum thalictroides)
has been noted in forests in The various introduced earthworms resemble each other but tend to be
the Great Lakes area as a result found at different depths in the soil. (Vinicius Tupinamba/Shutterstock.)
52 n INVERTEBRATES (ANNELID WORMS)

of earthworm invasions. The rare goblin fern or moonwort (Botrychium mormo) has been
extirpated from some areas
The litter is also habitat to a number of small vertebrates, invertebrates, and microbes.
When it disappears, so do they. A more direct link may be made between earthworm inva-
sions and declines in native salamanders that feed on worms. Larger worms such as leaf
worms and nightcrawlers will be consumed by adult salamanders, but are too big to be
ingested by juveniles. If the exotic earthworms replace native species, salamander recruit-
ment could be significantly reduced.
White-tail deer (Odocoileus virginianus) and earthworms may be connected to changes in
the species composition of forest communities. Where deer yard up during winter, their
feces accumulate and attract and nourish earthworms. An increase in earthworms reduces
the litter layer and makes forest herbs more vulnerable to deer browsing. Together, they con-
tribute to the loss of herbaceous plants and tree seedlings.
Their deposition of casts, cementing together of soil particles while burrowing, displace-
ment of other bioturbating organisms, and consumption of bacteria and fungi are some of
the means by which earthworms alter the soil environment. With time, earthworm activities
lead to soil compaction at depth. Compaction can bind some nutrients, prevent root pen-
etration, reduce aeration and available soil moisture, and increase runoff and removal of
phosphorus. The mixing of soil layers may change both the soil structure and its biochemi-
cal properties. It may increase weathering rates, but studies have yet to determine if the
result will be a greater release of greenhouse gases to the atmosphere or increased storage
of carbon in soils. Either negatively or positively, earthworms may be involved in climate
change processes.
Management. Prevention of the introduction of new species to the United States and the
spread of existing invasive earthworms are paramount. Gardeners should not dispose of
compost in natural areas, and fishermen should not dump unused live bait. Inspection of
vermiculture products could ensure that the species being sold is truly the one intended
and not some other that entered a batch through misidentification or contamination. The
horticultural trade in live plants needs better control of accidental migrants in pots and root
balls. Vehicle tires could be hosed down or otherwise sanitized before entering areas most
vulnerable to invasion.

Selected References
Baskin, Yvonne. Under-Ground. How Creatures of Mud and Dirt Shape Our World. Washington, DC:
Island Press, 2005.
Blatchford, John. “Earthworms Spread by Human Activity: Worms Improve Soil Fertility but Invasive
Species do Damage.” Suite 101.com, 2009. http://zoology.suite101.com/article.cfm/
earthworms_spread_by_human_activity#ixzz0eJavfI9t.
Dunne, Niall. “Invasive Earthworms—A Threat to North American Forests,” Plants and Gardens News:
19(1): 2004. http://chicagoconservationcorps.org/blog/wp-content/uploads2/2009/06/L40
%20Vermicomposting%20and%20Invasive%20Earth%20Worms.pdf.
“Earthworm Ecological Groups.” Natural Resources Research Institute, University of Minnesota,
Duluth, 2006. http://www.nrri.umn.edu/worms/identification/ecology_groups.html.
Halsey, Daniel. “Invasive Earthworms: Affects on Native Soils.” Soil5125, University of Minnesota,
2009. http://southwoodsforestgardens.blogspot.com/2009/01/paper-on-invasive-european
-worms.html.
 ASIAN CLAM n 53

Proulx, Nick. “Ecological Risk Assessment of Non-Indigenous Earthworm Species.” Final draft.
Minnesota Department of Natural Resources, 2003. http://www.nrri.umn.edu/WORMS/research/
publications/Proulx%202003.pdf.

n Mollusks
n Asian Clam
Also known as: Asiatic clam, gold clam
Scientific name: Corbicula fluminea
Family: Corbiculidae
Native Range. East Asia, from southeastern China to Korea and southeastern Russia.
Distribution in the United States. The Asian clam has established populations in all states
except Alaska, Maine, Montana, New Hampshire, North Dakota, Rhode Island, South
Dakota, and Wyoming. It also occurs in Puerto Rico.
Description. This small freshwater bivalve is distinguished by a rounded to somewhat tri-
angular shell with concentric, evenly spaced ridges. The umbo or raised area above the hinge
is high and centrally positioned on the shell. On the inside of the valves, long, straight, serrated
ridges, the lateral teeth, emanate from the hinge area, two on each side of the right valve and
one on each side of the left valve. Three cardinal teeth project below the umbo on each valve.
Two color morphs occur in the United States. In most parts of the country, shells are yellow-
brown, but in the southwest, they are black. In the lighter-colored specimens, the inside of
the shell is a glossy light purple nacre (mother-of-pearl); in dark morphs, the nacre is deep
blue. Adult shell length rarely exceeds 1.5 in. (40 mm); usually it is less than 1 in. (25 mm).
Related or Similar Species. Asian clams could be confused with native fingernail or pea
clams (Family Sphaeriidae). Fingernail clams lack both lateral teeth and cardinal teeth and
are typically less than an inch (25 mm) long. The Asian clam has a heavier shell than most
native species.
Introduction History. Asian clams were first collected in the United States in 1938 in the
Columbia River near Knappton, Washington, but it is believed that they had probably
occurred in some West Coast drainages since at least 1924. The clams were an important
food for Chinese people, who may have deliberately brought them along when they emi-
grated to North America. It is also possible that they arrived accidentally in contaminated
imports of the giant Pacific oyster (Crassostrea gigas), a shellfish first imported into
Washington from Japan in 1903 and now the basis of a major aquaculture industry.
Asian clams were discovered in the Ohio River in 1957 and quickly spread throughout
the Mississippi River drainage system and into Lake Erie and Lake Michigan. On the
Atlantic slope, they were first reported from Delaware in 1972, New Jersey in 1982, and
Long Island, New York, in 1984. They were not expected to be able to survive New
England’s cold winters, but were found in the lower Connecticut River in 1990 and in sev-
eral Massachusetts reservoirs in 2007.
In Hawai’i, the Asian clam first appeared at a farmer’s market on O’ahu in 1977. Within a
few years, it was established in many streams and reservoirs on O’ahu as well as on the
islands of Kaua’i, Maui, and Hawai’i.
 54 n INVERTEBRATES (MOLLUSKS)

One of the most recently

documented invasions has
occurred in Lake Tahoe. From
a relatively few individuals
spotted there in 2002, the pop-
ulation had exploded by 2009.
These mollusks can disperse
naturally within a water body,
since juveniles float on cur-
rents. Overland transport is
accomplished on contaminated
boats, trailers, and aquatic
sports gear, or, since they are
used as fish bait, when unused
bait is dumped into a stream,
lake, or reservoir.
Habitat. Asian clams inhabit
the bottom sediments of stre-
ams, rivers, lakes, and ponds
of all sizes, and will also invade
irrigation canals. The substrate
can be silt, sand, or gravel.
They require well-oxygenated
water, so they prefer flowing
water and are intolerant of pol-
luted water. However, they can
withstand brackish water with
salinities up to 13 ppt for short
periods of time; and estuarine
populations occur in the
Chesapeake Bay and in San
Top: Asian clams came from eastern Asia. Bottom: The Asian clam is Francisco Bay. Water tempera-
invasive in almost all states as well as in Puerto Rico. (Adapted from tures above 60°F (16°C) are
USGS 2001.) needed for spawning and
release of young. Asian clams
can tolerate water temperatures as high as 86°F (30°C); they do not survive when tempera-
tures drop below 35°F (2°C). In cold-weather regions such as the Great Lakes, there may be
large fluctuations in population size from year to year.
Diet. These filter-feeders are herbivores, consuming mostly phytoplankters sieved from
the water column.
Life History. Asian clams are hermaphrodites and capable of self-fertilization. Sperm are
released into the water column and are taken in by other individuals. Eggs are fertilized in
the inner gills, and there the larvae are incubated. Juveniles are released through the excur-
rent siphon when about 0.04 in. (1 mm) in size. They are poor swimmers and sink to the
bottom, where they begin their adult stage. Reproduction and release of juveniles usually
occurs twice a year, in the spring and again in late summer or fall. Juveniles become sexually
mature at shell lengths of 0.2–0.4 in. (6–10 mm). Lifespan is one to four years.
 ASIAN CLAM n 55

A. This small clam, usually less than 1.0 in. across, has a rounded shell with concentric, evenly spaced ridges
and a high, centrally positioned umbo. (U.S. Geological Survey.) B. Asian clams with a nickel for scale. (Shawn
Liston, Bugwood.org.)

Impacts. Asian clams are notorious biofouling organisms, clogging intake pipes of power
plants and other facilities. The weak-swimming juveniles descend to the base of the water
column from whence intake pipes usually withdraw water. Live animals, empty shells, and
dead clam bodies are sucked in. This has proved to be a particular problem at nuclear power
plants, where water is used for fire protection.
Another type of economic consequence occurs where stream beds are dredged for sand
and gravel to use as an aggregate in cement, as happens in Ohio and Tennessee. The clams
end up in the cement but worm their way to the surface as it starts to set, leaving empty tun-
nels that weaken the structure.
Large congregations of Asian clams can alter the benthic habitat, which they may come to
dominate, and may compete with native fingernail clams for food and space. Recent studies
in Lake Tahoe, for example, show an increase in a filamentous green alga (Zygnema sp.) in
clam beds, probably thriving on the bivalves’ waste products.
Management. Prevention of introduction to new bodies of water is the most effective way
to slow or stop the invasion of Asian clams. Boats and equipment should be inspected,
cleaned of any obvious attached plants and animals, and drained of all water on land before
they leave the launch area. Bait buckets should be emptied on land, not in water. Before
launching into another body of water, boats and all equipment should be cleaned in hot
water and dried for several days.
Chemical (bromine and chlorine) and mechanical (screens and traps) treatments can
control infestations in intake pipes.

Selected References
Balcom, N. C. “Aquatic Immigrants of the Northeast, No. 4: Asian Clam, Corbicula fluminea,”
Connecticut Sea Grant College Program, 1994.
Foster, A. M., P. Fuller, A. Benson, S. Constant, and D. Raikow. “Corbicula fluminea.” USGS
Nonindigenous Aquatic Species Database, Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/
FactSheet.aspx?speciesID=92.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Corbicula fluminea (Mollusc).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?fr=1&si=537.
56 n INVERTEBRATES (MOLLUSKS)

USGS. “Asian clam, Corbicula fluminea (Muller, 1774) (Mollusca: Corbiculidae)”. Nonindigenous
Species Information Bulletin, Florida Caribbean Science Center, 2001. http://fl.biology.usgs.gov/
corbicula4.pdf.

n Asian Green Mussel

Also known as: Green mussel, green-lipped mussel
Scientific name: Perna viridis
Synonyms: Mytilus viridus, Chloromya viridis
Family: Mytilidae
Native Range. Coastal waters of southern Asia from the Persian Gulf to southern China
and the Philippines.
Distribution in the United States. Gulf and Atlantic coasts of Florida northward through
coastal Georgia as far as Charleston, South Carolina.
Description. This large marine bivalve has a smooth, elongated, fan-shaped shell.
The thin valves clearly show concentric growth rings; the ventral margin is concave on one
side. The entire shell of juveniles is bright green, but in adults, it is brown with green mar-
gins. The inner shell surfaces are smooth and iridescent blue or blue-green. The beak, at
the hinge, has interlocking teeth: two on the left valve and one on the right. Juveniles and
adults secrete hairlike byssal threads with which to attach to hard substrates.
Related or Similar Species. A number of native mussels in the same family as the Asian
green mussel occur in states infested with this nonnative bivalve. They have similar shapes,
but none are green. The scorched mussel (Brachidontes exustus), ribbed mussel (Guekensia
demissa), and hooked mussel (Ischadium recurvum), the native species most apt to inhabit
the same sites as Asian green mussels, have ribbed rather than smooth shells. The shells
of the smaller horsemussels (Modiolus spp.) are brown on the outside and white on the
inside. The smooth outer shell of the tropical charru mussel (Mytella charruana), another
nonnative mussel found (rarely) in Florida’s waters, has a dark-brown wavy pattern on a
lighter background; the inner shell is iridescent purple.
Introduction History. Asian green mussels were first reported in Florida in Tampa Bay in
1999, when they clogged intake pipes at a power plant. This was probably the result of a dis-
charge of ballast water carrying larvae by a ship coming from the Caribbean. Asian green mus-
sels had arrived in Trinidad in 1990 and rapidly spread through the Caribbean. Drifting in
currents, larval green mussels have dispersed southward as far as Marco Island, near Naples.
A separate invasion event occurred near St. Augustine on the Atlantic side of the Florida penin-
sula in 2002. They likely arrived on a recreational or fishing boat transported overland from the
Gulf coast without proper decontamination of hull, live wells, or gear. Northward-flowing
ocean currents dispersed the mussel to Jacksonville, Florida, and along the entire Georgia coast
by the end of 2003. In 2006, they were reported at Charleston, South Carolina. Mussels also
dispersed southward; their range currently extends to near Titusville, Florida.
Habitat. Asian green mussels inhabit coastal and estuarine waters in the intertidal and subti-
dal zones to depths near 35 ft. (10 m). They tolerate a wide range of salinities, from hypersaline
(80 ppt) to brackish (12 ppt). Optimal salinity is reportedly 27–33 ppt. They also tolerate a wide
range of temperatures between 50° and 95°F (10–35°C), but do best in water temperatures of
79–90° F (26–32° C). Juveniles appear to prefer to settle in areas of high water flow about 1 ft.
(30 cm) below the low-tide mark. They seem to seek crevices or the undersides of floating objects.
Diet. These mollusks are sessile filter-feeders. An incurrent siphon draws in seawater that
is moved by cilia to the branchial cavity. Phytoplankters, zooplankters, and other organic
 ASIAN GREEN MUSSEL n 57

particles are filtered out. Mucus

is secreted to bind the food
items into a bolus that is direct-
ed by cilia to the mouth. Water
exits via an excurrent siphon.
Life History. The sexes are
separate animals that release
eggs and sperm into the water
column during spawning epi-
sodes that peak in spring and
autumn. Fertilization takes
place externally. The presence
of gametes in the water stimu-
lates other individuals to release
eggs and sperm to synchronize
spawning within a local popula-
tion. The dilution of seawater,
such as what happens at the
onset of a rainy season, also
seems to trigger spawning.
Within 8 hours of fertilization,
the embryos become ciliated,
free-swimming larvae and,
8–12 hours later, have a shell
and ciliated membrane called a
velum. The larvae metamor-
phose into juveniles and settle
8–20 days later. The juveniles
secrete byssal threads and
attach to hard surfaces. In
Tampa Bay, they reportedly
become sexually mature in Top: The Asian green mussel is native to coastal waters of southern Asia.
1–2 months when shell length Bottom: In the United States, the Asian green mussel is invasive along
is 0.6–1.2 in. (15–30 mm). both the Gulf and Atlantic coasts of Florida and has currently expanded
They live about three years. its range as far north as South Carolina. (Adapted from McGuire and
Impacts. Asian green mus- Stevely 2009.)
sels are marine biofouling organisms, acting much like the freshwater zebra mussel (see
Mollusks, Zebra Mussel). They can clog intake screens and pipes of power and desaliniza-
tion plants and cover the hulls of boats and submerged parts of buoys, bridges, piling, and
seawalls. Not only do they block the flow of water, but they also damage pumps, reduce heat
transfer efficiency, and accelerate the rate of corrosion of metal surfaces. Economic impacts
of increased plant maintenance can be significant.
Since the Asian green mussel can rapidly build up large populations that form mats on
the seafloor, it is suspected that they will complete for space and possibly for planktonic
food supplies with native bivalves. In Tampa Bay, green mussels seem to have expanded at
the expense of native oysters (Crassostrea virginica). Elsewhere, for example in Georgia,
where oysters live in habitats where they are exposed at low tide, such replacement does
not seem to be happening.
 58 n INVERTEBRATES (MOLLUSKS)

A. Asian green mussel shell shows distinct beak and concentric growth rings. (U.S. Geological Survey.) B. The
concave ventral margin and smooth inner shell surfaces are visible on some of these specimens. (U.S.
Geological Survey.)

The success of the invasion of Asian green mussels in the coastal waters off Florida sug-
gests they would be equally successful if accidentally introduced to other sites along the
Gulf coast or to southern California.
Management. Currently, most management consists of mechanically removing mussels
from intake screens and pipes or flushing those systems with chlorinated water. The main
focus is to prevent further introductions of the organisms by educating commercial fisher-
men and recreational boaters and fishermen to inspect and clean the hulls of their vessels
and to drain live wells and bilges on land after taking their boats out of the water.

Selected References
Masterson, J. “Species Report: Perna viridis.” Smithsonian Marine Station at Fort Pierce, 2007. http://
www.sms.si.edu/irlSpec/Perna_viridis.htm.
McGuire, Maia, and John Stevely. “Invasive Species of Florida’s Coastal Waters: The Asian Green
Mussel (Perna viridis).” Publication #SGEF 175, Florida Sea Grant College Program. Institute of
Food and Agricultural Sciences (IFAS), University of Florida, 2009. http://edis.ifas.ufl.edu/sg094.
Powell, Cindie. “Asian Invader Musseling in on U.S. Habitats.” Texas Sea Grant and NOAA, 2005.
http://www.oar.noaa.gov/spotlite/archive/spot_greenmussel.html.
Thornton-DeVictor, Susan, and David Knott. “The Asian Green Mussel: Recent Introduction to the
South Atlantic Bight.” Species of the Month, Southeastern Regional Taxonomic Center, South
Carolina Department of Natural Resources, n.d. http://www.dnr.sc.gov/marine/sertc/The%
20Asian%20Green%20Mussel.pdf.

n Chinese Mystery Snail

Also known as: Asian applesnail, Japanese mystery snail, black snail, trapdoor snail
Scientific name: Cipangopaludina chinensis malleata
Synonyms: Bellamya chinensis, Viviparus chinesis, Viviparous stelmaphora,
Paludina malleata et al.
Family: Viviparidae
 CHINESE MYSTERY SNAIL n 59

Native Range. Southeast

Asia. It occurs in Myanmar
(Burma), Thailand, Indonesia
(Java), Vietnam, China, Korea,
Japan, and the Philippines.
Distribution in the United
States. Chinese mystery snails
are reported in 27 states. In the
eastern United States, popula-
tions occur in New England
and the Great Lakes region,
and on the West Coast, they
can be found from the San
Francisco Bay area north to
Seattle. Mid-continent records
come from Colorado, Iowa,
Nebraska, Oklahoma, Texas,
and Utah.
Description. These gastro-
pods are distinguished by their
large size, adults reaching
lengths of 1.5 in. (60–65 mm)
from the tip of the whorl to the
lip of the shell. The shell has
6–7 whorls with rounded
shoulders and indented sutures;
in adults, it is a uniform olive
green, greenish brown, brown,
or reddish brown without band-
ing on the outside and white to
pale blue on the inside. The
dark solid operculum, a func- Top: The Chinese mystery snail is native to Southeast and East Asia.
tioning “trapdoor,” is marked Bottom: Chinese mystery snails are reported from 27 states. (Adapted
with concentric rings. The outer from Kipp and Benson 2007.)
lip is round to oval and black.
Related or Similar Species. Taxonomic confusion exists as to whether there are two
forms of the Chinese mystery snail in the United States, or whether the Japanese mystery
snail (C. japonica) is a separate species with a more elongate shell and other subtle morpho-
logical differences. The banded mystery snail (Viviparus georgianus), native to some parts of
the United States but an introduced species in the Great Lakes region and eastern states
north of the Carolinas, is smaller than the Chinese mystery snail, with a maximum shell
length of 1.75 in. (45 mm); its shell is encircled with obvious reddish-brown bands. The
brown mystery snail (Campeloma decisum), native to the eastern United States, only rarely
grows as large as the Chinese mystery snail and is much narrower than either the Chinese
or banded mystery snails. Its shell is usually olive green. The brown mystery snail is one of
very few native snails with an operculum.
Introduction History. The earliest record of Chinese mystery snails in the United States
dates to 1892 and San Francisco, where they were imported for the live-food market.
 60 n INVERTEBRATES (MOLLUSKS)

By 1911, a free-living population

was thriving in San Francisco
Bay. Snails may have been
accidentally introduced into
Massachusetts in the early 1900s
with goldfish released to control
mosquitoes. A population was
established in Boston by 1915,
perhaps a by-product of the local
Asian food market. Snails entered
Lake Ontario from the Niagara
River between 1931 and 1942.
Chinese mystery snails were
reported in Florida in 1950 and
were established in Texas and
Lakes Erie and Michigan and
their drainages by 1965. Snail
The shell of the Chinese mystery snail has indented sutures between the introductions initially seem to
rounded whorls. (Pieter Johnson, University of Colorado.) have been intentional releases
either to develop a local food
supply or from the freshwater aquarium trade, in which Chinese mystery snails are used to keep
fish tanks clean of algae growths. They may be unintentionally moved from one body of water to
another as contaminants in live bait (e.g., minnows and crayfish) or on plants and in water trans-
ported on recreational watercraft and boat trailers. It takes only one pregnant female to start a
new population.
Habitat. These freshwater snails prefer quiet waters with soft substrates of silt, sand, or
mud. They can be found in lakes, ditches, rice paddies, and slow-moving streams at water
depths of 1.5 to 15 ft. (0.5 to 5 m). They can tolerate pollution and may thrive in stagnant
water, but they cannot survive very low oxygen levels and experience major die-offs under
a combination of warm water and algal blooms that reduce dissolved oxygen content.
Diet. Chinese mystery snails feed on organic and inorganic material on the bottom of
water bodies and scrape algae from hard surfaces. They also consume zooplankters and
phytoplankters.
Life History. These snails are live-bearers. In the eastern United States, embryos develop
in the female between May and August, and the young are born in shallow water from June
through October. After birthing, females retreat to deeper water for the winter months. Each
female may produce as many as 100 tiny snails in a single brood. (It is the sudden appear-
ance of tiny, perfectly developed snails that may be the “mystery” of these snails.) Females
live up to five years and tend to have their largest broods in their later years. Males live on
average three years.
Impacts. The ecological impacts, if any, of this introduced species remain unknown. It is
possible they could compete with native snails for food and space, but little indication exists
that this is happening. Chinese mystery snails do carry parasites, including one that can
infect humans, but do not seem to be a vector for “swimmer’s itch” as some have feared.
The species can clog the screens of even large water-intake pipes.
Management. Prevention of new infestations is the best control measure available.
Eradication of existing populations is likely impossible. People should refrain from dumping
 COMMON PERIWINKLE n 61

bait and aquarium contents and should sanitize fishing and boating equipment before enter-
ing another body of water. Live animals of any sort should never be released into the wild.

Selected References
“Aquatic Invasive Species: Chinese Mystery Snail.” Indiana Department of Natural Resources, 2005.
http://www.in.gov/dnr/files/CHINESE_MYSTERY_SNAIL.pdf.
Kipp, Rebekah M., and Amy Benson. “Cipangopaludina chinensis malleata.” USGS Nonindigenous
Aquatic Species Database, Gainesville, FL, 2007. http://nas.er.usgs.gov/queries/FactSheet.asp
?speciesID=1045.
“Mystery Snail Monitoring Protocol.” Aquatic Invasive Species Monitoring Manual—Citizen Lake
Monitoring Network, 2009. http://www.uwsp.edu/cnr/uwexlakes/clmn/.

n Common Periwinkle
Also known as: Edible periwinkle, wrinkle winkle
Scientific name: Littorina littorea
Family: Littornidae
Native Range. Europe, from the coast of northern Spain, throughout the British Isles, and
north to Scandinavia and Russia.
Distribution in the United States. Established along the coast of the northeastern United
States, from Maine to Virginia and the Chesapeake Bay. On the West Coast, individuals have
been collected from California to Washington.
Description. This small marine snail has a conical shell that comes to a point at the apex.
Shells of young animals show distinct ridges spiraling up to a prominent point; in adults, the
ridges become indistinct and the shell appears smooth. Five or six somewhat swollen whorls
are outlined by shallow sutures. The base color of the shells is usually grayish brown or
black, getting paler near the top. Dark lines form a spiral pattern on much of the shell, but
the central axis or columella is white, as is the inside of the shell. The thick inner lip of the
aperture is bent over the columella. Average shell height is 0.6–1.5 in. (16–38 mm), but it
may be as great as 2.0 in. (52 mm). Juvenile stages have black barring on their flat, broad
tentacles.
Related or Similar Species. The ridged shells of young common periwinkles could be
mistaken for those of the rough periwinkle (Littorina saxatilis), which is native to the coasts
of eastern North America from the Chesapeake Bay northward. Rough periwinkles have
4–5 whorls and grow to only about 0.7 in. (18 mm). The exterior of the shell may bear a
checkered pattern; the inside is brown. They tend to inhabit salt marshes, but can be found
on rocky coasts. Rough periwinkles have been introduced to San Francisco Bay
Introduction History. Live common periwinkles were first identified in North America in
1840 at Pictou, Nova Scotia, Canada. From this site in the Northumberland Straits of the
southern Gulf of St. Lawrence, it moved to the Atlantic Coast, being discovered in Halifax,
Nova Scotia, in 1854. By 1861–1862, it had appeared at Eastport, Maine. Soon thereafter,
it was found on Cape Cod (1872), and by 1890, it had reached Cape May, New Jersey. It
appears that a separate introduction occurred sometime in the early 1950s on the
Delmarva Peninsula. Southward expansion from this point of entry was relatively slow; it
took about 25 years to reach the entrance to Chesapeake Bay (1978), its current southern
range limit.
 62 n INVERTEBRATES (MOLLUSKS)

It is generally agreed that the

common periwinkle arrived in
Nova Scotia from Great Britain
on rocks used as ballast in trans-
oceanic ships, possibly those
involved in the early and mid-
nineteenth-century timber trade.
The initial inoculation was fol-
lowed by rapid population
growth and range expansion
along the rocky coasts and
salt marshes of New England.
However, not all agree with this
scenario and raise the possibil-
ities that (a) the periwinkle was
actually native to northeast
Canada and experienced a range
expansion only after European
settlement; or (b) that it was car-
ried to Canada by the Vikings in
pre-Columbian times. Most evi-
dence to date, including informa-
tion from prehistoric and historic
archaeological sites and modern
genetic analyses, support a
nineteenth-century introduction.
Habitat. These mobile gas-
tropods inhabit coastal and
estuarine environments. The
most typical habitat for com-
mon periwinkles is on all but
Top: The common periwinkle is native to the Atlantic and Baltic coasts of the most exposed rocky coasts
Europe. Bottom: The common periwinkle is a dominant member of rocky in the intertidal zone from the
coast communities in the northeastern United States and has been spray zone of the upper littoral
collected in California and Washington State. (Adapted from Benson 2009.) to the sublittoral zone that is
exposed only during the lowest
low tides. In sheltered areas, they may found in salt marshes and on mudflats.
Diet. Common periwinkles are herbivores and graze on benthic diatoms and dinoflagel-
lates as well as macroalgae such as sea lettuce (Ulva lactuca) and other large green algae and
young and ephemeral red and brown algae. While grazing on rocks, they may ingest small
invertebrates such as barnacle larvae.
Life History. Common periwinkles have separate sexes and reproduce annually. Females
release their fertilized eggs into the ocean in horny capsules about 0.03 in. (1 mm) wide.
Each capsule is convex on both sides and usually contains 2–3 eggs, although as many as
9 eggs may be inside. The shedding of egg capsules coincides with spring tides (i.e., when
the tidal range is greatest). The larvae exit the capsules and settle within six weeks.
Juveniles mature at 2–3 years of age when shell height is about 0.4 in. (10 mm). The lifespan
of a common periwinkle is 5–10 years.
 COMMON PERIWINKLE n 63

A. Common periwinkles are one of the most common mollusks in intertidal communities north of New Jersey. (J.
Pederson. Reprinted with permission of the MIT Sea Grant College Program.) B. The conical shell spirals up into
a prominent point, while the inner lip of the aperture folds over the columella. (Amy Benson, U.S. Geological
Survey.)

Impacts. Common periwinkles quickly became one of the most common mollusks on the
Atlantic Coast north of New Jersey and an ecological dominant in intertidal communities. In
both exposed rocky coast habitats and estuarine situations densities may reach 20–45/sq. ft.
(200–500/m2). Its consumption of green algae has allowed Irish moss (Chondrus crispus), a
slow-growing foliose red alga, to proliferate. Constant grazing by periwinkles of young algae
prevents the reestablishment of the original algal canopy on rocks and removes a key food
item of native marine snails. On rocky coasts, common perwinkles have displaced the
eastern mudsnail (Nassarius obsoletus) from many habitats and caused a niche shift and pop-
ulation reduction in the rough periwinkle (Littorina saxatilis). In estuarine habitats, its bull-
dozing of soft sediments slows the accumulation of fine materials and stifles the growth of
the root mat of salt marsh cordgrass (Spartina alterniflora).
Management. Few management practices have been developed to deal with common
periwinkles, and little research is conducted on them. They are today well-established mem-
bers of intertidal communities. Indeed, they may gain value as a bio-indicator of contami-
nated marine habitats.

Selected References
Benson, A. J. “Littorina littorea.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL, 2009.
http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=1009.
Chapman, John W., James T. Carlton, M. Renee Bellinger, and April M. H. Blakeslee. “Premature
Refutation of a Human-Mediated Marine Species Introduction: The Case History of the Marine
Snail Littorina littorea in the Northwestern Atlantic.” Biological Invasions 9: 737–50, 2007.
doi:10.1007/s10530-006-9073-x.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Littorina littorea (Mollusc).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=400&fr=1.
 64 n INVERTEBRATES (MOLLUSKS)

n Giant African Snail

Also known as: Giant African land snail
Scientific name: Achatina fulica
Family: Achatinidae
Native Range. East Africa, particularly the coastal areas of Kenya and Tanzania.
Distribution in the United States. Established only in Hawai’i, but its potential to invade
the continental United States is a source of major concern. Snails have been collected in
California, several southern states, and in Michigan, Ohio, and Wisconsin. A small but rap-
idly expanding population in Florida was eradicated in 1973.
Description. This large ter-
restrial snail can reach lengths
near 8 in. (20 cm). The cone-
shaped shell makes up roughly
half the total length. An adult
giant African snail stands about
3 in. (7–8 cm) high. The shell
is usually reddish brown with
cream and dark-brown streaks
running perpendicular to the
whorls, but a light “café au lait”
color is also common. The
shells of adults narrow toward
the apex, which is barely drawn
out. They have 7–9 whorls with
indented sutures. The semi-
elliptical opening takes up less
than half the shell length and
has a sharp, thin outer lip. The
inner lip (columella and parietal
callus) is pale blue or whitish.
This land snail is much larger
than any native species.
Related or Similar Species.
Other large African land snails
could invade the United States.
These include the world’s largest
land snail, the Ghana tiger snail
(Achatina achatina); margies or
West African land snail (Archa-
chatina marginata); and the
Nigerian land snail (Limicolaria
aurora). None of these have been
Top: The giant African snail comes from coastal regions of East Africa.
Bottom: The giant African snail is an invasive species only in Hawai’i at
found free-living in the United
this time, but it is frequently smuggled into other parts of the United States to date.
States, risking establishment of new populations elsewhere. Introduction History. Twice,
giant African snails were illegally
 GIANT AFRICAN SNAIL n 65

A. Darker forms of the giant African snail infest this tree. (David G. Robinson, USDA APHIS PPQ, Bugwood.org.)
B. This large terrestrial snail can reach lengths of 8 in., making it much larger than any native snail species.
(Forest and Kim Starr.)

imported into hatcheries on Maui, Hawai’i, in 1936, once by mail and once brought in lug-
gage from Japan. It was bred there and, shortly afterward, on O’ahu. By the time they were
discovered by authorities, escapees had established large colonies on both islands. In 1958
the snails appeared on Kaua’i and Hawai’i. By 1963, they had been discovered on Moloka’i
and possibly Lana’i.
Between 1948 and 1958, snails were intercepted by quarantine officials on cargo coming
into San Pedro and other California ports at least 50 times. However, children returning
from visits to Hawai’i were more successful in smuggling the snails into Arizona (Mesa)
and Florida (Miami) in 1958 and 1966, respectively. The contraband was released outside
when discovered by family members. In Arizona, the snails were quickly eradicated, but
the case in Florida became a cautionary tale that still argues against allowing this species any-
where in the United States. The Miami family decided to keep the snails as pets and set them
out in the family garden. Within seven years, more than 18,000 giant African snails occu-
pied 42 city blocks in Miami. Another infestation was reported north of the city in
Hollywood. It took 10 years and a million dollars for Florida to eradicate them, mostly by
hand collection.
Snails still arrive at U.S. ports as part of an illegal pet trade and unintentionally in ship-
ping containers and on plants imported from Hawai’i. Snails appeared in the Midwestern
states of Michigan, Ohio, and Wisconsin as part of the pet and shell trades and in response
to requests from teachers looking for classroom animals and unaware of the threats posed to
agriculture, horticulture, and possibly human health by these attractive, easy-to-care-for gas-
tropods. Local dispersal of eggs, and snails if populations were established, would be facili-
tated by the movement of garden waste, rubbish, construction materials, vehicles, and other
equipment away from contaminated areas. The snails themselves are capable of moving 150
ft. (50 m) a night. It only takes one gravid female to start a new population.
Habitat. Native to humid tropical forests, the giant African snail is highly adaptable and
could survive in temperate climates, even those with snow. It can also withstand drier envi-
ronments because it can seal itself inside the shell and aestivate in loose soil during
66 n INVERTEBRATES (MOLLUSKS)

unfavorable conditions. It is active at temperatures of 48–84°F (9–29°C); at lower tempera-

tures, it becomes dormant. In the tropics, this land snail inhabits disturbed forests and forest
edge, shrublands, and wetlands and has invaded agricultural areas, gardens, and urban
areas. It is nocturnal; sunlight can be lethal.
Diet. Giant African snails are herbivores and detritivores and reportedly consume some
500 different types of medicinal and ornamental plants as well as food crops, including such
temperate-zone species as beans, cucumbers, melons, peanuts, and peas. Young snails with
shells up to an inch (3 cm) long are almost completely vegetarian; older individuals increas-
ingly feed on detritus and decaying plant material, although they still feed on live plants.
Life History. The giant African snail is hermaphroditic, each individual possessing both
male and female reproductive organs, but must breed with other individuals to produce fer-
tile eggs. Snails lay their first eggs at six months and are able to produce eggs for another
400 days, with as many as three clutches a year. During their first year, they may lay 100
eggs, and in their second year, up to 500. After that time, fecundity decreases. The usual life-
span of a snail is 5–6 years.
The large eggs are about 0.2 in. (4.5–5.5 mm) in diameter and deposited in nests exca-
vated in the soil or under leaves and loose stones. They hatch from a few hours to 17 days
after laying, depending upon temperature. They require temperatures above 59°F (15°C).
The hatchlings remain below ground for 5–15 days, where they consume their egg shells
and organic detritus. They remain close to the nest during this time, but within two months
will establish a new home range and begin feeding aboveground on green plants at night.
Impacts. The greatest threat posed by giant African snails is to commercial agriculture
because they are capable of devouring tree crops, ornamentals, and vegetables. Florida’s
tropical plant industry would be particularly vulnerable. The snails could also transmit para-
sites to humans. They are carriers for rat lungworm (Angiostrongylus cantonensis), a nematode
that produces eosinophilic meningitis in humans. In natural island habitats, such as in
Hawai’i, they may compete with native snails.
Because of its voracious appetite and importation into many tropical islands in the Pacific
and Caribbean as well as Southeast Asia, these snails are considered the world’s worst pest
snail and have been nominated by IUCN as among 100 of the world’s worst invaders.
Management. In Hawai’i and Florida, a number of biocontrol measures were tried, the
more successful ones involving the importation of carnivorous snails from various parts of
the world, including Euglandina rosea and Gonaxis quadrilateralis. These seemed to depress
recruitment in giant African snails but did not eliminate them altogether. Success in
Florida came with labor-intensive and thus expensive baiting and hand collection. (In other
parts of the world, pesticides are used to control populations.)

Selected References
Davis, C. J., and G. D. Butler Jr. “Introduced Enemies of the Giant African Snail, Achatina fulica
Bowdich, in Hawaii (Pulmonata: Achatinidae).” Proceedings, Hawaiian Entomological Society, 60(3):
377–89, 1964. Available online at http://scholarspace.manoa.hawaii.edu/bitstream/10125/10889/
1/18_377-390.pdf.
IUCN/SSC Invasive Species Specialist Group (ISSG). “Achatina fulica (Mollusc).” ISSG Global Invasive
Species Database, 2006. http://www.issg.org/database/species/ecology.asp?si=64&fr=1&sts=sss.
Robinson, D. G. “Identity: Achatina fulica Bowdich, 1882.” U.S. Department of Agriculture, Animal and
Plant Health Inspection Service, 2002. http://www.aphis.usda.gov/plant_health/plant_pest_info/
gas/downloads/achatinafulica.pdf.
 GOLDEN APPLE SNAIL n 67

Stokes, Heather. “Giant (East) African Snail (Achatina fulica).” Introduced Species Summary Project.
Columbia University, 2006. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/
inv_spp_summ/Achatina_fulica.htm.

n Golden Apple Snail

Also known as: Channeled apple snail, applesnail
Scientific name: Pomacea canaliculata
Family: Ampullariidae
Native Range. South Amer-
ica, from temperate Argentina
into the Amazon basin. It
occurs in Argentina, Paraguay,
and Uruguay in the La Plata
basin and in Brazil and Bolivia
in the Amazon basin.
Distribution in the United
States. Established on Maui,
Kaua’i, O’ahu, and Hawai’i
islands in Hawai’i; a couple of
isolated sites in southern
California; northern Florida
near Jacksonville; a three-
county area in southeastern
Texas at the heart of the state’s
rice-growing region; and a site
near Fort Worth in northern
Texas. It has been reported
from Mobile, Alabama; Yuma,
Arizona; Georgia; Louisiana;
North Carolina; and Oklahoma.
Description. This freshwater
snail is fairly large; its globular
shell measures 2.75–3.6 in.
(7.0–9.0 cm) high. Deeply
indented sutures or channels
separate one whorl from the
next. The apical spire pro-
gresses in steps to a prominent
point on top; the spire has
nearly flat shoulders. Shell color Top: The golden apple snail is native to the Amazon and La Plata basins of
in wild forms is olive brown to South America. Bottom: Golden apple snails are established on Hawai’i
yellowish brown, often with and in isolated locations of southern California and northeastern
dark bands; those bred in cap- Florida. They have been found not far from Texas’s rice-growing region,
tivity may have yellow and where they would pose a major threat to the crop. (Adapted from
green shells. The large opening USGS. “Pomacea canaliculata.” USGS Nonindigenous Aquatic Species
(aperture) may be round or Database, Gainesville, FL, 2010. http://nas.er.usgs.gov/queries/
FactSheet.asp?speciesID=980.)
 68 n INVERTEBRATES (MOLLUSKS)

A. The shell of this immature golden apple snail shows the indented sutures between whorls and the flat-
shouldered apex that distinguish this species. B. The large apertures are visible on specimens in this group of
adult snails. (Susan Ellis, Bugwood.org.)

oval. The thick, retractable operculum has a concentric structure with the nucleus near the
center. Eggs are bright pink.
Related or Similar Species. The taxonomy of apple snails is not completely resolved, and
it may be that a complex of closely related species exists. Several common and scientific
names have been used in the literature, and misidentification of invaders has added to the
general confusion. In Florida, there are four other apple snails, one of which, the Florida
apple snail (P. paludosa) is native to peninsular Florida. It has a distinctively low, rounded
shell spike and is much smaller (shell height 1.6–2.75 in. [4.0–7.0 cm]) than the golden
apple snail. The eggs of Florida apple snails are white and relatively large. Genetic studies
reveal that the most common introduced apple snail in that state is the island apple snail,
P. insularum, and not P. canaliculata as previously thought. The two are very similar in canal-
iculata appearance.
Introduction History. Golden apple snails were brought to Southeast Asia in 1980 as a
potential food source for local peoples and possibly an export to the gourmet markets of
Asia. They were taken to Hawai’i from the Philippines for similar reasons in 1989.
Elsewhere in the United States, they entered as part of the pet and aquarium trade, of which
they became part in the 1950s. It is likely that infestations in the continental United States
are the result of escapes or releases of pet snails. The snail (or a close relative) was in Palm
Beach County, Florida, in 1978 and in three other central Florida counties by the late
1990s. A small population existed in a reservoir in Rockingham County, North Carolina in
1993, but did not become established. Golden apple snails were found to be established
in Lake Miramar in San Diego County and in a small pond at the Norton Simon Museum
in Pasadena, California, in 1997. Live snails and egg masses were retrieved from a canal near
the Salton Sea in Riverside County, California, in 2001. Texas Parks and Wildlife discovered
the snail in a rice irrigation canal near Houston in 2001, and surveys revealed their presence
in canals and bayous in three counties in the southeastern part of that state. Flooding result-
ing from Tropical Storm Alison in June of that year probably dispersed them to other sites in
the area.
 GOLDEN APPLE SNAIL n 69

Habitat. Native to lakes and swamps, in the United States, the golden apple snail is found
in urban ponds, drainage and irrigation ditches, natural streams, bayous, wetlands, and rice
and taro fields, which are regularly flooded. It feeds at night and spends the day submerged
among aquatic plants. This apple snail withstands periods of drought by burrowing into the
bottom and retracting its operculum to seal the shell. They may aestivate as long as five
months and can also hibernate for long periods of time during cold weather.
Diet. Golden apple snails are herbivores and consume most types of aquatic vegetation.
They show a strong preference for taro and rice. Unfortunately, they will not feed on two
invasive aquatic plants, hydrilla (See Volume 2, Aquatic Plants, Hydrilla) and waterhyacinth
(See Volume 2, Aquatic Plants, Waterhyacinth).
Life History. Golden apple snails are sexually mature when they are about 1 in. (2.5 cm)
in size or between three months and two years of age. Females crawl out of the water at night
and deposit clusters of pink eggs on just about any solid surface protruding above the water-
line. The eggs are 0.09–0.14 in. (2.2–3.5 mm) in diameter, and each clutch may contain
200–600 eggs or more. Depending upon ambient temperature, the eggs will hatch in
7–15 days. A female will produce a new clutch every few weeks and can breed throughout
the year, although reproduction is depressed in cooler months. Golden apple snails will live
about four years.
Impacts. The golden apple snail is a major agricultural pest in Hawai’i, where it attacks
taro and rice. They feed on the corm and create holes that give access to bacteria and other
plant pathogens, either killing the plant or greatly reducing its yield. They also consume
young shoots of taro, rice, and other water plants. Elsewhere in the United States, they have
not become a problem yet because they are generally in nonagricultural areas or, as near
the Salton Sea, in areas where the crops grown are not those likely to be attacked by these
gastropods. Only the populations in southeastern Texas, close to rice fields, are cause for
concern and have prompted more widespread recognition of the potential harm caused by
these invaders. So far, crop damage has not been observed in Texas. (It should be noted that
golden apple snails have been very destructive of rice and taro crops in Southeast Asia. For
this reason, they have been nominated by the IUCN as one of 100 of the world’s worst
invasive species.)
In Florida, there is concern that exotic apple snails could outcompete the native apple
snail, P. paladosa, which is the primary food of the rare and endangered Everglades Kite
(Rostrhamus sociabilis).
Golden apple snails can transmit rat lungworm (Angiostrongylus cantonensis) if they are
improperly cooked, causing severe headache, fever, and even death in people who eat them.
Management. In Southeast Asia, where the golden apple snail is such a threat to agricul-
ture, several methods of biological control have been attempted with little success. Among
the predators brought in to eat snails are ducks, carp, Nile tilapia, and red ants, which eat
snail eggs. Toxic plants grown in rice paddies or left to float on the surface did kill the apple
snails, and control of water levels helped. Laborious hand-picking appears to be the most
effective means of control, but eradication is nearly impossible. Blocking new introductions
is the best way to prevent the spread of this pest.

Selected References
Cowie, R. H. “Ecology of Pomacea canaliculata.” ISSG Global Invasive Species Database, 2004. http://
www.issg.org/database/species/ecology.asp?si=135&fr=1&sts=.
70 n INVERTEBRATES (MOLLUSKS)

Ghesquiere, Stijn A. I. “Pomacea (pomacea) canaliculata (Lamarck, 1819).” The Apple Snail Website,
n.d. http://www.applesnail.net/content/species/pomacea_canaliculata.htm.
Howells, Robert G., and James W. Smith. “Status of the Applesnail Pomacea canaliculata in the United
States.” The Seventh International Congress on Medical and Applied Malacology (7th ICMAM) Los
Baños, Laguna, SEAMEO Regional Center for Graduate Study and Research in Agriculture
(SEARCA), Philippines, 2003. http://www.applesnail.net/pestalert/conferences/icam07/country
_report%20_usa.pdf.
Mohan, Nalini. “Apple Snail (Pomacea canaliculata).” Introduced Species Summary Project. Columbia
University, 2002. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/
Pomacea_canaliculata.html.
Strange, Lionel A., and Thomas R. Fasulo. “Apple Snails of Florida.” University of Florida, 2004. http://
entnemdept.ufl.edu/creatures/misc/gastro/apple_snails.htm.

n Naval Shipworm
Also known as: Atlantic shipworm, great shipworm
Scientific name: Teredo navalis
Family: Teredinidae
Native Range. Naval shipworms have been transported around the world in the hulls of
wooden sailing ships for centuries. It is difficult to know, therefore, just what their native
range is. The Atlantic coast of Europe from Iberia to Scandinavia is the most likely place of
origin. Today the species is cosmopolitan, found in tropical and subtropical waters of the
Atlantic and Pacific oceans in both the Northern and Southern hemispheres.
Distribution in the United States. Naval shipworms are found in all coastal regions of the
United States.
Description. Although called a worm, this animal is mollusk, a somewhat bizarrely
formed bivalve adapted to boring into and living in wood. The reddish body is long and
wormlike and topped by a very small shell that serves to chip away wood. Most contempo-
rary information on them comes from the Baltic Sea, where they regularly attain lengths of
8–12 in. (20–30 cm) and a width of 0.4–0.8 in. (1–2 cm). In tropical waters, they reportedly
become nearly 20 in. (50 cm) long. The shell is ridged and covers but a small part of the
body; it is usually 0.5–0.8 in. (12–20 mm) in length.
Frequently, the burrows that shipworms dig in submerged wood are the chief indication
that the animals are present. Each burrow is a long cylindrical tube lined with a calcareous
coating secreted by the shipworm. It is blocked near the entrance by a pair of calcareous
structures or “pallets” that are paddle-shaped. Burrows excavated by adults can be as much
as 3 ft. (1 m) long, but larvae with diameters of 0.04 in. (1 mm) also bore into wood. The
shipworm extends two siphons from its anterior end through a small opening between the
pallets for feeding, respiration, and excretion of wastes.
Related or Similar Species. There are native species of shipworm in several genera,
including six species in the genus Teredo, found along the Atlantic coast; and others that
are introduced or of unknown origin.
Introduction History. The naval shipworm was first reported in the United States in
1839, when it was detected in the sheathing of a foreign wooden ship in Massachusetts
Bay. Within 100 years, it was common all along the Atlantic coast north of the bay. Naval
shipworms showed up in Long Island Sound in 1869 in the hull of a sailing ship, and within
a few decades, they were common in New York Harbor. The first naval shipworms were
 NAVAL SHIPWORM n 71

detected in Chesapeake Bay in

1878, but it remained rare.
Later, it was collected from
North Carolina south to
Florida and along Texas’s Gulf
coast, as well as in Puerto Rico.
An invasion on the Pacific
coast in San Francisco Bay
occurred in 1913 with drastic
results (see under Impacts).
Today, the naval shipworm is
less abundant as a result of
many fewer wooden-hulled
ships and the widespread use
of chemically treated timbers in
waterfront construction.
Habitat. Shipworms live in
timbers submerged in seawater,
including the hulls of wooden
ships and pilings, piers, and
other wooden waterfront struc-
tures. They tolerate a wide range
of salinities (5–45 ppt), so they
also thrive in estuaries. Larvae
probably die in salinities less
than 5 ppt. Shipworms repro-
duce at water temperatures rang-
ing from 52–86°F (11–30°C),
but are known to survive in
water temperatures as low as
33°F (0.7°C). They are found in
tropical seas as well as cool tem- Top: Naval shipworms are found around the world as a consequence of
perate waters. They survive up global trade and explorations, especially in the days of wooden-hulled
to six weeks in oxygen-poor sit- ships. Their origins are obscured but are likely to have been in the
uations by sealing themselves Atlantic waters off Europe. Bottom: Shipworms are found in all coastal
into their burrow and metaboliz- waters of the United States.
ing stored glycogen.
Diet. Shipworms feed almost exclusively on wood with the aid of symbiotic bacteria
that digest cellulose and fix nitrogen. The bacteria reside in special cells in the gills. The
specially sculptured shell rasps away wood that is then transported to the mouth by cilia.
Shipworms may also be filter-feeders of plankton, drawing in water through the incurrent
siphon.
Life History. Adult shipworms occur as separate males and females. Spawning apparently
depends on temperature. It reportedly occurs from April to September in New Jersey and from
May to October at Woods Hole, Massachusetts. Males release sperm into the water column.
Other individuals take them in through their incurrent siphons, and fertilization
 72 n INVERTEBRATES (MOLLUSKS)

occurs internally in the chamber

housing the gills. Larvae are
brooded in the gills until the
velum (a ciliated organ that
provides locomotion) and a
straight-hinged shell form. Then
the larvae are released into the
water column, where they begin
a planktonic stage lasting up to
four weeks. During this phase,
they form siphons, gills, and an
obvious foot. The tiny (0.04 in.
or 1 mm) final-stage larvae
detect wood chemically and
swim toward it, attaching them-
selves with byssal threads. It is
not known how the soft-shelled
larvae bore into wood; maternal
enzymes may aid in softening
the surface. The settled larvae
rapidly metamorphose into
This peculiar bivalve has a long wormlike body and a tiny shell used to juvenile shipworms. They be-
bore into wood. (U.S. Geological Survey.) come sexually mature adults six
weeks after settlement.
Impacts. In the past, this animal was so abundant in harbors around the world that they
determined the lifespan of a wooden ship. It was reported in Wales that submerged ship timbers
were destroyed in eight years. The presence of shipworms necessitated covering ships’ hulls in
tar or cladding them in copper. The outbreak in San Francisco Bay from 1919 to 1921 destroyed
untold numbers of wharves, piers, and other waterfront structures. On average, a major struc-
ture was lost every two weeks with enormous economic consequences. In the Baltic Sea area,
there is concern today that thousands of ancient shipwrecks lying on the seafloor and of great
archeological value could be destroyed if naval shipworms invaded the sites.
If and how this species affects native or other introduced shipworms has not been stud-
ied, but competition for food is possible.
Management. The naval shipworm is less of a problem today than in the past as a result
of many fewer wooden-hulled ships and the widespread use of chemically treated timbers in
waterfront construction. However, in the absence of these defenses, it will still destroy
wooden structures below water.

Selected References
Masterson, J. “Teredo navalis.” Smithsonian Marine Station, 2007. http://www.sms.si.edu/irlSpec/
Teredo_navalis.htm.
NIMPIS. “Teredo navalis Species Summary.” National Introduced Marine Pest Information System,
edited by C. L. Hewitt, R. B. Martin, C. Sliwa, F. R. McEnnulty, N. E. Murphy, T. Jones, and S.
Cooper, 2002. Available at http://www.frammandearter.se/0/2english/pdf/Teredo_navalis.pdf.
“Shipworm (Teredo navalis).” Wreck Protect, n.d. http://wreckprotect.eu/fileadmin/site_upload/
wreck_protect/pdf/shipwormspdfnew.pdf.
 NEW ZEALAND MUD SNAIL n 73

n New Zealand Mud Snail

Also known as: Jenkin’s spire shell
Scientific name: Potamopyrgus antipodarum
Synonyms: Hydrobia jenkinsi, Potamopyrgus jenkinsi
Family: Hydrobiidae
Native Range. Freshwater streams and lakes and brackish habitats in New Zealand and
adjacent islands.
Distribution in the United States. Populations are found in all western states except New
Mexico and in four of the Great Lakes: Ontario, Michigan Erie, Michigan and Superior.
Description. This very small aquatic snail typically measures 0.25 in (4–5 mm) in the
western United States, although
it may be twice that large in its
native range. The cone-shaped
shell has right-handed coils and
contains 5–6 whorls separated
by deep indentation. A retract-
able plate (operculum) covers
the oval opening to the shell
and protects the animal when it
retreats into its shell. Shell color
ranges from gray to light or dark
brown.
Different morphs appear
among introduced populations.
Many from the Great Lakes and
western states have a slight keel
in the middle of each whorl,
whereas others from the Great
Lakes have spines on the who-
rls. One morph found in the
West is wider than usual and
so pale and transparent that
internal structures are visible.
Related or Similar Species.
The New Zealand mud snail
could be confused with a num-
ber of native snails in the western
United States and may be confi-
dently identified only by experts.
The shell is longer and narrower
than most native snails of the
same family and has more
whorls. No native western snails Top: The New Zealand mud snail is native to fresh and brackish water
have keels. Most native species habitats in New Zealand. Bottom: New Zealand mud snails have
have fewer whorls than the New invaded almost all western states and four of the Great Lakes. (Adapted
Zealand land snail. (The Aquatic from Benson and Kipp 2009.)
 74 n INVERTEBRATES (MOLLUSKS)

A. The retractable plate or operculum and deep indentations separating whorls are clearly visible on this
common form of the New Zealand mud snail. B. Spines occur on the whorls of many individuals from the
Great Lakes region. C. The living animal (D. L. Gustafson, Montana State University.)

Nuisance Species Research Program of the U.S. Army Corps of Engineers presents comparative
photos of native snails at http://el.erdc.usace.army.mil/ansrp/potamopyrgus_antipodarum.pdf.)
Introduction History. The exact means of introduction to U.S. waters is unknown, but
the snails likely arrived with shipments of fish eggs and live game fish from New Zealand
to some western states. Introduction to the Great Lakes, on the other hand, probably
occurred with the discharge of ballast water by ships coming from Europe. First records of
the snail came from the Snake River, Idaho, in 1987. Populations were reported near the
mouth of the Columbia River in Oregon in 1997, and they were established in the lower
Columbia in Washington by 2002. First reports from California come from the Owens
 NEW ZEALAND MUD SNAIL n 75

River; it is now widespread in that state. In 2002, the snails were in the Colorado River in
northern Arizona; in 2004, they were discovered in a small creek near Boulder, Colorado.
An established population existed in Lake Ontario in 1991 and in Lake Erie by 2005.
They were also found in Duluth harbor on Lake Superior in 2005.
The tiny New Zealand mud snail may be inadvertently transferred from one body of water
to another on boats, boots, waders, clothing, and other recreational gear. They may possibly
be spread by animals, because they can pass unharmed through the guts of fish and birds.
Even cattle wading in streams and ponds could transport these mollusks, since they are able
to withstand desiccation for a short period time by withdrawing into their shells behind a
closed operculum.
Within a river or lake system, the snails may float downstream or become attached to
floating mats of algae. They have also been shown to be capable of moving upstream under
their own volition.
Since most New Zealand mud snails reproduce asexually, it takes only one snail to begin
a new population. They spread rapidly and now have access to the eastern United States via
the Great Lakes.
Habitat. The New Zealand mud snail is found in a variety of freshwater and brackish-
water habitats at depths up to 150 ft. (45 m). It inhabits streams, lakes, estuaries, lagoons,
canals, ditches, and water tanks on just about any type of substrate, from mud and sand to
concrete and within many types of vegetation. It prefers the littoral zone of lakes and slow-
moving streams, but because it can burrow into sediments. It tolerates fast-moving water.
These snails thrive in salinities of 0–15 ppt and for short periods of time will tolerate salin-
ities near that of seawater. They also tolerate a wide range of temperatures from 32° to 110°F
(0–34°C). They are preadapted to live in degraded water systems with high nutrient content
and/or heavy siltation.
Diet. This small gastropod is a scraper/grazer that feeds nocturnally on plant and animal
detritus and on living algae and other organisms attached to the substrate (the periphyton).
Diatoms are prominent in its diet.
Life History. Most New Zealand mud snails in the United States are members of all
female populations, meaning that individuals in a population are clones and genetically
identical. Males are not needed for reproduction; indeed, the female is born with 20–120
developing embryos already in her reproductive system. Sexual reproduction does occur,
but sexual males and females are very rare. Females generally release live young in summer
and autumn. Average lifespan is about one year. Sexual males and females are mature when
about 0.1 in. (3 mm) in size.
New Zealand mud snails have a high reproductive potential and it is not unusual for pop-
ulation densities to exceed 5,290/ft.2 (100,000/m2). In the Madison River near Yellowstone
National Park in Montana, densities three times greater are reported.
Impacts. The New Zealand mud snail has had negative impacts in other parts of the world
and is expected to have them in the United States as well. Concerns center on its potential to
decrease populations of native herbivorous invertebrates—including native snails—through
competition for food and space and to reduce the periphyton cover that serves as nourish-
ment and substrate for some aquatic organisms. Evidence from Montana suggests a decrease
in larval mayflies, stoneflies, caddisflies, and black flies when the snails become abundant.
These insects are important food for native trout and other fish, and their loss could
negatively impact the important recreational fishing industry in many western states.
In the Snake River, Idaho, five native mollusks (Utah valvata snail [Valvata utahensis],
Idaho springsnail [Pyrgulopsis idahoensis], Snake River physa [Haitia natricina], Banbury
76 n INVERTEBRATES (MOLLUSKS)

Springs lanx [Lanx sp.], and Bliss Rapids snail [Taylorconcha serpenticola]) are listed as threat-
ened or endangered, possibly as a consequence of competition from New Zealand mud snails.
A decrease in the periphyton would alter the physical environment and affect ecosystem
processes beginning with primary production. Very dense populations of the mud snails
could clog intake filters at facilities withdrawing lake or river water.
Management. Little can be done to eradicate established populations of the New Zealand
mud snail. Management is focused on the prevention of its spread to additional bodies of
water. Boats and fishing and other gear should be scrubbed before leaving an infested body
of water and thoroughly dried for at least 24 hours or treated with heat ( > 85°F [30°C])
before being used elsewhere.

Selected References
Benson, A. J., and R. M. Kipp. “Potamopyrgus antipodarum.” Nonindigenous Aquatic Species Fact Sheet,
U.S. Geological Survey, 2009. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=1008.
Crosier, Dani, and Dan Malloy. “New Zealand Mudsnail (Potamopyrgus antipodarum).” United States
Federal Aquatic Nuisance Species Task Force, 2005. http://www.anstaskforce.gov/spoc/nzms.php.
Crosier, Danielle, and Daniel P. Molloy. “New Zealand Mudsnail–Potamopyrgus antipodarum.” Aquatic
Nuisance Species Research Program, U.S. Army Corps of Engineers, n.d. http://el.erdc.usace
.army.mil/ansrp/potamopyrgus_antipodarum.pdf.
Richards, David C., Billie L. Kerans, and Daniel L. Gustafson. “New Zealand Mudsnail in the Western
USA.” Montana State University-Bozeman, 2004. http://www.esg.montana.edu/aim/mollusca/nzms/.

n Quagga Mussel
Scientific name: Dreissena rostriformis bugensis
Family: Dreissenidae
Native Range. Dnieper and Bug rivers and Dnieper-Bug Estuary, Ukraine.
Distribution in the United States. Established in all of the Great Lakes. Also found in
inland bodies of water in Arizona, California, Colorado, Illinois, Iowa, Kentucky,
Michigan, Minnesota, Missouri, Nevada, New York, Ohio, and Pennsylvania.
Description. The quagga mussel is a small freshwater bivalve with asymmetrical valves and
a convex ventral side that prevents it from maintaining a stable upright position. The byssal
groove on the ventral side is small and positioned near the hinge, and the angle between the
ventral and dorsal surfaces is rounded. There are usually dark concentric bands on the shell
that fade near the hinge, but shell patterns are quite variable and the bands may be black,
cream, or white. In Lake Erie is a type that is completely white. Two distinct forms are found
in the Great Lakes. One type, the so-called “epilimnetic” or shallow water form, has a high flat
shell; the other, the “profunda” or deep, cold-water form, has a somewhat elongated, globular
shape. The epilimnetic type attaches to hard surfaces with hairlike byssal threads and forms
dense colonies or druses; they may attach to the shells of other quagga mussels and create
mats 4–12 in. (10–30 cm) thick that encrust or clog pipes and other manmade features.
The profunda type, while it can attach to objects with byssal threads, may also partially bury
itself in soft sediments and extend a long incurrent siphon above its shell to draw in sus-
pended organic matter. Quagga mussel shells grow to lengths of about 1.5 in. (4 cm).
Related or Similar Species. Quagga mussels are quite similar to the closely related zebra
mussel (Dreissena polymorpha; see Mollusks, Zebra Mussel). In fact, the two were not
immediately identified as separate species. The quagga mussel shell has a convex ventral side
 QUAGGA MUSSEL n 77

in contrast to the flat or convex

ventral side of zebra mussels.
The quagga mussel shell is
rounded, whereas that of the
zebra mussel is more triangular.
The zebra mussel also is distinct
by virtue of bilateral symmetry
along a midventral line and a
long byssal groove in the
middle of the ventral side.
The Asian clam (Corbicula flu-
minea), another small invasive
bivalve (see Mollusks, Asian
Clam), lacks byssal threads and
has a thicker, ridged shell. It is
light-yellowish brown to dark
brown and usually is not striped.
Introduction History. The
first known occurrence of
quagga mussels in the Great
Lakes dates to September 1989,
although at the time the col-
lected specimen was believed to
be a form of zebra mussel. The
specimen was obtained near
Port Colburne, Lake Erie. In
1991, a mussel taken from the
Erie Canal was positively identi-
fied as a new species and
dubbed the quagga mussel after
a less-striped relative of the
Plains zebra that is now extinct. Top: Quagga mussels are native to the Dnieper and Bug rivers and estuary
The mussel was most likely in Ukraine. Bottom: Quagga mussels are established in all of the Great
introduced in discharged ballast Lakes and in many inland bodies of water across the lower 48 states.
water from transoceanic freight- (Adapted from Benson, Richerson, and Maynard 2010.)
ers. The mussel spread through
the lower Great Lakes and reached Duluth Harbor, Lake Superior, by 2005. Quagga mussels
were discovered in the Mississippi River between St. Louis, Missouri, and Alton, Illinois, in
1995. Larvae can float downstream in rivers or on currents in lakes. They can leapfrog from
lake to lake attached to boats and boat trailers or in anything that holds water. Adults are able
to withstand exposure to air for 3–5 days.
In January 2007, quagga mussels showed up in the major impoundments on the
Colorado River, first in Lake Mead, near Boulder City, Nevada, and then in Lakes Mohave
and Havasu along the lower Colorado River. The same year, they entered the Colorado
River Aqueduct, which diverts water from the Colorado River to southern California. Since
then, they have been found in 15 reservoirs in southern California and also in six reservoirs
in the state of Colorado. They are also in the Nevada State Fish Hatchery on Lake Mead and
at Willow Beach National Fish Hatchery, just downstream from Hoover Dam on the
 78 n INVERTEBRATES (MOLLUSKS)

A. Quagga mussel shells are asymmetrical; pattern and color vary. (U.S. Geological Survey.) B. The living animal.
(M. Quigley, GLERL.)

Colorado, which raises trout and threatened native fishes to stock Lakes Mead and Mohave
and to restock the Colorado River.
Habitat. Quagga mussels are found in the upper reaches of estuaries, in lakes and reser-
voirs and connecting waterways. They do best in freshwater with salinities less than 1 ppt,
but can still reproduce in brackish water with salinities near 3 ppt. (They die in salinities
greater than 6 ppt). Quaggas apparently have a broader range of temperature tolerance than
zebra mussels and hence inhabit a wider range of water depths. As a consequence they
occupy both shallow warm waters and deeper cold-water habitats. In North America they
are colonizing waters where the temperature remains between 39° and 48°F (4–9°C) all
year. Adults attach to natural hard surfaces of rock, shell, and wood or to large aquatic plants
as well as to manmade structures of concrete, metal, nylon, fiberglass, and wood. Some
morphs also bury themselves in soft lake bottoms.
Diet. Filter-feeders, quagga mussels ingest food particles by pulling water into the shell
cavity with cilia. The water passes into an incurrent siphon where phytoplankters, zoo-
plankters, and other suspended organic matter are extracted. Indigestible particles are
cemented together with mucus and ejected from the incurrent siphon as pseudofeces; water
is discharged from an excurrent siphon.
Life History. Individuals are either male or female and reproduce externally when eggs
and sperm are released into the water column. Microscopic embryos develop into larvae in
a few days. The larvae soon sport tiny bivalve shells and drift as part of the plankton for
3–4 weeks. When suitable substrates are found, juveniles settle and attach themselves with
byssal threads. Adults are also sessile, but able to detach and move to new sites.
Spawning usually peaks in spring and autumn in the Great Lakes, but elsewhere may
occur throughout the year. In Lake Havasu on the Colorado River, quaggas reportedly
spawn six times a year. The average lifespan is five years.
Impacts. Major alteration of aquatic ecosystems can result from heavy infestations of
quagga mussels. Their removal of much of the phytoplankton and other suspended organic
matter from the water column reduces the food supply of zooplankters and thereby affects
the rest of the food chain. Their production of large amounts of pseudofeces transfers much
of the energy in the system from the upper parts of the water column to benthic habitats.
Bottom-feeding fish increase at the expense of plankton-feeders. In addition, when large
accumulations of pseudofeces decompose, oxygen is withdrawn from the water. In the cen-
tral basin of Lake Erie, dead zones of oxygen-depleted water have appeared for decades in
late summer at depths greater than 40 ft. (12 m). Even with the general amelioration of
severe nutrient pollution problems that affected the lake in the 1970s, these dead zones
 VEINED RAPA WHELK n 79

persist. One hypothesis is that changes in nutrient cycles brought on by the introduction of
quagga and zebra mussels may be a cause.
The reduction of planktonic green algae increases water clarity and allows greater
growth of macrophytes in shallow waters, changing the species composition of the plant
community and the physical structure of the aquatic environment as well as available food
sources. Emergent and floating plants become nursery areas for a variety of aquatic
organisms.
Indications are that quagga mussels outcompete their close relative the zebra mussel and
are replacing them in shallow waters. They also foul native mollusks and other hard-shelled
invertebrates, but their presence in American waters is too brief for any impacts to be
realized.
Mussels bioaccumulate pollutants in the water. These pollutants collect in pseudofeces
and from there can pass up the food chains to game fish and other wildlife. The bivalves
themselves are consumed by some crayfish, fish, and diving ducks, offering another path-
way for concentrated toxins to move into food chains.
Quagga mussels biofoul organisms and clog the pipes and screens of water intake struc-
tures. This can reduce pumping capacity and damage equipment and cost industries and
communities millions of dollars to mitigate. Fouling of water-intake screens and sluice gates
has rapidly become a major problem along the Colorado River, which supplies drinking and
irrigation water via long aqueducts to the major cities and agricultural regions of southern
California, southern Nevada, and southern Arizona.
Management. A variety of control methods are available, but so far none has proved both
effective and environmentally sound. Biological controls that would target quagga (and
zebra) mussels and interrupt their reproductive cycle or interfere with the settling of the lar-
vae are being researched.

Selected References
Benson, A. J., M. M. Richerson, and E. Maynard. “Dreissena rostriformis bugensis.” USGS Nonindigenous
Aquatic Species Database, Gainesville, FL, 2008. http://nas.er.usgs.gov/queries/FactSheet.asp
?speciesID=95.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Dreissena bugensis (Mollusc).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?si=918&fr=1&sts=sss.
Richerson, Myriah. “Dreissena species FAQs, a closer look.” USGS, 2009. http://fl.biology.usgs.gov/
Nonindigenous_Species/Zebra_mussel_FAQs/Dreissena_FAQs/dreissena_faqs.html.

n Veined Rapa Whelk

Also known as: Asian rapa whelk
Scientific name: Rapana venosa
Synonym: Rapana thomasiana
Family: Muricidae
Native Range. Estuarine and marine waters of the western Pacific Ocean. It is known
from Vladivostok, Russia, south to Taiwan in the Sea of Japan, the Yellow Sea, the Bohai
Sea, and the East China Sea. Genetic studies of whelks from the Chesapeake Bay suggest they
arrived in American waters via the Black Sea, to which they were introduced in the 1940s.
 80 n INVERTEBRATES (MOLLUSKS)

Distribution in the United

States. Established in Chesa-
peake Bay from the mouth of
the Rappahannock River south
to the bay’s entrance. Based on
water temperatures similar to
that of their native distribution
area, and the existence of a high
volume of coastwise trade out of
Hampton Roads, their potential
range extends from Cape Cod,
Massachusetts, to Charleston,
South Carolina.
Description. This large
marine snail is most readily iden-
tified by the bright orange color
of the inside of its shell, visible
at the large, oval aperture that
opens on the right side of the
shell. The heavy shell is globular
in shape and has a short spire,
broad flat columella, and deep
umbilicus (the hollow on the
ventral side). Smooth spiral ribs
terminate in blunt knobs adorn-
ing the shoulder, and the edge
of the outer lip is finely toothed.
The outer shell ranges from gray
to brown and bears dark brown
lines that create an interrupted
pattern over the entire shell.
Top: The veined rapa whelk is native to estuarine and marine waters from Dark veining may occur on the
southeastern Russia to Taiwan. Bottom: Veined rapa whelks have been inside of the shell. In their native
found in the Chesapeake Bay. (Adapted from International Council for range, they reach a length of 7 in.
the Exploration of the Sea 2004.) (180 mm). Adults slightly over
6.5 in. (170 mm) have been col-
lected in the Chesapeake Bay.
The egg cases of the veined rapa whelk are also distinctive, usually described as resem-
bling a yellow shag rug. They are about 1.2 in. (30 mm) high and individually attached to
the seafloor to form a mat.
Related or Similar Species. Two large native whelks inhabit the potential range of the
veined rapa whelk along the East Coast of the United States. Neither the knobbed whelk
(Busycon carica) nor the channeled whelk (Busycon canliculatum) has a broad flat columella
or the dark veining on the shell. Both native whelks are more elongated, have thinner shells,
and display less ribbing on the outside of their shells than does the veined rapa whelk. Egg
cases of the knobbed whelk are strung together in a long chain, a “mermaid’s necklace.”
Introduction History. The first scientific evidence of veined rapa whelks in Chesapeake
Bay came in 1998, when researchers from the Virginia Institute of Marine Science collected
 VEINED RAPA WHELK n 81

a specimen in Hampton Roads,

Virginia. The large whelk was
probably 10 years old, strongly
suggesting the presence of
whelks in the bay since 1988, a
fact confirmed anecdotally by
local watermen. The species
likely arrived in ballast waters
in empty coal vessels returning
from the eastern Mediterra-
nean, since Norfolk, Virginia,
is a major coal-exporting port.
Genetic studies support this
contention. The long larval
period when the whelk floats
in the water column facilitates
the long distance dispersal of
this gastropod by currents Adult veined rapa whelks found in the Chesapeake Bay have reached just
in the bay and by container over 6.5 in. long. Larger specimens occur in their native range. (George
Chernilevsky.)
ships and other vessels that take
up ballast water before heading
to ports all along the East Coast.
Habitat. Subtidal zones of coasts and estuaries. Adults prefer sandy bottoms into which
they burrow, but juveniles tend to dwell on hard substrates, including oyster reefs, and only
migrate to areas with sandy bottoms when shell length is about 2.75 in. (70 mm). In its
native range, it tolerates a wide annual range of temperatures, from 39–80ºF (4–27ºC); in
cooler climates, it may migrate to deeper and warmer water during winter. Adults and larvae
can withstand low salinities and polluted and/or oxygen-poor water.
Diet. Larvae consume phytoplankters, but juveniles and adults are carnivores. Prey
includes other shellfish that live in soft sediments (the infauna), such as clams and mussels,
and oysters. Most vulnerable in the Chesapeake Bay are native hard clams (Mercenaria merce-
naria), but mussels (Mytilus edulis), soft-shell clams (Mya arenaria), and oysters (Crassostrea
virginica) are also taken. The whelk engulfs its prey whole; and, when the bivalve opens, it
sucks out the soft body of the victim, leaving behind a clean shell with few telltale scars.
Life History. In its native range, mating occurs during winter and spring, and eggs are
laid from April to late July. Each egg case is attached by its base to a hard substrate. From
200 to 1,000 eggs are contained in each case; each mat may consist of 50- to 500-egg cases.
A female can lay several mats a year from a single mating. Initially, the egg cases are white,
but as the larvae become visible swimming inside, they turn yellow. After an incubation
period of 14–17 days (depending upon temperature), the egg cases blacken, open, and the
larvae swim free out of the top of the case to become part of the plankton. The larval stage
lasts for a variable length of time, extending as long as 80 days. Upon metamorphosis, they
settle onto attached invertebrates such as bryozoans and barnacles and become tiny, hard-
shelled, cryptically colored whelks. The young whelks grow quickly so that within three
weeks, their shell’s length approaches 0.02 in. (0.5 mm); within a year, they are more than
2.4 in. (60 mm) long. Veined rapa whelks begin to reproduce at two years of age, when they
have a shell length of about 3 in. (74 mm). The largest whelks collected from the
Chesapeake Bay are estimated to be about 10 years old.
82 n INVERTEBRATES (MOLLUSKS)

Impacts. The veined rapa whelk has a history of negative impacts in European waters,
especially in the Black Sea, where its predation was implicated in the rapid decline of native
edible bivalves and near extinction of oysters (Ostraea edulis) on the Guadata oyster bank.
Concern is therefore high that similar impacts could ensue at commercially important
fisheries in the Chesapeake Bay and elsewhere along the East Coast of the United States. In
the Chesapeake Bay, it appears that hard clams are most threatened, since oysters are
confined by disease to very low-salinity sites.
These nonnative whelks might compete with the native oyster drill (Urosalpinx cinerea), a
species struggling to recover from a massive die-off caused by an influx of freshwater related
to Hurricane Agnes in 1972. A shift in relative abundance of hermit crab species is also pos-
sible. The shells of the veined rapa whelk are well suited to house the striped hermit crab
(Clibanarius vittatus), but less so to the currently dominant flat-clawed hermit crab
(Pagurus pollicaris). The former is predator of oysters, and its increase would add to pres-
sures already affecting the bay’s remaining oyster populations.
Management. It appears unlikely that established populations can be eradicated, so the
emphasis of research is on identifying sites vulnerable to invasion and trying to prevent
the organism’s spread. Several states on both coasts of North America have issued alerts
for their citizens to be on the lookout for veined rapa whelks and report any sightings to
authorities. For a time, researchers on Chesapeake Bay offered a bounty on whelks caught
by fishermen, in part to monitor the status and spread of the whelk population and in part
to discourage fishermen from throwing the by-catch back into the water.

Selected References
Chesapeake Bay Introduced Species Database. “Rapana venosa.” NEMESIS (National Exotic Marine and
Estuarine Species Information System). Smithsonian Environmental Research Center, 2009. http://
invasions.si.edu/nemesis/CH-TAX.jsp?Species_name=Rapana%20venosa.
International Council for the Exploration of the Sea. “Alien Species Alert: Rapana venosa (Veined
Whelk),” edited by Roger Mann, Anna Occhipinti, and Juliana M. Harding. ICES Cooperative
Research Report No. 264, p. 14, 2004. http://www.ices.dk/pubs/crr/crr264/crr264.pdf.
Richerson, Myriah. “Rapana venosa.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL,
2006. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=1018.
Southeast Ecological Center. “Veined Rapa Whelk, Asian Rapa Whelk, Rapana venosa (Valenciennes,
1846) Mollusca: Gastropoda, Muricidae.” Nonindigenous Species Information Bulletin, U.S.
Geological Survey, 2009. http://fl.biology.usgs.gov/Nonindigenous_Species/Rapa_whelk/rapa
_whelk.html.

n Zebra Mussel
Scientific name: Dreissena polymorpha
Family: Dreissenidae
Native Range. Eastern Europe and western Asia in the drainage systems of the Black and
Caspian seas and the Sea of Azov. U.S. populations originate from mussels transported from
the southern limit of their range.
Distribution in the United States. Zebra mussels are now found in waterways in or bor-
dering at least 30 states in the continental United States. They are established in all of the
Great Lakes and in most navigable drainages in the Mississippi River system, including the
Missouri River in Nebraska and South Dakota. They are also established in the Hudson
 ZEBRA MUSSEL n 83

River and found in numerous

inland lakes in the Midwest.
The most recent introductions
have occurred in San Justo
Reservoir in central California,
two reservoirs in Colorado,
Lake Texoma on the Texas-
Oklahoma border, and a lake
in western Massachusetts.
Description. Zebra mussels
are small freshwater bivalves
with triangular shells. The bot-
tom of the shell where the hinge
lies is flat or concave, and the
shell margin is sharply angled.
When placed upright on a flat
surface, they are stable. Many
bear dark stripes on the shell,
but this pattern can be quite
variable and even absent. Some
will be plain and cream-
colored; others black. Fila-
ments or byssal threads extend
from the bottom of the shell
and bind them to each other or
to hard surfaces. Usually zebra
mussels occur in dense clusters
or colonies known as druses.
Shells of adults generally range
in length from 0.2–1.8 in.
(6–45 mm), with a maximum
length of about 1.9 in. (0.5 cm). Top: Zebra mussels are native to the drainage systems of the Black, Azov,
Related or Similar Species. and Caspian seas. Bottom: Zebra mussels are a scourge in at least 30 states
Two other dreissenids occur in and are established in all the Great Lakes and much of the Mississippi
North America, the native false River system. (Adapted from Benson and Raikow 2009.)
dark mussel (Mytilopsis leauco-
phaeata) and the introduced quagga mussel (Dreissena rostriformis bugensis; see Mollusks,
Quagga Mussel). Each has a convex bottom edge and a rounded shell margin, which pre-
vents it from balancing in an upright position.
Introduction History. The introduction of zebra mussels probably stems from a single
exchange of ballast water by a commercial ship entering the Great Lakes from the northern
Black Sea. The first reports came from Lake St. Clair, which connects Lake Erie and Lake
Huron, in 1988. It is likely they had been in the lake for 2–3 years before having been
detected. By 1990, the passive drifting of larval mussels and the ability of juveniles and
adults to attach to barges and other vessels had allowed them to disperse into all the Great
Lakes. In 1991, they were in the Illinois River, gateway to the Mississippi River, and within
a year they were established in the Arkansas, Cumberland, Ohio, and Tennessee rivers. They
were also in the Hudson River and the Finger Lakes of New York by 1991. Overland
 84 n INVERTEBRATES (MOLLUSKS)

A. Zebra mussels overgrow the shells of native mollusks. (Eric Engbretson.) B. A living zebra mussel is in a stable
position when placed upright on its hinge. (M. Quigley, GLERL.) C. The dense clusters in which zebra mussels
usually congregate are called druses. (Eric Engbretson.) D. Some of the variation found in shell patterns of
Dreissena polymorpha. (U.S. Geological Survey.)

dispersal has undoubtedly been facilitated by recreational boaters trailering their watercraft
from infested bodies of water to pristine waters without decontaminating hulls, trailers, live-
wells, and engines. Zebra mussels were first recorded in Virginia in a quarry pond in 2002.
Other states with first records in the twenty-first century include Kansas (Perry Lake and the
Kansas River) in 2007; California (San Justo Reservoir, San Benito County) and Colorado
(Pueblo Reservoir—an impoundment on the upper Arkansas River near Pueblo and four
reservoirs west of Denver) in 2008; and Texas and Oklahoma (Lake Texoma), and
Massachusetts (Laurel Lake, Berkshire County) in 2009.
Habitat. Zebra mussels can be found in freshwater lakes, ponds, and rivers. Usually they
occur in shallow, algae-rich water at depths of 6–30 ft. (2–9 m). They require hard sub-
strates for settling and attachment. They thus may occur on rocks; in and on metal, concrete,
or other manmade structures; on the shells of other mollusks and the exoskeletons of
 ZEBRA MUSSEL n 85

crustaceans; and even on the cases of caddisfly larvae. In soft sediments, they may attach to
pieces of shell or wood and other plant material or to stones and still be able to establish a
druse. Adults do not survive freezing temperatures and die when water temperatures are
above 75°F (24°C) for extended periods of time. They also do poorly in oxygen-poor waters.
Optimal calcium-ion concentrations for zebra mussels are 45–55 mg/l, and optimal pH for
adult growth lies between 7.4 and 8.0.
Diet. Larval, juvenile, and adult zebra mussels are filter-feeders, capturing primarily algae
and zooplankters in the water column. They also feed on bacteria and detritus.
Life History. Female mussels release eggs into the water column, where they are fertilized
by sperm released by males. Most spawning occurs between June and September, when
water temperatures are at an optimal 54–68°F (12–20°C). Three distinct periods character-
ize the life cycle of the zebra mussel. The microscopic larvae are part of the plankton and
undergo four stages when they are essentially clam-like in shape. In the final larval stage, a
larva settles on a substrate and crawls to a suitable attachment site. There it undergoes meta-
morphosis and begins its juvenile period with a small (1–3 mm), triangular shell. Gametes
become fully developed juveniles in 8–180 days; the colder the water, the slower the devel-
opment time. The adult period starts when the mussel becomes sexually mature, usually at
one year of age and a shell length of 0.3–0.35 in. (8–9 mm). A zebra mussel lives on average
2–3 years after attachment.
Impacts. Dense populations of zebra mussels have the potential to alter freshwater eco-
systems and change the physical environment by eliminating or reducing the numbers of
native organisms. The great filtering capacity of these small mussels reduces the phytoplank-
ton and increases water clarity. This allows large, rooted aquatic plants to increase and
changes not only the food supply, but the physical structure of the system. In some instan-
ces, the sessile juveniles and adults even outcompete zebra mussel larvae, which rely upon
phytoplankters for nourishment. A diminished supply of phytoplankters such as diatoms
leads to a reduction in the zooplankters upon which many fishes depend. Larval stages of
fish such as the bluegill (Lepomis macrochirus) in inland lakes may be strongly affected. At
the same time, the prodigious amount of feces and pseudofeces produced in large druses
increases the food supply for macroinvertebrate bottom feeders and thus for fish that feed
on these animals. Bacterial productivity also increases to break down the mussels’ wastes,
which may increase food available to the mussels themselves. Zebra mussels are eaten by
native mollusk-eating fish such as freshwater drum (Aplodinotus grunniens), lake sturgeon,
yellow perch, and catfish, which have been shown to increase in heavily infested waters.
Thus, a change in the composition of the fish community occurs as benthic-feeding fish
replace planktivorous species.

Zebra Mussels

T he invasion of zebra mussels in the 1980s sparked new interest in the problems
associated with exotic species. The new science of invasion biology developed
within the discipline of ecology, the Nonindigenous Aquatic Nuisance Prevention
and Control Act was passed, and public perception of the threats posed by intro-
duced species to both aquatic and terrestrial ecosystems grew. Invasions became
a common topic in the media and increasingly a subject of scientific research. For
some, the zebra mussel is the “poster child” of biological invasions.
86 n INVERTEBRATES (CRUSTACEANS)

The attachment of zebra mussels to the shells of native clams (unionids) makes it more
difficult for the clams to move through sediments in search of food and optimal concentra-
tions of oxygen. In contrast, encrusted bivalves in wave-impacted areas are easily dislodged
and washed away from prime sites. Furthermore, mussels can obstruct the openings of the
clams’ shells and prevent food intake and the release of gametes or prevent defensive closing
of the shell. During the very early invasion of Lake Erie, reports came of thousands of zebra
mussels on a single unionid. Native clam populations in the western basin of Lake Erie and
in Lake St. Clair have plummeted.
The zebra mussel is the most serious biofouling organism to have been introduced into
American waters. It clogs water intake and distribution pipes in industrial and power plants
that withdraw water from lakes and rivers, increases the corrosion rates of iron and steel
pipes and rivets, and obstructs pumps, valves, weep holes, screens, and the like, damaging
equipment and entire facilities. After storms on the Great Lakes huge windrows of mussel
shells accumulate on beaches; the decaying mollusks release methane and produce a
noxious stench. The razor-sharp shells are hazards to beachgoers.
Management. Control of zebra mussels is problematic. Chemical molluscicides; manual
removal; ultraviolet, hot water, and CO2 treatments; filters; screens; and electrical currents all
have been tried. Preventing their spread into new waterways is the target of most management
schemes. This can be best achieved through the decontamination of diving and fishing gear and
of boats and boat trailers, and stopping the dumping of unused bait in lakes and rivers.

Selected References
Benson, A. J., and D. Raikow. “Dreissena polymorpha.” USGS Nonindigenous Aquatic Species Database,
Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=5.
Nichols, S. Gerrine. “Zebra Mussel Identification.” Zebra Mussel Research Program. Zebra Mussel
Information System, Environmental Laboratory, Engineer Research and Development Center,
United States Army Corps of Engineers, 2002. http://el.erdc.usace.army.mil/zebra/zmis/zmishelp4/
zebra_mussel_identification.htm.

n Crustaceans
n Chinese Mitten Crab
Scientific name: Eriocheir sinensis
Family: Varunidae (formerly Grapsidae)
Native Range. Subtropical East Asia, from Fukien Province, China, north to the Korean
Peninsula.
Distribution in the United States. The Chinese mitten crab is established only in
California, where it occurs in the Sacramento-San Joaquin Delta and tributary streams.
Specimens have been collected in Delaware, Louisiana, Maryland, New York, Ohio, and
Washington. Until 2007, all those collected outside of California were males; but recently
females with eggs have appeared in the Mid-Atlantic states, so breeding populations may
now exist there.
Description. The Chinese mitten crab is most easily identified by the character that gives
it its name: light brown, hairy claws of equal size that have white tips and make it look like
 CHINESE MITTEN CRAB n 87

the animal is wearing mittens.

The brown hairs (setae) are evi-
dent in juveniles when the cara-
pace is about an inch (25 mm)
wide. The carapace is round in
outline and convex, but bumpy
on the dorsal surface. There is
a distinct notch between the
eyes. On the edges of the front
part of the carapace are four
spines, the fourth of which is
quite small. Females have a wide
abdominal flap shaped like a
beehive; males have a narrow,
bell-shaped abdominal flap.
These differences are apparent
in animals larger than 0.4 in.
(10 mm). The legs are more than
twice as long as the carapace is
wide. The crab is brownish
orange to greenish brown in
color. Carapaces of adults are
1.2–4.0 in. (30–100 mm) wide.
In California, adults reach about
3 in. (80 mm).
Juvenile Chinese mitten crabs
will dig burrows into vertical,
clay river banks within the inter-
tidal zone. The burrows have
oval openings and may be 1–3
in. (2.5–7.6 cm) wide and as
much as 8 in. (20 cm) deep. Top: The Chinese mitten crab is native to subtropical regions of east Asia.
They slope downward and may Bottom: The Chinese mitten crab is invasive only in California, although
have two entrances. individuals have been collected elsewhere. Females with eggs have been
Related or Similar Species. reported from Mid-Atlantic states, suggesting populations may be becom-
The Harris mud crab (Rhithro- ing established there. (Adapted from Benson and Fuller 2007.)
panopeus harrisii), similar in
size and general appearance and native to the Atlantic coast of North America, could be
mistaken for a young Chinese mitten crab. However, it lacks the hairy claws and the notch
between the eyes and is most commonly found in estuaries. The Harris mud crab has
ridges on the back of the carapace, which is brown or black. Maximum carapace size is
0.75 in. (19 mm). The entrances to its burrows are circular and about 1.5 in. (38 mm)
wide.
Occasionally, the very similar Japanese mitten crab (Eriocheir japonicus) is reported in the
United States, but this species is not widely established outside its native range. (Some scien-
tists think it is the same species as the Chinese mitten crab.)
The Chinese mitten crab is the only crab found in freshwater in the United States. Its
carapace shape differs from all other true crabs that occur here.
 88 n INVERTEBRATES (CRUSTACEANS)

Introduction History. A sin-

gle Chinese mitten crab was
taken by a shrimp trawler in
South San Francisco Bay
in 1992. They were captured in
San Pablo Bay in 1994, Suisun
Marsh in 1996, and the Delta
in the fall of 1996. Several dozen
were trapped upstream at the
U.S. Bureau of Reclama-
tion’s Tracy Fish Collection
Facility at the Delta Mendota
canal in 1996. Migrating crabs
in the fall of 1997 numbered in
the tens of thousands; and the
following year, the population
Chinese mitten crabs get their name from their hairy claws. (© N. Sloth/
was estimated to be 775,000
Biopix.)
and threatened to shut down
fish salvage operations at the canal pump station. By 2000, Chinese mitten crabs were
reported 30 mi. (50 km) upstream from the Bay-Delta, with no halt to their expansion
expected.
It is likely that this popular Asian delicacy was deliberately introduced as live crabs for
human consumption or to develop a local food resource, although the release of untreated
ballast water could also disperse these animals. Specimens collected in other parts of the
United States suggest that the illegal import and release of live crabs continues, although
so far, populations have not become established outside of California. The crab was found
in Hawai’i in the 1950s; and Chinese mitten crabs have been reported from the mouth of
the Columbia River, Washington, from Yaquina Bay in central Oregon, and from the Great
Lakes region since the 1960s. More recently they have shown up in Chesapeake Bay
(2005–2007), Delaware Bay (2007), the Hudson River in New York, (2007–2009), and
the Shrewsbury River in New Jersey (2008–2009) on the East Coast of the United States.
Only in New York have both male and female adult and juvenile crabs been collected. Live
crabs have not been detected in Chesapeake and Delaware bays since 2007.
Habitat. The Chinese mitten crab is catadromous and spends most of its life cycle in
freshwater. Juvenile and adult crabs are also able to walk on dry land. Adults live in the bot-
toms and banks of freshwater streams, but they reproduce in the brackish water of estuaries.
Late larval stages float in the upper part of the water column and are transported by currents
toward the mouths of estuaries. Settling and metamorphosis into juveniles occurs on the
seafloor along coasts and in embayments. A temperate-zone species, it is adapted to chang-
ing water temperatures and salinities and thus tolerates pollution. Optimal temperatures
are 75–82°F (24–28°C) for juveniles, 59–64°F (15–18°C) for larvae. Larvae in early stages
of development tolerate a wide range of salinities.
Diet. Usually described as opportunistic omnivores, Chinese mitten crabs consume algae,
macrophytes, detritus originating on land, and invertebrates. They scavenge dead fish and
are notorious for stealing fishermen’s bait.
Life History. In San Francisco Bay, Chinese mitten crabs mate in the late fall and winter
when water salinity is greater than 20 ppt. Some 24 hours later, a female spawns eggs, which
 CHINESE MITTEN CRAB n 89

are affixed underneath her abdominal flap. A female will carry 250,000 to one million eggs
through the winter until they hatch in the spring or summer. Hatchlings become part of
the plankton for 1–2 months. They settle in salt or brackish water in late spring or early
summer, and the juveniles migrate to freshwater, perhaps aided by tidal currents. When
living in tidal freshwater rivers, the juveniles congregate in dense colonies and burrow into
the bank for protection against predators and desiccation at low tide. In nontidal areas,
they apparently do not burrow. Older juveniles occur farther upstream than younger ones.
While migrating upstream, crabs will leave the water and walk across banks and levees to
bypass dams and other obstacles. When 1–4 years old, the males and females migrate
downstream in late summer or fall to brackish water, where they become sexually mature.
The males arrive first; mating begins as soon as the females arrive. Adults die soon after
mating.
Impacts. In the San Francisco estuary, Chinese mitten crabs may compete for food and
shelter with such commercially important species as the red swamp crayfish (Procambarus
clarkia) and the signal crayfish (Pacifasticus leniusculus). However, it is primarily a nuisance
species for shrimp trawlers and both commercial and recreational fishermen, stealing bait,
tearing nets, getting entangled in gear, and eating the catch. During downstream migra-
tions, the large numbers of crabs involved will clog water intakes, reducing water flow
and potentially causing power plant systems to overheat. The burrowing of juveniles can
accelerate river bank and levee erosion and collapse. Chinese mitten crabs are intermedi-
ate hosts of a mammalian lung fluke (Paragonimus spp.) that could affect humans.
However, neither the parasite nor its primary hosts, Asian freshwater snails, have been
found in the United States.
In China, the mitten crab is an agricultural pest in rice paddies, eating young shoots and
damaging levees. There is concern that should the crab become established in the rice-
growing areas of the Gulf coast, it could have similar negative impacts.
Management. In Germany, where these crabs have become a major problem, some suc-
cess in controlling them has been achieved with traps on the upstream side of dams captur-
ing juveniles during their upstream migration. Similarly, the use of traveling screens and
trash racks at water intakes can capture large numbers of migrating crabs. Periodic back-
flushing can remove crabs from water intakes.

Selected References
Benson, A. J., and P. L. Fuller. “Eriocheir sinensis.” USGS Nonindigenous Aquatic Species Database,
Gainesville, FL, 2010. Revised July 20, 2007. http://nas.er.usgs.gov/queries/FactSheet.aspx
?speciesID=182.
“Chinese Mitten Crab.” ProjectUFO, 2009. http://www.projectufo.ca/drupal/Chinese_Mitten_Crab.
Chinese Mitten Crab Survey Program. “Chinese Mitten Crab Update, U.S. Atlantic Coast Bays and
Rivers.” Smithsonian Environmental Research Center, 2009. http://www.serc.si.edu/labs/marine_
invasions/news/CHINESE_MITTEN_CRAB_UPDATE_APR21_09.pdf.
Chinese Mitten Crab Working Group. “National Management Plan for the Genus Eriocheir (Mitten
Crabs).” Aquatic Nuisance Species Task Force, 2003. http://www.anstaskforce.gov/Species
%20plans/national%20mgmt%20plan%20for%20mitten%20crab.pdf.
“Life History and Background Information on the Chinese Mitten Crab.” California Department of Fish
and Game, 1998. http://www.dfg.ca.gov/delta/mittencrab/life_hist.asp.
 90 n INVERTEBRATES (CRUSTACEANS)

n Green Crab
Also known as: European green crab, European shore crab
Scientific name: Carcinus maenas
Family: Portunidae
Native Range. The green crab is native to the western Baltic Sea and the Atlantic coasts of
northwest Europe and North Africa from Iceland and Norway to Great Britain and south
along Spain and Portugal to Morocco and northernmost Mauritania.
Distribution in the United States. On the Pacific coast, the green crab is established in
most bays and estuaries from Monterey Bay, California, north to Grays Harbor,
Washington. Populations, how-
ever, have remained small.
Along the Atlantic seaboard,
green crabs are established along
the coast of New England. They
are found south along the
Atlantic coasts of Maryland and
Virginia, but have not been col-
lected in Chesapeake Bay. A sin-
gle green crab was collected in
Hawai’i in 1973.
Description. The green crab
has a somewhat hexagonal
carapace. Five “teeth” or blunt
spines edge the carapace be-
hind each eye. Three bumps or
rounded teeth lie between the
eyes. The carapace has a granu-
lar texture and is usually a mot-
tled dark green or brown with
white or yellowish spots. The
carapace is broader than it is
long; its width is 2.4–3.9 in.
(6–10 cm). The ventral surface
varies in color from green to
yellow, orange, or red, depend-
ing on molt status. The second
and third pairs of walking legs
are the longest and are almost
twice as long as carapace
length. The fourth pair is the
shortest; they are relatively flat
Top: Green crabs are native to the Atlantic coast of Europe and North
and bear hairs (setae).
Africa. (Adapted from map by National Introduced Marine Pest System Related or Similar Species.
[NIMPIS], Australia.) Bottom: The green crab is established in bays and On the West Coast, some native
estuaries from Monterey Bay, California, northward along the west coast crabs are green and could be
and from Maine to the mouth of the Chesapeake Bay along the east mistaken for green crabs. One
coast. (Adapted from Perry 2010.)
 GREEN CRAB n 91

such native is the helmet crab

(Telmessus cheiragonus), which
is distinguished by six spines
behind each eye, a pair of long
antennae and a body covered
in stiff hairs. The green crab is
the only shore crab on the
Pacific coast with five spines at
the edge of the shell behind
each eye.
Introduction History. Green
crabs arrived in the United
States in the early 1800s at or
near Cape Cod, Massachusetts.
The first known report dates to
1817. They began to extend
their range northward in the
early twentieth century, enter-
ing Maine’s waters in the
1950s, and reaching Nova
Scotia, Canada, in the 1960s.
They were first collected in
New Jersey in 1929 and later
in Maryland and in Chinco-
teague Bay, Virginia. East Coast
introductions could have come The green crab occupies a wide variety of habitats on sheltered coasts. (P.
in ballast water or by crabs Erickson. Reprinted with permission of the MIT Sea Grant College
living amongst fouling organ- Program.)
isms on the hulls of transoce-
anic vessels or in the burrows of shipworms living in wooden sailing ships.
The green crab was first reported on the Pacific coast in 1989 in Estero Americano,
Sonoma and Marin counties, California. About the same time, it appeared in San Francisco
Bay. Genetic analyses suggest they came from the east coast of North America and did not
represent a direct introduction from Europe or North Africa. It is probable that they arrived
in algae used to pack New England baitworms (Nereis virens and Glycera dibranchiata).
Green crabs were first collected in Bodega Harbor in 1993 and Humboldt Bay, 1995. They
appeared in Oregon, first in Coos Bay (1997), then in Tillamook, Netarts, and Yaquina bays
in 1998. Also in 1998, they were first collected in Grays Harbor and Willapa Bay,
Washington. It is believed that strong El Niño–derived currents transported green crab lar-
vae northward in 1997–1998. In many of the invaded areas, populations have remained
small or have died out. The highest densities occur in Bodega Harbor, but these are slight
compared to East Coast infestations.
Habitat. Green crabs can be found on protected and semi-protected coasts under both
marine and estuarine conditions. They will occupy both rocky and soft sediment shores, sea-
grass meadows, and tidal marshes. In the United States, they are important members of
rocky intertidal communities on the East Coast, but on the West Coast, they do not occupy
exposed rocky shores. Juveniles and adults occur in intertidal and subtidal zones to depths
of 180 ft. (55 m). Adult green crabs tolerate wide ranges of salinity (4–52 ppt) and
92 n INVERTEBRATES (CRUSTACEANS)

temperature (32–86°F or 0–30°C), but larvae require salinities of 26–29 ppt and tempera-
tures of 52–77°F (11–25°C) for successful development.
Diet. Green crabs are primarily predators that consume bivalves, including mussels,
clams, oysters, and scallops; snails; other crabs; barnacles; and isopods. However, they
also eat algae. They may dig nearly 6 in. (15 cm) into sandy or muddy bottoms in search
of prey.
Life History. Mating occurs after the females molt. The time of the molt varies with geo-
graphic location, but most commonly is between June and October. A male carries a smaller
female underneath his body until she molts. At that time, she turns over and releases a
mass of eggs that she will keep on her abdomen. The male then releases sperm and fertil-
izes them. The eggs are held by the female through the winter until spring or early
summer. When the larvae hatch, they become part of the plankton and drift in the water
column for several weeks or months. It appears that tidal currents transport the larvae
away from shore and into the open sea. During the planktonic stage, larvae molt and pass
through several stages of the life cycle. During the final larval stage, tides bring them back
to shore, and they metamorphose and settle to the bottom as juvenile crabs. The juveniles
grow to a carapace width of about 0.25 in. (6 mm) before their first winter and 0.5–1.0 in.
(13–25 mm) before their second winter. They are sexually mature at 2–3 years of age and
may live for 5 years.
Impacts. The green crab is believed largely responsible for the decline of New England’s
soft shell clam (Mya arenaria) fishery in the 1950s. It is also implicated in declines of the
northern quahog (Mercenaria mercenaria) and a scallop (Argopecten irradians).
The cultivated Manila clam (Venerupis philippinarum) harvest in Tomales and Humboldt
bays, California, declined by 40 percent after the establishment of green crabs. In Bodega
Harbor, decreases in the density of small clams (Nutricola spp.) and shore crabs
(Menigrapsus oregonensis) are attributed to the introduction of the green crab. It is feared that
the commercially important Dungeness crab (Cancer magister) could be similarly affected if
the green crab were to expand its range and numbers along the west coasts of the United
States and Canada, since it could prey on juveniles.
Management. Fencing, trapping, and poisoning were generally ineffective where tried on
the East Coast. Washington is attempting to stem the influx of green crabs legislatively by
declaring it a deleterious exotic species and prohibiting the possession and transportation
of the animal in the state. Aquaculture in Washington is under restrictions that limit the
transfer of shells, living animals, and equipment from infested Willaba Bay and Grays
Harbor to waters the crab has not yet invaded. Imports of shellfish seed from out-of-state
waters where green crabs occur are also restricted.

Selected References
“Carcinus maenas (European Green Crab).” Invasive Species Fact Sheet. Washington Department of
Fish and Wildlife, 2005. http://wdfw.wa.gov/fish/ans/identify/html/index.php?species=carcinus
_maenas.
Cohen, Andrew N. “Carcinus maenas (Linnaeus, 1758).” Guide to the Exotic Species of San Francisco Bay.
San Francisco Estuary Institute, Oakland, CA, 2005. http://www.exoticsguide.org/species_pages/
c_maenas.html.
Perry, Harriet. “Carcinus maenas.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL,
2010. Revised April 25, 2008. http://nas.er.usgs.gov/queries/FactSheet.aspx?SpeciesID=190.
 RUSTY CRAYFISH n 93

n Rusty Crayfish
Also known as: Crawdad, crawfish
Scientific name: Orconectes rusticus
Family: Cambaridae
Native Range. Rusty crayfish are native to the Ohio, Tennessee, and Cumberland drain-
age systems in the states of Kentucky, Indiana, Illinois, Ohio, and Tennessee.
Distribution in the United States. Outside their native range, rusty crayfish are native
transplants in the Great Lakes region, Iowa, Maryland, Missouri, New Jersey, New York,
Pennsylvania, and all of the New England states except Rhode Island.
Description. The rusty crayfish has relatively large claws that are smooth and have black
tips. When closed, the pincers leave an oval gap between them. Color varies from gray-green
to reddish brown. The most distinctive feature is a pair of dark-red or rusty spots on either
side of the carapace just in front of the abdomen (“tail”); however, these spots are not always
present. Maximum length of the body exclusive of the claws is about 4 in. (10 cm). Males are
larger than females.
Related or Similar Species. Some 65 species of the genus Orconectes are native to the
United States. The rusty crayfish has larger, smoother claws than most others. Among those
most apt to be found where rusty crayfish have been introduced, the northern clearwater
crawfish (O. propinquus) is most similar. However, it lacks the two spots and instead has a
dark patch on its tail. Another close relative, the virile crayfish (O. virilis), an invasive in part
of its range, can be distinguished by the white bumps on the claws and a very narrow gap
when the claws are closed. The calico or papershell crayfish (O. immunis) shows a distinct
notch in the closed claws. Neither the virile nor the calico crayfish has black bands at the tips
of their claws.
Introduction History. The expansion of the rusty crayfish out of its native range is not
well documented, but it is assumed that nonresident fishermen, familiar with them in their
homes states within the Ohio River basin, brought them to new areas as bait and released
them intentionally or accidentally. Rusty crayfish are also popular in school laboratories
and sold by biological supply companies. Releases from classrooms are another way rusty
crayfish may have entered nonnative waters. Rusty crayfish are commercially harvested for
human food, and this provides an economic incentive to introduce them to new locations.
Since female crayfish store sperm, a single animal could start a new population.
Their introduction to Wisconsin lakes and streams occurred sometime in the 1960s. The
first report from southern Minnesota came in 1967. In both of these states, the range has
increased dramatically in ensuing years. The most recent introduction was in Maryland,
where they were found in the Monocacy River and Susquehanna drainage in 2007.
Habitat. These freshwater crustaceans require permanent bodies of water and adequate
cover such as rocks, tree limbs, or logs. They inhabit lakes and streams, where they can be
found in both still pools and fast-flowing stretches. A variety of bottom types from silts
and sands to gravel or rock are used. They do not dig deep burrows as do some of their rel-
atives and so cannot escape when intermittent streams dry up.
Diet. Rusty crayfish have unusually high metabolic rates and are therefore voracious and
opportunistic feeders. They consume aquatic plants; bottom-dwelling invertebrates such as
aquatic worms, snails, bivalves, insect larvae, and crustaceans; detritus; fish eggs; and small
fish. Juveniles concentrate on invertebrates such as mayfly, stonefly, and midge larvae, and
freshwater shrimp.
 94 n INVERTEBRATES (CRUSTACEANS)

Life History. Rusty crayfish

mate in late summer or early
fall, and sometimes in early
spring. The male transfers his
sperm to the female, which
stores them in a seminal recep-
tacle on her belly until the water
begins to warm in late April or
May. External fertilization takes
place when eggs and sperm
are simultaneously released by
the female. As this happens,
she secretes a mucus-like sub-
stance called glair that forms
white patches on the underside
of the tail fan. The fertilized
eggs attach to the swimmerets
on the underside of the female’s
abdomen by means of the glair.
Females produce 80–575 eggs.
It takes 3–6 weeks for the
eggs to hatch. The young con-
tinue to hold on to the mother’s
swimmerets for several weeks
as they go through 3–4 molts.
After the young leave the
female, they undergo another
8–10 molts before they become
adults, usually the year after
they hatch. Sexual maturity
occurs at a total body length of
Top: Rusty crayfish are native to the Ohio, Tennessee, and Cumberland 1.4 in. (3.5 cm). Growth slows
river systems. Bottom: Rusty crayfish are invasive native transplants in once the crayfish is mature.
many drainages of the Great Lakes region, the central United States, and Females molt only after re-
the Northeast. (Both maps adapted from USGS 2008.) leasing their young, usually in
June or early July. Males molt
twice a year, once in the spring into a sexually inactive form, and again in summer into a
reproductive form. The twice-annual molt allows males to become larger than females.
Rusty crayfish live 3–4 years.
Impacts. Rusty crayfish are aggressive and displace native crayfish, such as the virile cray-
fish and the northern clearwater crayfish, from shelter and compete with them for food.
When the smaller natives are forced from their hiding places, they become vulnerable to
increased predation by fish. Declines in these two native crayfish have occurred in lakes in
Wisconsin (and parts of Ontario, Canada). Another close relative, Sanborn’s crayfish
(Orconectes sanbornii), has been displaced in Ohio in waters where it was not native.
Hybridization with the northern clearwater crayfish contributes to the decline of that species.
The high metabolic rate of the rusty crayfish means it consumes a lot of food for its size.
The destruction of submerged vegetation that provides shelter for invertebrates, young game
 SPINY WATER FLEA n 95

fish and panfish, and forage

species for other fish can be
particularly damaging to the
aquatic ecosystem as it elimi-
nates important and sometimes
scarce habitat. They may
directly harm native fish by
consuming their eggs and
reducing the amount of inverte-
brate prey items upon which
juvenile fish depend. Declines
have been documented in larval
midges, mayflies, and stoneflies
and in fish such as bluegill
(Lepomis macrochirus), pump-
kinseed (Lepomis gibbosus), The rusty crayfish is so named because of the rusty patches of color on
smallmouth bass (Lepomis gib- either side of the carapace just in front of the tail. (Jeff Gunderson,
bosus), largemouth bass (Micro- Minnesota Sea Grant.)
pterus salmoides), lake trout
(Salvelinus namaycush), walleye (Sander vitreus), and northern pike (Esox lucius) when rusty
crayfish invade.
Large numbers of rusty crayfish become a nuisance to swimmers, who are in danger of
stepping on them and being pinched by their large claws.
Management. Few environmentally sound means of control are available. Commercial
harvests may reduce numbers and keep them in check. Some researchers believe that popu-
lation explosions of rusty crayfish are in part due to the overfishing of predatory fishes, and
they recommend restoring healthy populations of sunfish and bass as a way of reducing the
impacts of rusty crayfish. The best management practice is to prevent further introductions
by educating people about the ways in which this little animal threatens their local waters.

Selected References
Gunderson, Jeff. “Rusty Crayfish: A Nasty Invader.” Minnesota Sea Grant, 2008. http://
www.seagrant.umn.edu/ais/rustycrayfish_invader.
Pappas, J. “Orconectes rusticus.” Animal Diversity Web, 2002. http://animaldiversity.ummz.umich.edu/
site/accounts/information/Orconectes_rusticus.html.
U.S. Geological Survey. “Orconectes rusticus.” USGS Nonindigenous Aquatic Species Database,
Gainesville, FL, 2008. http://nas.er.usgs.gov/queries/factsheet.aspx?SpeciesID=214.

n Spiny Water Flea

Scientific name: Bythotrephes longimanus
Order: Cladocera
Family: Cercopagidae
Native Range. Northern Eurasia, from Great Britain and Scandinavia across Russia to the
Bering Sea. Since the spiny water flea has been widely dispersed by humans in Europe and
Asia to lakes in which it was not native, its true natural range is difficult to reconstruct.
 96 n INVERTEBRATES (CRUSTACEANS)

Distribution in the United

States. This invader is estab-
lished in all the Great Lakes
and has been found in inland
lakes in Ohio, Michigan (Long
Lake), Minnesota (Greenwood
and Flour lakes), New York
(Great Sacandaga Lake) and
Wisconsin (Stormy Lake and
Gile Flowage).
Description. The spiny water
flea is not an insect, but a
crustacean. Many of its features
are difficult to discern with-
out magnification. Because
they assemble into gelatinous
clumps, they often resemble
wet cotton batting full of tiny
black spots. Each individual in
the mass has a well-developed
abdomen and a long, thin tail
spine. The head is clearly sepa-
rated from the trunk and bears
a single large black compound
eye. Two swimming antennae
used to propel the animal
through water are attached just
behind the head. The spine is
straight and accounts for more
than 70 percent of total body
length. The spine bears 1–4
Top: Spiny water fleas occur across northern Eurasia. The exact native pairs of barbs depending upon
range is uncertain because people have introduced the species to many the animal’s age. Juveniles have
lakes in both Europe and Asia. Bottom: The spiny water flea is invasive a single pair; additional pairs
in all of the Great Lakes as well as inland lakes in adjacent states. are added above the spine when
(Adapted from Liebig and Benson 2007.) the exoskeleton covering the
trunk of the body is shed to
allow for growth. Two morphs
representing different reproductive modes (see under Life History) are present.
Parthenogenically reproduced individuals, all females, have longer tail spines with an
obvious “kink” in the middle of the spine and, when fully developed, have three pairs of
barbs resulting from two molts. Sexually reproducing females will acquire four pairs of
barbs during three molts. (The parthenogenically reproduced females were once believed
to be a separate species and identified as B. cederstroemi.) Females are also identifiable
because they carry their eggs in a brood pouch that balloons from the back of the body.
Males will not gain a new pair of barbs during their final molt and end up with only two
pairs.
 SPINY WATER FLEA n 97

Spiny water fleas have four

pairs of legs. The first pair is
longer than the others and used
for snaring prey. The others are
used to grasp the prey while it
is being eaten. Average adult
size is 0.4 in. (1 cm). The spiny water flea is a crustacean barely visible with the naked eye.
Related or Similar Species. (Microscopy by Howard Webb.)
This tiny crustacean has a
unique body shape with a long tail spine that clearly distinguishes it from freshwater zoo-
plankters native to the Great Lakes. Another exotic cladoceran, the fishhook water flea
(Cercopagis pengoi), also collects into masses, but individuals can be separated from spiny water
fleas by virtue of their angled tails that end in a distinct loop or “fishhook” and their tiny eyes.
Introduction History. The first live specimens of spiny water flea were retrieved from
Lake Huron in 1984. The next year, they were found in Lakes Erie and Ontario. By 1986,
they had made their way to Lake Michigan, and by 1987, they had invaded Lake Superior.
By early 2002, they had been reported from 66 lakes in the northern United States and
southern Canada. In 2007, spiny water fleas were discovered in Stormy Lake, Wisconsin,
and in 2008, they were found in Great Sacandaga Lake, in New York’s Adirondack
Mountains.
Genetic analysis has traced the source of the invasion to the port of St. Petersburg,
Russia. Spiny water fleas are native to nearby Lake Lagoda. During spring snowmelt, they
are carried down the Neva River to the harbor, where the water becomes fresh or only
slightly brackish. Freighters, which had brought wheat from the United States, return
empty and therefore take up ballast water and any zooplankton contained in it at St.
Petersburg. They then discharge that water when they next take on grain in American ports
in the Great Lakes.
Local dispersal from the Great Lakes to inland lakes probably occurs by means of
contaminated fishing gear and boats.
Habitat. Spiny water fleas do best in deep lakes in temperate climates. They will inhabit
both large and small lakes, and may live in shallow lakes and rivers as well as in brackish
waters. They are limited to water temperatures of 39–86°F (4–30°C) and salinities of
0.04–8.0 percent. Optimal conditions are temperatures of 50–75°F (10–24°C) and salinities
of 0.04–0.4 percent. Females are conspicuous to predatory fish during daylight hours
because of their large eyes and brood pouches, and thus undergo daily vertical migrations.
During daylight hours, they move lower into the water column and are usually at depths
of 30–65 ft. (10–20 m). They swim up closer to the surface where food is more abundant
and at night, when most will be found within the top 30 ft. (10 m) of the water column.
Diet. Spiny water fleas are tiny predators that feed on smaller, herbivorous zooplankters.
Daphnia spp. and other water fleas are preferred prey, but they also eat copepods and roti-
fers. Studies show that a single spiny water flea can consume 20 prey items a day.
Life History. Spiny water fleas have a complex life cycle involving both sexual and asex-
ual reproduction. Most of the time, females reproduce by cloning (parthenogenesis) and
produce 1–10 eggs that develop into new females without fertilization by males. A new
generation may be produced every two weeks in the summer, when temperatures are
warm and food abundant. Under these conditions, males are rare in the population.
However, sex of the young is determined not by genetics, but by the environment. When
98 n INVERTEBRATES (CRUSTACEANS)

conditions deteriorate in the fall, males begin to be produced. They mate with females that
then produce “resting eggs.” At first, the resting eggs are carried in the brood pouch, but even-
tually they are released into the water and settle to the bottom of the lake, where they go into a
near-dormant phase (diapause) in order to survive the cold of winter. As water temperature
rises to 39°F (4°C) and above in the spring, the eggs hatch into females that will reproduce
parthogenetically and make possible rapid population growth during favorable conditions.
The brief sexually reproducing part of the life cycle promotes both genetic diversity and dis-
persal capability.
Impacts. The initial impact of the introduction of spiny water fleas was as a nuisance to
fishermen. Hundreds of these cladocerans can clump onto fishing lines and downrigger
cables, especially at connections and swivels, and clog the first guide of an angler’s rod.
Sometimes it becomes impossible to reel in the catch, and the line must be cut. With time,
ecological impacts became evident. The community structure of the zooplankton commu-
nity can be altered by heavy predation on small herbivorous crustaceans such as Daphnia,
copepods, and rotifers. In Lake Michigan, both D. retrocurva and D. pulicaria populations
collapsed after the introduction of water fleas, and populations of another tiny cladoceran,
Holopedium gibberum, declined dramatically. In consuming these animals, spiny water fleas
also compete for food with the plankton-feeding larvae of several fish species as well as with
other cladocerans, such as the giant water flea (Leptodora kindtii), which declined in some
lakes when spiny water fleas were introduced. Such results are not always duplicated in
smaller lakes.
Impacts on fish populations are not clear. Juvenile fish of species that feed heavily on
Daphnia may suffer, affecting annual recruitment rates of species such as yellow perch
(Perca flavescens). The barbed tail of the spiny water flea cannot be accommodated in the
small mouth gape and throats of young fish, so they cannot shift their diets to include the
invader species. On the other hand, adult yellow perch, as well as white perch (Morone
americana), bass (Micropterus spp.), and alewife (Alosa pseudoharengus) selectively prey on
spiny water fleas and could be helped by a new, seasonally abundant forage species. These
fish in turn are eaten by larger game fish such as Chinook salmon (Oncorhynchus tshawyt-
scha) and lake whitefish (Coregonus clupeaformis), so economically important fisheries could
be affected in a positive manner.
Management. Control measures are aimed at preventing the spread of the spiny water
flea to other inland lakes. Educational campaigns encourage fishermen to clean gear prop-
erly and empty bilges, livewells, and bait buckets responsibly before moving from one lake
to another.

Selected References
Caceres, Carla E., and John T. Lehman. “Spiny Tailed Bythotrephes: Its Life History and Effects on the
Great Lakes.” Minnesota Sea Grant College Program. University of Minnesota, 2004; last modified
January 26, 2010. http://www.seagrant.umn.edu/exotics/spiny.html.
Liebig, Jim, and Amy Benson. “Bythotrephes longimanus.” USGS Nonindigenous Aquatic Species
Database, Gainesville, FL, 2010. Revised January 25, 2007. http://nas.er.usgs.gov/queries/
FactSheet.aspx?SpeciesID=162.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Bythotrephes longimanus (Crustacean).” ISSG Global Invasive Species Database, 2005.
http://www.issg.org/database/species/ecology.asp?si=151&fr=1&sts=sss.
 HONEYBEE TRACHEAL MITE n 99

Sikes, Benjamin. “Spiny Water Flea.” Invader of the Month. Institute for Biological Invasions,
University of Tennessee, Knoxville, 2002. http://invasions.bio.utk.edu/invaders/fleas.html.

n Arachnids
n Honeybee Tracheal Mite
Scientific name: Acarapis woodi
Order: Trombidiformes
Family: Tarsonemidae
Native Range. Uncertain. Honeybee tracheal mites were first described from collections
made in dying bee colonies on the Isle of Wight in the English Channel in 1921. They were
responsible for wide-ranging honey bee mortality in Europe in the early part of the twentieth
century.
Distribution in the United States. Throughout the continental United States, except
Alaska, in both managed and feral honey bee colonies.
Description. Invisible to the naked eye, tracheal mites have white oval bodies with a
shiny, smooth cuticle. The body and legs have several long fine hairs. The mouth parts are
beak-like and elongated. Females are 143–147 microns long and 77–81 microns wide;
males are 125–136 microns long and 60–70 microns wide.
Few outward symptoms are evident in infected bees until the infestation is severe. Then
bees may develop disjointed, “K” wings and become unable to fly. Honey production may
decline. Sudden death of the hive during winter is a sign that the honeybee tracheal mite
may be present. Verification requires dissection of the dead bees and examination of the tra-
cheae, which will be brown instead of clear or white as in healthy bees.
Introduction History. Honeybee tracheal mites were unknown in North America prior to
1980, when they were detected in Mexico, some 200 miles (320 km) south of the U.S. bor-
der. They were first discovered in the United States at a commercial beekeeping enterprise in
Weslaco, Hidalgo County, Texas, in early July 1984. The following month, they were found
in New Iberia, Louisiana, and by October 1984, they were being reported in Florida,
Nebraska, New York, North Dakota, and South Dakota. By August 1985 they infested bee
colonies in 17 states. The rapid spread of the mite was facilitated by migratory beekeepers,
who truck bee colonies from southern states northward to pollinate various crops. The com-
mercial bee business also contributed to the spread by selling queens and package bees.
Mites move quickly through a colony via bee-to-bee contact in the hive. Workers and
drones drifting from hive to hive disperse the mite through entire apiaries or from one apiary
to another. Bees also encounter mites when they rob honey from other hives; and colonies
weakened by heavy mite infestations are particularly vulnerable to robbing. Since mites can-
not live more than a day outside the bee host, they readily leave dead bees and hitchhike to
new colonies inside robber bees. Normal swarming of bees can also spread the tracheal mite
to new areas.
Habitat. Honeybee tracheal mites are internal parasites that live within the breathing
tubes (trachea) of adult bees. They prefer the larger tubes at the base of the bees’ wings that
provide oxygen to the flight muscles. Occasionally, they occur in the air sacs. They infest
only European honey bees (Apis mellifera), Africanized honey bees (Apis mellifera scutellata;
see Insects, Africanized Honey Bee) and Asian honey bees (Apis cerana).
100 n INVERTEBRATES (ARACHNIDS)

Diet. Tracheal mites punc-

ture the walls of the breathing
tubes and feed on the blood
(hemolymph) of workers,
drones, and queen bees.
Life History. The life cycle
consists of four stages: egg,
larva, a resting stage nymph,
and adult. The female mite
enters a bee shortly after it
emerges from its cell by moving
through the first thoracic spi-
racle into the breathing tubes.
She remains in the host for the
rest of her life or until the bee
dies. Three or four days after
arrival, she lays 5–7 eggs, which
hatch in another three or four
days. Adult male mites are
mature 11–12 days after the
eggs are laid; females are mature
in 14–15 days. Gravid females
crawl to the tip of a bee hair
and “jump” to a new host,
entering through the breathing
pores (spiracles) to lay their
eggs. They tend to attack young
bees less than 24 hours old.
Mite populations tend to be
cyclical; heaviest infestations
occur in winter, when the bees
Top: The origins of the honeybee tracheal mite are unknown. It was first are confined and crowded in
reported affecting domestic honey bees in England. Bottom: The the hive, and then decline in
honeybee tracheal mite is found throughout the lower 48 states and on summer.
O’ahu, Hawai’i. Impacts. Initially, the intro-
duction of the tracheal mite
caused widespread losses of bee colonies and even entire apiaries throughout the United
States. Problems were especially severe in the more temperate parts of the country, where
bees cluster in confinement within the hive during the winter months. Highest bee mortality
usually occurs in late winter. Infected bees often show no signs of a problem, though pollen
collection and honey production may decline as a bee weakens.
The mites kill by clogging the breathing tubes with their bodies and waste products.
Normally elastic tracheae become stiff and brittle, and flight muscles atrophy.
Furthermore, hemolymph contains agents that act as antifreeze, so as the mites consume
the bee’s body fluid, it reduces its ability to withstand low temperatures.
The effects of mite infestation, known as acarine disease, may remain in a colony for years
with little damage. Worker bees and queens become less susceptible to infestation as they
age. It appears that most honey bees in the United States have developed some resistance
 HONEYBEE TRACHEAL MITE n 101

A. View of honeybee tracheal mite from above. B. Lateral view of honeybee tracheal mite. C. Mites clogging a
bee’s trachea. (Pest and Diseases Image Library, Bugwood.org.)

to or tolerance of mite infestations and that the problem of acarine disease is not as severe as
it was in the 1980s and 1990s. Its impacts may have been overshadowed by the introduction
of the varroa mite (Varroa destructor) in 1987 (see Arachnids, Varroa Mite).
Management. After the initial discovery of mites in Texas in 1921, the federal Honey Bee
of Act of 1922 was passed. It prohibited the importation of any honey bee into the United
States. The law has been relaxed somewhat since then to allow the import of bees from
Canada, where mite infestations are unknown, and a few other mite-free countries.
Chemical control may involve fumigation with menthol crystals. Grease patties, made of
vegetable shortening and sugar and available commercially, keep mites from infesting young
bees because the oily bees apparently do not “smell” right to mites. Evaporating formic acid
is also very effective in controlling tracheal mites and has the added bonus of reducing infes-
tations of the varroa mite.
102 n INVERTEBRATES (ARACHNIDS)

Several stocks of honey bees, including the Buckfast bee, have been bred to resist tracheal
mites. The bees still become infested, but only at levels too low to cause significant damage.
Use of these bees eliminates the need to treat the colony with chemicals.

Selected References
Ambrose, John T., and Michael Stanghellini. “Tracheal Mites.” Note 2.02. Beekeeping. Insect Pest
Management, Department of Entomology, North Carolina State University, 2001. http://
www.cals.ncsu.edu/entomology/apiculture/PDF%20files/2.02.pdf.
Collison, Clarence H. “Honey Bee Tracheal Mite.” Publication 1753, Extension Service of Mississippi
State University, 2009. http://msucares.com/pubs/publications/p1753.htm.
Hunt, Greg. “Parasitic Mites of Honey Bees.” Beekeeping. E-201-W. Department of Entomology, Purdue
University Cooperative Extension Service, 2006. http://extension.entm.purdue.edu/publications/
E-201.pdf.
Sammataro, Diana. “An Easy Dissection Technique for Finding the Tracheal Mite Acarapis woodi
(Rennie) (Acari: Tarsonemidae), in Honey Bees, with Video Link.” International Journal of
Acarology, 32(4), 2006. Available online at http://entnemdept.ufl.edu/HoneyBee/files/speaker
_notes/exp_wksp_Tracheal_Mite_Dissection_Notes.pdf.

n Varroa Mite
Scientific name: Varroa destructor
Order: Parasitiformes
Family: Varroidae
Native Range. Mainland Asia. It is known to be endemic to Japan, Korea, and Thailand,
where its host is the Asian honey bee Apis cerana. At some point it shifted hosts to the
European honey bee; and, in 1963 in Singapore, it was first identified on European honey
bees as a new species. Previously it had been classified as Varroa jacobsoni, a parasitic mite
of the Asian honey bee known from much of mainland and insular Asia.
Distribution in the United States. In all 50 states, in both wild and managed honey bee
colonies.
Description. This small arachnid is an external parasite of honey bees. Tick-like, it has a
flattened oval body and eight legs. Adult females are reddish brown and measure about 0.06
in. (1.5 to 1.99 mm) in width, approximately the size of the head of a pin. Their bodies are
curved to allow them to squeeze into the abdominal folds of a bee and thereby be protected
from the bee’s normal cleaning habits. Males are smaller and spherical in shape; they are yel-
lowish and have tan legs. Both sexes are visible to the unaided eye.
Symptoms of severe varroa infestations include an accumulation of dead bees at the
entrance to a hive, the uncapping and destruction of brood cells by worker bees, and
deformed legs and wings on adult bees.
Related or Similar Species. The bee louse (Braula coeca), a tiny wingless fly, is similar in
size and color, but has six legs instead of eight. It is rare in most hives today since it suc-
cumbs to treatments used to control the varroa mite.
Introduction History. A single varroa mite was discovered in Maryland in 1979, its
source unknown. It showed up again in 1987, this time in Florida and Wisconsin. It has
since spread rapidly throughout the United States. The last state invaded was Hawai’i, where
the first infestation occurred in O’ahu in 2007; the following year, the varroa mite was found
in honey bee colonies on the Big Island.
 VARROA MITE n 103

Mites are transported from

hive to hive by drifting workers
and drones. Worker bees also
pick up mites when they rob
honey from smaller colonies.
People can move mites from
place to place when they trans-
port infested colonies of bees
to fields and orchards for polli-
nation and through the impor-
tation of infested package bees.
Swarming bees may carry mites
to other apiaries or to feral pop-
ulations. It is also possible that
mites are spread short distances
on bumblebees and other
nectar-feeding insect hosts.
Habitat. Varroa mites are
usually found on the thoraxes
and abdomens of larvae, pupae,
and adults of all races of the
European honey bee (Apis melli-
fera) and the Africanized honey
bee (A. mellifera scutellata; see
Insects, Africanized honey
bee). All life stages inhabit the
brood cells of honey bees; adult
females will emerge with the
young bee and live on drone
and worker bees. The varroa
mite also occurs on the Ame-
rican bumblebee (Bombus penn- Top: The varroa mite is known to be native to Japan, Korea, and Thailand
sylvanicus), flower fly (Palpada and perhaps other parts of Asia. It originally infested the Asian honey bee
vinetorum), and rainbow scarab but later adopted the European honey bee as its host. Bottom: Varroa mites
beetle (Panaeus vindex), but are found in all 50 states, where they infest feral and managed bee colonies.
cannot reproduce on them. It
is also a parasite of Asian honey bees (A. cerana and A. koschevnilovi).
Diet. Varroa mites suck the hemolymph (“bee blood”) of developing honey bee larvae
and mature adults. They pierce the soft skin of larvae and the tougher integument of adults
to obtain this fluid. Like other arachnids, they inject enzymes that predigest a bee’s tissues so
they can consume them also.
Life History. The life cycle of varroa mites is synchronized with that of its host, the honey
bee. Females enter the brood cells of honey bees just before worker bees cap the cell, when
the bee larva is five days old and about to pupate. About three days later, she lays her first
egg, which usually is unfertilized and becomes a male. Later, she will lay a fertilized egg
every 30 hours; these become females. All immature mites feed on the bee larva and must
mature and mate before the bee emerges from its cell. It usually takes five to eight days for
the females to mature and a few days less for the male. The male mite dies in the cell, but
104 n INVERTEBRATES (ARACHNIDS)

A. Varroa destructor. B. Varroa mites at base of honey bee brood cell. C. Mite on honey bee pupa. (Scott Bauer,
USDA Agricultural Research Service, Bugwood.org.)

mated females will leave the cell on the host bee to seek new brood cells, preferably those of
drone larvae, where the process begins again. The average lifespan is about 50 days during
the breeding season. In winter, when brood-rearing by bees declines, mites live solely on
adult bees in the hive. Adult mites can survive only a few days without bees.
Impacts. Heavy infestations of varroa mites cause young bees just emerging from their
brood cells to have malformed wings, legs, and bodies. On adult worker bees, they may
affect the flight behavior, orientation, and success in returning to the hive laden with pollen
and nectar. As worker bees die off, fewer bees are available to tend to the brood and collect
nectar, and the colony weakens. Weak colonies are susceptible to having their honey stores
robbed by stronger hives. Eventually, the colony dies. In temperate climates, the death of a
colony may take 3–5 years; but in the milder climate of Florida, infested bee colonies have
died within seven months.
Mites transmit several RNA viruses that kill bees by compromising their immune systems.
These include deformed wing virus (DWV), acute bee paralysis virus (APV), and slow
 VARROA MITE n 105

paralysis virus (SPV). They result in a condition known as parasitic mite syndrome, which
can destroy a colony in a few months.
Without treatment, colonies—indeed, entire apiaries—can be wiped out by varroa mites.
In the United States, most wild (feral) bee populations, where treatment is not an option,
have been decimated by this invasive arachnid. Pollination of numerous field and orchard
crops depend upon honey bees. Their loss would have devastating economic impacts.
Management. Chemical means are available to detect varroa mites and control them in a
bee colony. A common treatment involves hanging strips impregnated with fluvalinate
(Apistan) in the brood nest area for about a month. As dead mites fall off bees, they can be
collected on sticky paper or a fine mesh placed screen on the bottom of a hive. This product
allows detection of low-level infestations. Such strips can also be used as a control measure.
A problem with this method is the development of fluvalinate-resistant mites. Another
chemical treatment using coumaphos (CheckMite+) is also very effective, but has the disad-
vantage of employing a dangerous organophosphate. Two formulations of thymol (oil of
thyme—Api-Life VAR and Api-Guard) and one using fenpyroximate (Hivastan) are also
available to beekeepers. All should be applied with careful adherence to instructions.
Research suggests biological controls may be implemented in the future. One candidate is
a strain of the fungus Metarhizium anisopliae that is lethal to varroa mites. Another possibility
is genetic engineering to develop a bee resistant to mite infestation. Asian bees (Apis cerana),
the original host for varroa mites, are aggressive groomers and cleaners and remove mites
from other bees by grooming and chewing the mite. They decap infested brood cells and
clean them out, discarding dead and dying larvae at the entrance to the colony. Bees that
develop faster in the brood cell can outpace the developing mites and emerge with fewer
of them. Bee breeders can select for queens that produce workers that are good at grooming,
that clean out brood cells, and that have quicker development times than average European
honey bees.
It is essential to stem the spread of infested colonies through constantly surveying bees for
the presence of the mite. Captured swarms, package bees, and other new colonies should be
quarantined away from the rest of the apiary and examined for mites before being allowed
contact with existing colonies.

Selected References
Bessin, Ric. “Varroa Mites Infesting Honey Bee Colonies.” ENTFACT-608. College of Agriculture,
University of Kentucky, 2001. http://www.ca.uky.edu/entomology/entfacts/ef608.asp.
Hunt, Greg. “Parasitic Mites of Honey Bees.” Beekeeping. E-201-W. Department of Entomology, Purdue
University Cooperative Extension Service, 2006. http://extension.entm.purdue.edu/publications/
E-201.pdf.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Varroa destructor (Arachnid).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?si=478&fr=1&sts=sss.
Reid, Brendan. “Varroa Mite (Varroa destructor).” Introduced Species Summary Project, Columbia
University, 2004. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/
varroa_destructor.html.
Sanford, M. T., H. A. Denmark, H. L. Comroy, and L. Cutts. “Featured Creatures: Varroa mite (Varroa
destructor).” University of Florida Institute of Food and Agriculture, 2007. http://www.entnemdept
.ufl.edu/creatures/misc/bees/varroa_mite.htm.
106 n INVERTEBRATES (INSECTS)

n Insects
n Africanized Honey Bee
Also known as: AHB, Killer bee
Scientific name: Apis mellifera scutellata
Order: Hymenoptera
Family: Apidae
Native Range. The African parent of the hybrid Africanized honey bee derives from
eastern and southern Africa, where Apis mellifera scutellata occurs from Kenya south to the
Indian and Atlantic coasts of South Africa. Bees from Tanzania were taken to Brazil, where
hybrids or Africanized honey bees were developed in Rio Claro in the state of Sao Paulo.
From there, they spread throughout tropical South America and Central America and into
the United States (see under Introduction History).
Distribution in the United States. As of July 2009, Africanized honey bees were estab-
lished from Texas and Oklahoma west through New Mexico and Arizona to southern
Nevada and southern California. Isolated swarms have been reported in southern Utah,
Arkansas, Louisiana, and Florida.
Description. The Africanized honey bee is a hybrid species and nearly indistinguishable
from the common domesticated European honey bee, one of its parents. The hybrid form
requires inspection and analysis in a laboratory to positively identify it. Behavioral traits do
set it apart, however. It is usually recognized by its extremely aggressive nature, the guarding
of a large area around its nest, frequent swarming, and nest location. Worker bees, all sterile
females, are about 0.75 in. (19 mm) long, imperceptibly smaller than European honey bees.
Drones, all males, have narrower bodies and are somewhat longer. Queens are larger still
and more robust, with enlarged abdomens.
The bees’ bodies are covered with a yellow-brown fuzz and marked with black stripes.
They have four clear wings and six legs, all attached to the middle body segment (thorax).
The abdomen is larger than the thorax and ends in a stinger. The venom is not as toxic
as that of European bees, but since large numbers of bees mass to defend their nest, the
multiple stings that result can be dangerous, indeed lethal, especially to sensitive persons.
Related or Similar Species. The European honey bee (Apis mellifera) is more a docile
domesticated animal selected for characteristics that suit it for easy handling by humans. It
produces and hoards more honey than Africanized honey bees and swarms less often.
European honey bees nest in large cavities in trees, in hollow walls, and in beehives, where
they develop from egg to adult a bit more slowly than their hybrid descendents. European
honey bees have adapted to the cold winters of the mid-latitudes, in part by storing large
amounts of honey in their hives.
Introduction History. African honey bees were brought to test sites in Rio Claro, Brazil, in
1956 by a Brazilian geneticist, Dr. Warwick Kerr. European honey bees had not fared well in
subtropical and tropical parts of South America. The hope was that crossing them with bees
from tropical Africa would improve both their honey production and survival rates by
selecting for the climatic adaptations of the African parents and the docility and honey-
producing attributes of the European ones. In 1957, 26 African queens swarmed with
European worker bees and escaped the experimental apiary. These bees hybridized with
commercial and feral populations of European honey bees, but in the wild retained most
of the traits of the African line, particularly the aggressive defense of their nests. They rapidly
 AFRICANIZED HONEY BEE n 107

spread through all of subtropical

and tropical South America
except Chile, and north through
Central America. They were first
identified in Mexico in 1985.
The rate of spread averaged
125–180 mi. (200–300 km) a
year. This dispersal was largely
unaided by humans. In 1990,
Africanized honey bees reached
Hidalgo, Texas, on the Mexican
border. By 1993, they were in
Arizona, and by 1994, they were
discovered near Blythe in
southern California. In 1996
they were found to the north in
Lawndale, California. Since that
time, the dispersal rate has, for
unknown reasons, slowed.
Africanized honey bees dis-
persed across South and
Central America by swarming
and by hybridizing with
existing feral and domestic col-
onies of European honey bees.
Anecdotal evidence suggests
European queens breed prefer-
entially with Africanized drones
and that Africanized queens
will enter the hives of Euro-
pean honey bees and kill and
replace the European queen. Top: The African parent of the Africanized honey bee has its origins in
It is believed that cold winter southern and eastern Africa. The hybrids between African bees and the
temperatures will prevent their European honey bee were developed in Rio Claro, Brazil. Bottom:
establishment very far north of Africanized honey bees are established in southern states from
the present U.S. distribution California to Florida. Short-lived colonies may appear as far north as
Virginia. (Adapted from “Spread of Africanized Honey Bees by Year, by
limits, and that high rainfall will
County.” USDA Agricultural Research Station, 2009. http://www.ars
restrict their spread east of .usda.gov/Research/docs.htm?docid=11059&page=6.)
Texas. Traits adaptive to tropi-
cal environments will be
selected against in colonies living in temperate climates. Disjunct populations in Florida
may have resulted from swarms arriving on ships; Africanized honey bees have been trapped
regularly at Florida’s deep-sea ports. Occasionally, temporary infestations occur as far north
as Virginia, perhaps the result of bees being transported accidentally on trains.
Habitat. The hybrid Africanized bees are less selective in habitat than the European
honey bee and nest in smaller numbers. Their nests may hang from exposed tree limbs or
under the eaves of buildings, or may be constructed in old tires and empty containers,
cement blocks, rotted logs, animal burrows, rock piles, and so forth—all places avoided
108 n INVERTEBRATES (INSECTS)

by European honey bees. They are apparently restricted to tropical and subtropical climates,
particularly those with distinct wet and dry seasons, and are seldom found poleward of 34°
latitude. Along the Gulf coast, their failure to become well established east of Texas has been
attributed to the higher annual precipitation in the Gulf coast states (greater than 50 in. or
1,270 mm a year) and possibly to greater rates of parasitism by varroa mites (Varroa destructor;
see Arachnids, Varroa Mite) in those more humid areas.
Diet. Africanized honey bees consume nectar and pollen. Nectar is converted to honey
and becomes the bees’ main source of carbohydrates. Pollen constitutes the main source of
protein. Most flowering plants serve as food sources.
Life History. A queen mates with a drone to produce a fertilized egg, which develops into
a female worker bee. Unfertilized eggs become male drones. If larvae are fed royal jelly—a
mixture of glandular secretions—they develop into queens. The life cycle involves egg, larva,
pupa, and adult. Eggs are attached to the bottom of cells in the comb and hatch in approx-
imately 60 hours. The larvae are then fed secretions from the heads of young nurse bees. If
the egg is to become a worker, it is sealed into the cell eight days after hatching. Within
24 hours, the larva spins a cocoon, and the next day enters the prepupal stage. A day later,
the white, immobile pupa has formed. Adult worker bees emerge 18.5 days after the egg
was laid; queens about 16 days after laying.
Worker bees live about 30 days, drones 5–10 weeks, and queens 1–3 years.
As a colony grows, it will split into two or more colonies. A queen and a group of workers
leave the nest together in a “swarm.” A new queen hatches in the original nest, mates, and
begins egg-laying. Africanized bees swarm frequently (as often as every six weeks) and at
any time of year. Their survival strategy, evolved in an African cultural environment in
which people were and still are bee hunters and honey thieves rather than beekeepers, is
to invest energy into large numbers of offspring and produce many reproductive swarms.
They fiercely defend their nests, but they also abandon them readily and start new colonies
elsewhere.
Impacts. The most direct impacts of Africanized honey bees are on the beekeeping indus-
try and agriculture. These hybrid bees compete with domestic European honey bees and, by
mating with European queens, take over hives. The Africanized bees not only are more

A. The Africanized honey bee is a hybrid species and can only be distinguished from the European honey bee by
laboratory analysis. (Jeffrey W. Lotz, Florida Department of Agriculture and Consumer Services, Bugwood.org.)
B. Africanized honey bee (left) and European honey bee (right). Normally color differences do not allow
distinction between the two. (Scott Bauer, USDA Agricultural Research Service, Bugwood.org.)
 AFRICANIZED HONEY BEE n 109

difficult to handle and much more apt to abandon their hives, but they also produce less honey
than European honey bees. Honey bees are extremely important pollinators of many orchard
and field crops, and beekeeping is a commercial enterprise of its own. One estimate has honey
bees adding $10 billion or more each year to the value of some 90 crops in the United States.
The problem in handling the aggressive hybrid bees has led some to give up beekeeping
altogether and others to adopt more labor-intensive and therefore costly management practices
(see under Management). Africanized honey bees are best suited to subtropical areas of the
United States, the very states where the packaged bee and queen industry is located.
Africanized honey bees are dangerous to people who venture too close to their nests or oth-
erwise disturb them because of the multiple stings victims receive from the hundreds of bees that
attack en masse at the slightest provocation. Their preferred nesting locations bring them into
close proximity to humans, increasing the likelihood of bee-human interaction. The bees can
sense intruders more than 50 ft. (15 m) away and be irritated by the vibrations of power equip-
ment more than 100 ft. (30 m) distant. They may chase people whom they feel are threats for
0.25 mi. (0.4 km). Some people have died from as few as 100–300 stings, but usually it takes
500–1,000 stings for the attack to be fatal. Livestock and pets are at risk as well as humans.
Effects on natural ecosystems and native pollinators are unknown. Competition from
Africanized honey bees could displace other nectar- and pollen-feeding invertebrates from
their more important food resources.
Management. As the invasion spread through South and Central America, beekeeping
and honey production suffered significant declines at first. Then beekeepers learned new
skills to better manage their Africanized hives and reduce the frequency of bee attacks.
Among the benefits to agriculture of these new practices was the production of pollinators
many farmers feel are superior to the European honey bee. A simple change was to keep
each bee colony on its own, separate hive stand so that one hive could be worked without
disturbing neighboring colonies. More smoke is used well ahead of working with the bees
to calm them and possibly mask alarm pheromones. In addition, a change from traditional
black bee veils to white ones as part of the beekeeper’s protective gear minimizes attacks.
In the United States, the strategy has been to try to diminish the infusion of African genes
into bee populations by either “drone-flooding” or frequent requeening with known
European queens. Drone-flooding involves adding European drones to a colony to lessen
the chances of the queen mating with an Africanized one.
To avoid painful or even deadly encounters with Africanized bees, people in areas where
these bees now occur should remain alert to their potential presence when outdoors and stay
away from known nests or swarms. Anyone knowing the location of hives or swarms should
obtain professional help to remove them. If an attack occurs, immediately run away in a
straight line, covering your face if possible, and seek shelter in a car or building. Do not swat
at them or try to hide in a pool of water. Scrape stingers off the skin; do not squeeze them, as
that will cause them to release more venom. Wash and apply ice to stings and see a doctor if
breathing becomes labored. People can bee-proof their homes and yards by sealing open-
ings, regularly inspecting eaves, and removing other potential nest sites.

Selected References
“Africanized Honey Bee.” Oklahoma Invasive Species Site. Oklahoma State University, 2007. http://
oklahomainvasivespecies.okstate.edu/africanized_honey_bee.html.
“Africanized Honeybee Pest Profile.” California Department of Food and Agriculture, 2010. http://
www.cdfa.ca.gov/phpps/pdep/target_pest_disease_profiles/ahb_profile.html.
110 n INVERTEBRATES (INSECTS)

Ellis, Jamie, and Amanda Ellis. “African Honey Bee.” Featured Creatures, Publication Number EENY-
429, University of Florida Institute of Food and Agricultural Sciences, 2008 http://
entomology.ifas.ufl.edu/creatures/misc/bees/ahb.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialty Group
(ISSG). “Apis mellifera scutellata (Insect).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=325&fr=1&sts.
Sanford, Malcolm T., and H. Glenn Hall. “African Honey Bee: What You Need to Know.” Fact Sheet ENY-
114, Entomology and Nemotology Department, Florida Cooperative Extension Service, Institute of
Food and Agricultural Sciences, University of Florida, 2005. http://edis.ifas.ufl.edu/mg113.
Villa, J. D., T. E. Rinderer, and J. A. Stelzer. “Answers to the Puzzling Distribution of Africanized Bees
in the United States or ‘Why Are Those Bees Not Moving East to Texas?’ ” American Bee Journal
142(7): 480–83, 2002. http://www.ars.usda.gov/research/publications/Publications.htm
?seq_no_115=133427.

n Argentine Ant
Scientific name: Linepithema humile
Synonym: Iridomyrmex humilus
Family: Formicidae
Native Range. South America: The Argentine ant is native to northern Argentina,
southern Brazil, Uruguay, and Paraguay. Genetic studies reveal that those ants in the
continental United States derive from the southern Paraná River region of Argentina.
Distribution in the United States. Argentine ants are widely established in California.
They are also found in southern Arizona. In the Gulf coast states, they are common from east
Texas to Florida. They have also been reported in Georgia, South Carolina, and North
Carolina. In Hawai’i, they are spreading through Haleakala National Park on Maui.
Populations also exist on the Big Island.
Description. The Argentine ant is medium to dark brown and about 0.1 in. (2–3 mm)
long. The workers all have the same morphology. Their bodies are smooth and shiny with-
out hairs on the back of the head or thorax. The petiole or waist segment consists of a single
upright scale. When viewed head-on, the eyes do not protrude beyond the outline of the
face. These ants lack stingers or acidopores. They move quickly along strong foraging trails,
often in large numbers. When crushed, they give off a musty odor.
Related or Similar Species. Most other ants give off an acidic odor when crushed.
Introduction History. The Argentine ant entered the United States at New Orleans in the
early 1890s. The first record dates to 1891. It probably arrived in shipments of coffee or sugar
from Argentina and may have spread across the southern United States on trains. It was first doc-
umented in California in 1907. Reintroductions to the Southeast may have occurred in the early
1900s in cargo arriving at Gulf ports from California. Argentine ants came to Hawai’i during
World War II, presumably with shipments of goods on troop ships from California.
Local dispersal may be facilitated by the ants’ predilection to move their nests frequently.
Therefore, they can rapidly take advantage of new opportunities for new nest sites in potted
plants or in refuse, items that humans will further disperse. They also “raft” when their hab-
itat is flooded and thereby move to new areas. New colonies can be established with as few
as 10 workers and a queen.
The invasive success of Argentine ants has been attributed to the genetic similarity of all
individuals, a consequence of either just one or very few colonizing events. The argument
has been that the ants fail to recognize individuals born of different queens as “others,” so
all can live together in massive supercolonies formed by budding. In other words, they lack
 ARGENTINE ANT n 111

population-control measures
that would have resulted from
competition among colonies.
Others maintain that the reason
supercolonies in areas to which
the Argentine ant has been
introduced are so much larger
than in their native range is
due to a reduced parasite load
and reduced competition from
other ant species in the intro-
duced range. Those two factors
allow supercolonies to survive
longer and attain great sizes, an
aid when invading new territory.
Habitat. Argentine ants pre-
fer moist environments with
moderate temperatures. They
can be found in a variety of
natural, disturbed, and man-
made habitats, from agricultural
areas (especially citrus orch-
ards) to natural forests and
grasslands, and in urban areas
both indoors and outdoors. In
drier regions, such as southern
California, they prefer to be near
perennial streams or irrigated
fields and human dwellings.
They are the most common
“house ant” in California, par-
ticularly in urban San Diego Top: Argentine ants are native to the Paraguay-Paraná River basin of
and Los Angeles. Usually re- northern Argentina, southern Brazil, Paraguay, and Uruguay. Bottom:
stricted to lower elevations, they Argentine ants are established in California and the Gulf coast states; dis-
are invading alpine areas in junct populations occur north of these areas. In Hawai’i, they occur on
Hawai’i. Argentine ants nest in Mau’i and the island of Hawai’i. (Adapted from “Sources of Introduced
Argentine Ants Tied to Reduced Genetic Variation.” University of
mulch or soil, in rotting wood,
California, San Diego, news release, n.d. http://ucsdnews.ucsd.edu
and in garbage. Nests are often ?rewsrel/science/mcants.htm.)
found in soil under objects or
near tree roots and in potted
plants or under walkways.
Diet. This ant prefers sweet substances. It tends aphids and consumes the honeydew that
these and other insects such as mealybugs and scales secrete. It also feeds on nectar and the
body fluids of dead animals. Occasionally, it will capture small, slow-moving insects or take
their eggs. In homes it seeks sugar, proteins, and water.
Life History. Queens and drones mate in the nest, unlike many ants that perform nuptial
flights. All worker ants are sterile. Nonetheless, worker ants can rear eggs and direct the
development of early instar larvae into reproductive females without queens being present.
112 n INVERTEBRATES (INSECTS)

A colony expands and disperses

by budding new subcolonies.
The result can be a huge super-
colony with many queens. The
production of males apparently
is determined by the quantity
of food available to larvae.
Workers may kill the queen
after a year and accept a newly
mated queen in her place.
Impacts. Argentine ants are
aggressive and have displaced
native ant species in various
parts of the world. In California,
local extinctions of natives such
as army ants (Neivamyrmex
spp.) and harvester ants (Messor
The petiole or waist of the Argentine ant has a single upright scale, clearly spp. and Pogonomyrmex spp.)
visible in this photograph. (Eli Sarnat.) have occurred.
Ants and other native insects
are often important seed dispersers and plant pollinators. If the natives are severely reduced
by invading Argentine ants, native plants may also decline. This is a major concern in
Haleakala National Park, Hawai’i, where Argentine ants are encroaching upon the habitat of
the endemic giant silversword (Argyroxiphium sandwicense), which is pollinated by yellow-
faced bees (Hylaeus spp.). It is feared ant predation on bee larvae will decimate the native bee.
Competition between Argentine and native ants for food and habitat in California has
affected the coastal horned lizard (Phrynosoma coronatum), which has a diet specializing on
native harvester and carpenter ants. The loss of native ants has contributed to the decline
of this once-common lizard. Decreases in the numbers of the native desert shrew
(Notiosorex crawfordi) are also attributed, in part, to the presence of the Argentine ant.
Argentine ants can be serious pests in commercial apiaries. Ants invade honey bee (Apis mel-
lifera) hives, causing the bees to abandon it. The ants then remove the honey and bee larvae to
their own nest. They can destroy 1–2 hives a day and soon overwhelm an entire bee yard.
This ant has been nominated as by IUCN as among “100 of the ‘World’s Worst’ invaders.”
Management. Once a colony is established, it is difficult if not impossible to eradicate it.
Chemical controls such as DDT, chlordane, and dieldrin that once helped control ant pop-
ulations are no longer legal to use in the United States. Bait traps can help, since the poison
works slowly and ants bring it back to their nests, where it will kill all workers and queens.
Prevention, through the inspecting of cargo and refuse, sealing cracks and other points of
entry to buildings, and maintaining landscapes that offer no nest sites, coupled with swift
action to eliminate any new infestations, is the most effective management strategy to limit
the further spread of the Argentine ant.

Selected References
Krushelnycky, Paul, Andrew Suares, and IUCN/SSC Invasive Species Specialty Group (ISSG).
“Linepithema humile (Insect).” ISSG Database, 2006. http://www.issg.org/database/species/
ecology.asp?fr=1&si=127.
 ASIAN LONGHORNED BEETLE n 113

Sarnat, E. M. “Linepithema humile.” PIAkey: Identification Guide to Ants of the Pacific Islands, Edition
2.0, Lucid v. 3.4. USDA/APHIS/PPQ Center for Plant Health Science and Technology and
University of California–Davis, 2008. http://keys.lucidcentral.org/keys/v3/PIAkey/Fact_Sheets/
Linepithema_humile.html.
Spring, Joe. “Argentine Ant (Linepithema humile) (Mayr, 1868).” Introduced Species Summary Project,
Columbia University, 2004. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/
inv_spp_summ/Linepithema_humile.html.
Westervelt, David, and Eric T. Jameson. “Argentine Ant, Liniepithema humile Mayr (Hymenoptera:
Formicidae).” Florida Department of Agriculture and Consumer Services, 2009. http://
www.doacs.state.fl.us/pi/pest_alerts/liniepithema_humile.html.

n Asian Longhorned Beetle

Also known as: ALB
Scientific name: Anoplophora glabripennis
Order: Coleoptera
Family: Cerambycidae
Native Range. East Asia: It is widely distributed in China from Sichuan and Gansu prov-
inces north and east to the vicinity of Beijing. It is also found on the islands of Hainan and
Taiwan, throughout Korea, and in Japan.
Distribution in the United States. The major infestations of Asian longhorned beetles are
in urban trees in Brooklyn, Queens, Manhattan, Long Island, Staten Island, and Prall’s
Island, New York; and in suburban Chicago. They have also been found in Hoboken,
Jersey City, and Middlesex/Union counties, New Jersey; and in and around Worcester,
Massachusetts.
Description. The adults are large, glossy black beetles with irregular white spots on the
wing covers (elytra). The scutellum, a small triangular plate between the attachment points
of the forewings, is black. The antennae are longer than the body and marked with distinct
black and white bands. Their bodies are 0.75–1.0 in. (20–35 mm) long and 0.25–0.5 in.
(6–12 mm) wide. Adults are seen in the open from late spring into fall.
The presence of Asian longhorned beetles is often recognized by signs on infested trees.
Shallow oval borings in the bark of trees ooze sap. Sawdust-like debris collects at the base
of trees or in the crotches of large branches. Circular holes the width of an adult are the exit
holes of newly emerging beetles.
Eggs are off-white and oblong, with slightly concave ends. They are 0.2–0.3 in. (5–7 mm)
long. Larvae are pale yellow elongate grubs that reach lengths of nearly 2.0 in. (50 mm) before
pupating. The pupae are off-white and about 1.2–1.3 in. (30–33 mm) long and 0.4 in.
(11 mm) wide. All of these life-cycle stages are hidden deep inside living trees.
Related or Similar Species. The whitespotted sawyer (Monochamus scutellatus), a native
beetle found in most forested areas of the United States, is about the same size as the Asian
longhorned beetle and also has long antennae. The body, however, is a dull or bronzed
black, and the antennae are only faintly banded. The scutellum is white. Larvae are very sim-
ilar to those of the Asian longhorned beetle. Whitespotted sawyers infest conifers, whereas
Asian longhorned beetles prefer hardwoods.
Another large native long-horned beetle is the boldly black-and-white cottonwood borer
(Plectrodera scalator) of the eastern United States and the Great Plains. Its antennae are solid
black and equal in length to the body. Larvae dwell in and near the roots, and pupation takes
place below ground level in roots; therefore, no exit holes appear in tree trunks or branches.
114 n INVERTEBRATES (INSECTS)

Introduction History. Asian

longhorned beetles were first
reported in the United States in
the Greenpoint section of
Brooklyn, New York, in 1996,
although they may have been
present but undetected for
10 years before then. They
probably arrived in wooden
pallets or other solid-wood
packing materials with pipes
that were brought in from
China in the late 1980s for a
sewer project. From Brooklyn,
they spread to Amityville on
Long Island, Queens, and Man-
hattan. The introduction to
Long Island was the result of a
tree care company inadvertently
disposing contaminated wood
from Brooklyn at several loca-
tions. A separate introduction
gave rise to the population in
Chicago, first reported in 1998.
Asian longhorned beetles
showed up in Jersey City, New
Jersey, in 2002 and had expan-
ded their distribution to other
towns (Woodbridge and Rah-
way) by 2004. They appeared
on Staten and Prall’s islands,
Top: Asian longhorned beetles are native to eastern China, Korea, Japan, New York, in 2007, and in
and Taiwan. Bottom: Asian longhorned beetles infest urban trees in and Worcester, Massachusetts, in
around Chicago, Illinois; the boroughs of New York City; New Jersey; 2008. Live beetles continue to
and Worcester, Massachusetts. be discovered at ports and ware-
houses in various states.
Local dispersal from primary sites of infestation is most likely aided by the transport of
infested firewood by vacationers, hunters, and others. Larvae hidden deep inside trees may
be accidentally moved in live trees and cut timber as well. On their own, Asian longhorned
beetles disperse very slowly. They were found exclusively on trees in cities and towns until
2008, when they were discovered in natural forests outside Worcester, Massachusetts.
Habitat. Asian longhorned beetles usually infest urban broadleaf deciduous trees, but
since 2008, they have been found in natural hardwood forests.
Diet. Asian longhorned beetles differ from most other members of their family by eating
the living wood of at least 18 species of broadleaf tree. (Most temperate-zone long-horned
beetles have one preferred host and feed on dead and dying trees.) Maples (Acer spp.)
are preferred, but Asian longhorned beetles have been reported to attack horse-chestnuts
and buckeyes (Aesculus spp.), willows (Salix spp.), ash (Fraxinus spp.), birches
 ASIAN LONGHORNED BEETLE n 115

A. Adult Asian longhorned beetle. (Donald Duerr, USDA Forest Service, Bugwood.org). B. Asian longhorned
beetle larvae. (Michael Bohne, Bugwood.org.)

(Betula spp.), and sycamores (Platanus spp.). Larvae eat healthy cambium, phloem, and
xylem, forming long tunnels in the sapwood and heartwood of host trees.
Life History. In both New York and Chicago, adults reportedly emerge from pupae
between July and October and bore their way through the bark of the host tree. This activity
leaves telltale piles of frass and dime-size circular holes in the bark. The beetles usually fly
only 150–250 ft. (50–70 m) away from the tree where they hatched. They feed on the bark
of twigs on a new host tree for 2–3 days and then mate somewhere on the trunk or branches.
The female chews a shallow, oval cavity into the cambium in which she lays a single egg; she
will repeat this process 25–90 times. In 10–15 days, the eggs hatch into larvae that feed at
first in the cambium but soon tunnel 4–12 in. (10–30 cm) deep into the tree. The larvae
metamorphose into pupae and, about 18 days later, into adults inside the tunnels. Eggs, lar-
vae, and pupae are able to overwinter if necessary. Adults remain active until late fall and
then die. The life cycle is completed in 12–18 months.
Impacts. Large numbers of larvae infesting a tree weaken the host. Their tunneling girdles
the tree, and dieback of the crown becomes noticeable. Weakened branches can break in
strong winds. Ultimately, the tree dies.
The egg-laying cavities and exit holes joined to a gallery of tunnels inside the tree also
open the host tree to infections and infestations of other kinds. Control efforts have resulted
in the destruction of thousands of shade and ornamental trees in New York and Chicago.
There is concern that if let loose in the broadleaf deciduous forests of New England, the
Asian longhorned beetle could become a pest on the scale of gypsy moths (see Insects,
Gypsy Moth) or chestnut blight (see Fungi, Chestnut Blight Fungus). Economic impacts in
urban areas—where the beetles could rival Dutch elm disease (see Fungi, Dutch Elm
Disease) in changing urban landscapes—involve diminished aesthetics and declining prop-
erty values. Timber, maple syrup, nursery, and tourist industries are threatened in New
England and elsewhere.
Management. The chief legal and effective way in the United States to deal with urban
infestations is to cut down all trees known to host the beetle, chip and burn the wood,
and grind out the stump. The trees that are removed are replaced with nonhost species.
Infested areas are quarantined, and firewood, tree trimmings, and other tree by-products
116 n INVERTEBRATES (INSECTS)

are prohibited from being transported out of the area. Contact insecticides are of little value,
because much of the life cycle of the beetle is spent inside the tree where sprays cannot reach
it. Systemic insecticides that can be taken up by the roots and distributed into stems and foli-
age have some potential, as does the application of parasitic fungi in bands on tree trunks to
control adult beetles.
The Animal and Plant Health Inspection Service (APHIS) of the U.S. Department of Agriculture
has inspectors at key ports of entry to check for beetle-infested cargo, targeting especially ship-
ments from China with solid-wood packing materials. Since 1998, all such material is required
to be treated prior to shipment to the United States, and compliance has been exceptionally high.
Federal quarantines were imposed in Chicago, New York, and New Jersey. In quarantine
areas, only arborists certified to remove Asian longhorned beetles are allowed to cut down
infested trees. No wood from the area can be used, moved, or disposed of without an inspec-
tion permit. New trees to be planted should be nonhost species. The quarantine zones in
Hoboken and Jersey City, New Jersey, were deregulated in 2005; and the quarantine in
Chicago was lifted in 2006. However, the quarantine area in the Middlesex/Union County
area of New Jersey was expanded when new trees infested with the beetle were discovered.
In July 2008, a new regulated area was established in Worcester, Massachusetts, that covers
62 sq. mi. (160 km2). Visual surveys of trees and other means of early detection of and rapid
response to new infestations are underway to attempt to prevent the spread of this pest into
the natural forests of Massachusetts and surrounding states.

Selected References
“Asian longhorned beetle.” University of Vermont, 2005. http://www.uvm.edu/albeetle/index.html.
Cavey, Joseph F. “Anoplophora glabripennis.” National Information Center for State and Private Forestry,
U.S. Forest Service, 2000. http://spfnic.fs.fed.us/exfor/data/pestreports.cfm?pestidval=53&lang
display=english.
Muruetagoiena, Tamara. “Asian Longhorned Beetle (Anoplophora glabripennis).” Introduced Species
Summary Project, Columbia University, 2004. http://www.columbia.edu/itc/cerc/danoff-burg/
invasion_bio/inv_spp_summ/Anoplophora%20glabripennis.html.

n Asian Tiger Mosquito

Scientific name: Aedes albopictus
Synonym: Stegomyia albopictus
Order: Diptera
Family: Culicidae
Native Range. Asia. The Asian tiger mosquito occurs naturally from tropical Southeast
Asia north through China, South Korea, and Japan, and on islands in the Pacific and
Indian oceans, including Madagascar.
Distribution in the United States. Asian tiger mosquitoes are established in 26 states in the
continental United States and in Hawai’i. Primarily a species of the Southeast, discontinuous
outlying populations are recorded as far north as Minnesota and as far west as Nebraska,
Kansas, Oklahoma, and Lubbock and Val Verde counties in Texas. On the East Coast, the
northern limit is currently New Jersey, but that could change with a warming climate.
Description. The adult is a small mosquito with a black body and conspicuous silvery-
white bands on the legs and feelers (palpus). A single white stripe runs along the back and
 ASIAN TIGER MOSQUITO n 117

head. Males are smaller than

females and have feathery anten-
nae and mouthparts modified
for nectar-feeding. The female
has unadorned feelers and a long
proboscis adapted to biting and
feeding on blood. As in all mem-
bers of the genus Aedes, the
abdomen narrows to a point.
Adults are a little less than 0.25
in. (5 mm) long.
The wormlike larvae, or “wig-
glers,” are easily disturbed by
vibrations or a passing shadow
and quickly descend to the bot-
tom of the water container in
which they live. They must
periodically rise to the water
surface to obtain oxygen through
a somewhat inflated-looking
breathing siphon. Fully grown
wrigglers are about 0.25 in.
(5 mm) long. The pupae are dark
brown and curled into the shape
of a comma. When disturbed,
these “tumblers” roll end over
end through the water.
Related or Similar Species.
The yellow fever mosquito,
Aedes eagypti, like the Asian tiger
mosquito, is active by day and
breeds in water held in contain- Top: The Asian tiger mosquito’s native range extends from Southeast Asia
ers. Its hind legs also bear white to Japan and includes islands in the Pacific and Indian oceans. (Adapted
bands. In the field, the yellow from map by Gando, http://en.wikipedia.org/wiki/File:Albopictus
fever mosquito is distinguished _distribution_2007.png.) Bottom: Asian tiger mosquitos are currently
by the lyre-shaped pattern of established in 27 states. (Adapted from “Distribution of Aedes albopictus
in the United States, by County, 2000.” Centers for Disease Control and
white scales on its back and the
Prevention, Division of Vector-borne Infectious Diseases. www.cdc.gov/
white scales on the head of the ncidod/dvbid/arbor/albopic_new.htm.)
female on a structure above its
proboscis. In female Asian tiger
mosquitoes, this structure is black. The yellow fever mosquito originated in Africa and probably
came to the New World on ships in the early days of trans-Atlantic trade and exploration.
Introduction History. Asian tiger mosquitoes probably arrived in Hawai’i soon after
World War II. Their first verified occurrence in the continental United States, indeed
in the New World, came in 1985 in Houston, Texas. It was accidentally transported
from Japan in used tires imported for recapping. During the 1980s, importations of
used tires increased, and Asian tires were preferred because of their high rubber content.
They arrived in containers that were inadequately inspected at the ports of entry. The first
118 n INVERTEBRATES (INSECTS)

discovery of the mosquito in

Maryland occurred in 1987 at
a tire processing plant in
Baltimore. The movement of
used tires within the United
States dispersed the insect to
other eastern and midwestern
states. Used tires are typically
stored outdoors for long peri-
ods of time, and those that can-
not be used for recapping often
end up in illegal tire dumps. In
both instances, they collect
rainwater and become prime
Asian tiger mosquito (male). (Susan Ellis, Bugwood.org.) Asian tiger mosquito breeding
grounds.
In California, Asian tiger
mosquitoes were discovered in 2001 in containerized shipments of lucky bamboo
(Dracaena sp.) from southern China. The popular ornamental plants had been transported
in standing water so they would survive the long voyage.
The mosquitoes have been locally dispersed in moist vegetation and water containers,
including cemetery flower pots. In several Florida counties, the mosquitoes were first found
in cemeteries where fresh-cut flowers were placed at grave sites in plastic floral baskets.
When the flowers died, they were discarded; however, the baskets were reused, often at dif-
ferent cemeteries. Eggs laid when flowers were fresh and standing in water in one cemetery
would hatch later in another when water was again added.
Habitat. The Asian tiger mosquito likely evolved in forests where it could breed in tree-
holes and in epiphytes and other places where water pooled. It still prefers the shade of
densely vegetated areas, but exploits artificial containers in urban and suburban areas for
breeding sites. It reproduces in bird baths, cemetery flower pots, empty soda cans, and other
abandoned containers that hold water, but it is associated especially with old tires that
collect water and organic-rich debris. The water only needs to be 0.25 in. (5–6 mm) deep.
Northern range limits may be set by cold temperatures (daily mean temperatures in
January of 32°F or 0°C) that prevent overwintering of eggs. Initially in the United States,
the southern limit was established by summer day lengths shorter than 13.5 hours, but
the species has overcome that controlling factor.
Diet. Asian tiger mosquitoes acquire energy by feeding on nectar. The females, however,
need protein to produce eggs, and they feed also on blood. Humans are the chief host, but
the females will feed on a variety of other mammals including dogs and cats, and on birds,
reptiles, and amphibians. They generally feed at ground level, biting people around the
ankles and knees.
The larvae are filter-feeders, consuming particulate organic debris and bacteria.
Life History. Mating occurs soon after adults emerge from pupae. The female mates only
once and stores sperm for future egg production. A female feeds on blood for 4–5 days
before laying eggs. She then deposits 40–150 eggs on the sides of containers just above water
level. She will continue a weeklong cycle of feeding on blood and laying eggs for the
 ASIAN TIGER MOSQUITO n 119

remainder of her life. The eggs do not hatch until water level rises and covers them. Eggs
may overwinter and not hatch until the following spring or summer, when they become
flooded by water warmer than 60° F (15.5° C). The wriggling larvae develop through four
instars and change into pupae about 10 days after hatching. Unlike many other insects, mos-
quito pupae are mobile, but they do not feed. Adults emerge from the pupae 10–14 days
later. Adult mosquitoes live several days to a few weeks depending upon the weather. The
lifespan is reduced during hot, dry spells. During their lives, adults rarely move more than
a few hundred yards (or meters) from the container in which they hatched.
Impacts. The arrival of the Asian tiger mosquito raised major concerns of its potential as a
vector for virus-based dengue fever. Dengue fever is a disease of the tropics. It once posed a
sporadic health problem in southern states, but was eradicated in the United States in the
1940s. However, dengue fever is common in the Caribbean and tropical South and
Central America, and could easily be reintroduced. A warming climate might enhance its
ability to spread. It now appears that the Asian tiger mosquito is less likely than its close rel-
ative, the yellow fever mosquito, to transmit the dengue fever virus from human to human
and cause major outbreaks of the disease. This is because the Asian tiger mosquito has
multiple mammalian, avian, and reptilian hosts and is not restricted to a diet of human
blood as is the yellow fever mosquito. If, after biting a human, the mosquito moves on to a
bite another type of host, transmission of the virus is halted.
In laboratory studies, the Asian tiger mosquito has proved to be a competent vector for a
number of other viruses, including eastern equine encephalitis, yellow fever, West Nile disease
(see Microorganisms, West Nile Virus), LaCrosse encephalitis, St. Louis encephalitis, and
western horse encephalitis. It is also a carrier of the nematode known as canine heartworm.
Field populations have been found to carry eastern equine encephalitis virus and West Nile
virus, but so far, no evidence exists that this mosquito has transmitted disease to any person.
The Asian tiger mosquito’s greatest impact has been as an extreme nuisance. It is much
more annoying than the yellow fever mosquito. An aggressive and persistent biter that feeds
during the day, it tends to rest in shrubs in shady areas, where people also seek relief from
the summer sun. The female interrupts her feeding to bite the same person several times
and, because of her agility, is difficult to swat. She injects an anticoagulant from their sali-
vary glands that causes the bite to swell and itch. Sometimes the itch may last for a week.
The Asian tiger mosquito has displaced the yellow fever mosquito from many areas in
which the distribution areas of the two overlapped.
Management. The best defense against the Asian tiger mosquito is for individuals to
remove potential breeding sites, such as old tires, empty buckets, or other containers that
could fill with rainwater. Bird baths and wading pools should be emptied at least once a
month. Roof gutters should be kept free of debris that would dam up water. The larvae
can be killed with applications of the bacterium Bti (Bacillus thuringiensis israelensis). Adults
are not readily trapped by contrivances that attract other mosquitoes, and they are resistant
to many insecticides.
Biological controls include stocking predators of the larvae such as mosquitofish
(Gambusia spp.; but see Fish, Mosquitofish) and developing a genetically engineered male
that will pass a lethal gene to its female offspring.
Since 1988, the U.S. Public Health Service has required that all used tires imported into
the United States from areas known to have Asian tiger mosquitoes be dried, cleaned, and
fumigated.
120 n INVERTEBRATES (INSECTS)

Selected References
“The Asian Tiger Mosquito in Maryland.” Maryland Department of Agriculture, n.d. http://
www.mda.state.md.us/plants-pests/mosquito_control/_asian_tiger_mosquito_md.php.
Crans, Wayne J. “Aedes albopictus (insect).” ISSG Database, IUCN/SSC Invasive Species Specialist
Group (ISSG), 2009. http://www.issg.org/database/species/ecology.asp?si=109&fr=1&sts.
Crans, Wayne J. “The Asian Tiger Mosquito in New Jersey.” Fact Sheet. Rutgers Cooperative Research
and Extension, New Jersey Agricultural Experiment Station, Rutgers, the State University of New
Jersey, 1996. http://njaes.rutgers.edu/pubs/download-free.asp?strPubID=FS845.
O’Meara, G. F. “The Asian Tiger Mosquito in Florida.” Entomology and Nematology Department,
Cooperative Extension Service, Institute of Food and Agricultural Sciences, University of Florida,
2005. http://edis.ifas.ufl.edu/mg339.
Rios, Leslie, and James E. Maruniak. “Featured Creatures: Asian Tiger Mosquito.” University of Florida,
2008. http://entnemdept.ufl.edu/creatures/aquatic/asian_tiger.htm.

n Brown Marmorated Stink Bug

Also known as: Yellow-brown stink bug; BMSB
Scientific name: Halyomorpha halys
Family: Pentatomidae
Native Range. East Asia: China, Japan, Korea, and Taiwan
Distribution in the United States. This rapidly spreading bug was reported in California,
Delaware, Maryland, Missouri, New Jersey, New York, North Carolina, Oregon,
Pennsylvania, Tennessee, Virginia, West Virginia, and the District of Columbia by 2010.
Description. Adults have the shield body shape typical of all stink bugs. The back is a
marbled (marmorated) brown, and the underside a lighter tan color. Small round depres-
sions on the head and first segment of the thorax (the pronotum) are coppery or bluish met-
allic in color. The edges of the pronotum are smooth, unlike the jagged margin of native
stink bugs of the genus Brochymena. The next-to-last segment of the antennae bears pale
bands, a distinguishing characteristic. The abdominal segments that extend beyond the
wings have alternating bands of black and white; and the brown legs have faint light bands.
Scent glands are located between the first and second pair of legs on the underside of the
thorax. Adults are about 0.5 in. (12–17 mm) long and almost as wide.
Immatures or nymphs develop through five stages or instars. The first instars
have yellowish-red abdomens with black bars and dark red eyes; they look like ticks and
are about 0.09 in. (2.4 mm) long. Relatively inactive, they tend to remain clustered near
the hatched egg mass. Successive instars become larger and more closely resemble adults.
Their legs and antennae are black with white bands. The final, fifth instar has a whitish
abdomen with red spots and is about 0.47 in. (12 mm) long. Knobs protrude in front of
the scent glands on the underside, and spines occur on the legs, in front of the eyes, and
on the sides of the thorax.
The pale green eggs are elliptical and measure about 0.04 in. (1 mm) in diameter. They
are usually found on the underside of leaves in masses of 20 to 30.
Related or Similar Species. Brown marmorated stink bugs are easily confused with native
species of stink bug, including the brown stink bug (Euschitus servus) and the green stink
bug (Acrosternum hilare); with rough stink bugs (Brochymena spp.); and some leaf-footed
or squash bugs (Family Coreidae). The Western conifer seed bug (Leptoglossus occidentalis)
also invades buildings in the autumn in order to winter over, but it can be distinguished
from the BMSB by the flattened structures on the lower segments of its hind legs.
 BROWN MARMORATED STINK BUG n 121

Introduction History. This

bug was accidentally intro-
duced in packing materials
to Allentown, Pennsylvania,
where it was first collected in
September 1998. It may have
been present for a couple of
years before it was discovered
and identified. In 2000 and
again in 2002, it was collected
in New Jersey and, soon after-
ward, appeared in Delaware,
Connecticut, and New York. In
2005, an established popula-
tion was documented in
Oregon. The stink bug has be-
come established in many pla-
ces on the East Coast of the
United States and continues to
expand its range. By 2009, it
was established in Maryland,
Virginia, and West Virginia
and had been collected in
California, Florida, Mississippi,
Missouri, North Carolina, Ohio,
Tennessee, and Washington,
D.C. It is suspected that they are
moving across the country in
motor vehicles.
Habitat. Originally, popula-
tions were restricted to urban
and suburban landscapes where Top: The brown marmorated stink bug is native to East Asia. Bottom:
they were associated with orna- This new invader was reported in 12 states and the District of Columbia
mental plants, garden crops, by 2010.
and fruit and shade trees. They
have since expanded into agricultural areas, including orchards. Brown marmorated stink
bugs seek winter shelter in buildings, entering in the fall through any small opening. They
remain dormant much of the winter and leave houses and other buildings in the spring to
reproduce and forage outside.
Diet. These bugs are sucking insects that feed on a wide variety of plants. They pierce
fruit and leaves with specially adapted mouthparts and withdraw plant juices. Nymphs feed
on fruits and seed pods. They are known to eat apples, peaches, pears, blackberries, toma-
toes, green peppers, beans, sweet and field corn, and soybeans. Among ornamentals, they
attack shade trees such as Norway maple (Acer platanoides) and ornamentals such as princess
tree (Paulownia tomentosa), butterfly bush (Buddleia spp.), Rugosa roses (Rosa rugosa), and
honeysuckle (Lonicera sp.)
Life History. Mating takes place in early summer, approximately two weeks after adults
emerge from hibernation. Egg-laying begins in early summer and will continue through
122 n INVERTEBRATES (INSECTS)

September in the Mid-Atlantic

states. The female lays new egg
masses on the undersides of
leaves about once a week
throughout the summer. The
eggs hatch in four or five days
and first instar nymphs usually
stay near their egg mass until
after the first molt. Each nym-
phal stage lasts about one week
and is followed by a molt that
produces the next instar.
About two weeks after the final
molt, the adults are sexually
mature. Only one generation
appears to be produced each
The brown marmorated stink bug is rapidly expanding its range year. However, since several
in the eastern United States. (David R. Lance, USDA APHIS PPQ, generations are produced each
Bugwood.org.) year in the subtropical regions
of its native range, the same
may be expected when the stink bug becomes established in the Southeast. Adults live sev-
eral years, and a single female may lay up to 400 eggs during her lifetime.
Impacts. Brown marmorated stink bugs are primarily nuisances in homes and other
buildings, where they congregate in autumn and then overwinter. On warm days, they
may rouse from hibernation and fly around, and in the spring, they fly to windows and
doorways as they try to leave the premises. The bugs do no structural damage, do not lay
eggs inside buildings, and are not known to carry any diseases. However, when crushed,
they secrete a most unpleasant odor from their scent glands.
The stink bug can damage ornamental and garden plants and is becoming an agricultural
pest in apple and peach orchards. Their feeding methods causes puckering, scarring, and
deformation of fruits—a condition known as cat facing, which renders them unfit for sale
as fresh market items. The fruits may still be processed, however, for juices, jams, sauces,
and the like. Leaf-stippling will occur on leaves. The spread of the bug could threaten com-
mercial apple, peach, and soybean crops. Damage to orchard crops has already been
reported in Maryland and West Virginia.
Management. Preventing stink bugs from getting into buildings is the best line of
defense. All potential cracks and crevices around windows and doors, chimneys, siding, fas-
cia, and holes through which wiring passes should be caulked. Screens on windows and
doors and on roof vents and weep holes should be kept in good repair. Insecticides applied
around such openings in early fall when the bugs swarm and try to gain entry have short-
term effects. Most degrade rapidly. A professional exterminator should be consulted for
the best and safest results. With light infestations, individual bugs can be swept into a pail
of water.
Once the insects are indoors, vacuuming individual stink bugs is the best remedy. Empty
vacuum bags immediately to prevent a buildup of the stench injured, dying bugs exude. Do
not spray insecticides in an attempt to kill bugs hiding in the house. The dead bugs will
attract other pests such as carpet beetles, which will later feed on woolen items, dry goods,
and other natural materials.
 COMMON BED BUG n 123

Spot treatment of garden plants with insecticides will reduce the damage from these
insects.

Selected References
Day, Eric R., and Dini Miller. “Brown Marmotated Stink Bug, Homoptera: Penatomidae: Halyomorpha
halys.” Virginia Cooperative Extension, 2009. http://pubs.ext.vt.edu/2902/2902-1100/2902
-1100.html.
Gyeltshen, Jamba, Gary Bernon, and Amanda Hodges. “Common Name: Brown Marmorated Stink
Bug.” Featured Creatures, Department of Entomology and Nematology, University of Florida,
2010. http://entnemdept.ufl.edu/creatures/veg/bean/brown_marmorated_stink_bug.htm.
Jacobs, Steve. “Brown Marmorated Stink Bug,” Entomological Notes. College of Agricultural Sciences,
Cooperative Extension, Pennsylvania State University, 2010. http://ento.psu.edu/extension/fact
sheets/brown-marmorated-stink-bug.
MIPSP. “Brown Marmorated Stink Bug.” Massachusetts Introduced Pests Outreach Project, 2008. http://
www.massnrc.org/pests/pestFAQsheets/brownmarmoratedstinkbug.html.

n Common Bed Bug

Also known as: Bedbug, chinche, mahogany flat, redcoat, wall louse
Scientific name: Cimex lectularius
Order: Hemiptera
Family: Cimicidae
Native Range. Uncertain: possibly eastern Mediterranean. May have originated as a para-
site of bats that transferred to humans in caves occupied by nomadic Stone Age peoples. Bed
bugs apparently did not become a problem until people began to live in villages and cities.
Distribution in the United States. Throughout.
Description. Adult bed bugs are small, oval, wingless insects. Before they feed, bed bugs
are flat and brown, but when engorged after a blood meal, they are swollen and dark red.
Eyes are dark. The upper surface of the first segment of the thorax (the prothorax) forms a
wide collar-like plate that curves slightly around the broad head. Antennae are prominent
and have four segments. The abdomen has 11 segments. Before feeding, adult bed bugs
are about 0.25 in. (4–9 mm) long and 0.06–0.1 in. (1.5–3 mm) wide.
Newly hatched nymphs are the size of poppy seeds and colorless. Later nymphal instars
are smaller, paler versions of the adult. They become bright red after feeding. The cream-
colored oval egg is about 0.04 in. (1 mm) long.
Bed bugs are rarely seen. Evidence of their presence develops in areas where they congre-
gate and hide during the daytime. Such harborages will accumulate dark spots and staining
from their feces, eggs, and eggshells, shed skins of maturing nymphs, and adult bugs. Red
stains on bedding and mattresses occur when engorged bed bugs are accidentally crushed
by their sleeping human hosts. Bed bugs do exude a distinct odor, variously described as
resembling crushed coriander, rotting raspberries, or a sweet musty smell; but only when
infestations are heavy can this be detected by people.
Bed bug bites are indicated by a patch of redness on the skin with a darker raised welt in
the center. However, half of the people who are bitten experience no reaction; and among
those who do show visible signs, the size and itchiness of the bite varies greatly. Bites usually
occur on the exposed skin of the face, neck, arms, and hands and may be clustered or occur
in a line. Identification of the presence of bed bugs from the bites alone is problematic.
124 n INVERTEBRATES (INSECTS)

Related or Similar Species.

Most other bed bug species pre-
fer other mammals and birds as
hosts. C. lectularius prefers
humans, but will feed on bats,
cats, dogs, and rodents as well
as chickens and other birds. In
tropical regions of the world,
C. hemipterus is the more
common parasite of humans. It
has a noticeably narrower pro-
thorax than C. lectularius. (The
bed bug found in Hawai’i is
C. lectularius.) Bed bugs could
be confused with small ticks or
cockroaches. Their bites can be
mistaken for those of mosqui-
toes, fleas, and spiders.
Introduction History. The
dispersal history of bed bugs is
not well documented, but they
became widespread in the tem-
perate regions of the Northern
Hemisphere during the colonial
period and likely came to the
United States in the 1700s
when infestations on sailing
ships out of Europe were noto-
rious. They were known from
Greece as early as 400 BC but
were not mentioned in accounts
Top: The actual origins of the common bed bug remain unknown, but it from Germany until the elev-
may derive from the eastern Mediterranean. Bottom: Common bed bugs enth century or from France
are found throughout the United States. until the thirteenth century.
Although first noted in England
in 1583, they did not become common until 1670. Presumably, sometime after that
they made their way to what is now the United States. They had appeared in Jamaica,
another British colony, by 1726.
In the early twentieth century, bed bugs were so common that they were deemed one of
the top three household pests. The widespread use of DDT after World War II essentially
eradicated bed bugs from the United States and other developed countries in the 1950s.
However, bed bug populations quickly developed resistance to DDT, and the use of the pes-
ticide was banned altogether in the United States in 1972 because of its deleterious effects on
birds. In the 1990s, after an absence of nearly 50 years, bed bugs began a rapid increase, not
only across the United States but in all the industrialized countries of the world. Once asso-
ciated with poverty and underdevelopment, bed bugs now also occupy high-end hotels
and residences. The resurgence may be a consequence of increases in international travel
and immigration; the development of pesticide resistance, especially to pyrethroids; and
 COMMON BED BUG n 125

A. Adult bed bug. (James D. Young, USDA APHIS PPQ, Bugwood.org.) B. Bed bug eggs. (Mohammed El Damir,
Pest Management, Bugwood.org.)

changing control practices for other vermin, such as cockroaches and ants, that no longer
rely on broad-spectrum, persistent, and highly toxic pesticides, but focus on safer methods
of bait-trapping. One study has traced the origin of the latest outbreak of bed bugs to poultry
facilities in Arkansas, Delaware, and Texas. Workers could have unknowingly carried the
pests home or elsewhere when they left the workplace. Another study suggests that recent
infestations began in the Northeast and spread south and west from there. The last states
to report major infestations were Hawai’i, Oregon, and Washington.
Habitat. These bugs occur where people sleep. All they need are a source of blood and a
place to hide. They tend to hide close to where they feed. Their flat shape and small size
allow them to congregate in tiny cracks and crevices. They prefer dry, rough surfaces such
as wood and paper and avoid wet and hard surfaces such as stone, metal, and plaster.
Favored sites include the seams of mattresses, box springs, bed frames, and cracks in mold-
ing. In hotels, they are often first found behind headboards. They may be found among
items stored under beds; along and under the edges of wall-to-wall carpeting; behind pic-
ture frames, switch plates, and outlets; behind wallpaper; and in phones, smoke detectors,
and televisions. In large infestations, they will spread to upholstered furniture and among
clutter accumulated in closets.
While bed bugs were once associated in the popular mind with urban poverty and filth,
this is not the case. Today’s outbreaks commonly occur in fine hotels, vacation resorts,
and cruise ships as well as in hospitals, college dormitories, and well-maintained apartment
buildings and single-family residences.
Diet. Common bed bugs are obligate feeders on blood. Humans are their preferred hosts.
They feed mainly at night while their hosts are asleep. They respond to warmth and carbon
dioxide to locate their blood meal. Finding exposed skin, they pierce it with their mouth-
parts and inject their saliva, which contains an anticoagulant and numbing compound.
A full meal takes 5–10 minutes. Typically during that time, a bed bug makes a row of three
bites: “breakfast,” “lunch,” and “dinner.” Adults may live several months without feeding.
Life History. After mating, a female lays 2–3 eggs per day in cracks and crevices. The
female continues to lay throughout her lifetime, which may last 6–12 months, producing a
total of 200–500 eggs. At room temperature (68°F or 20°C), nymphs hatch out in 1–2 weeks
and immediately begin to feed. Bed bugs undergo incomplete metamorphosis in that the
126 n INVERTEBRATES (INSECTS)

nymphs are simply tiny adults. Each of the five juvenile or nymphal states requires a blood
meal in order to molt into the next stage. Full maturation takes from 9 to 18 weeks.
Several generations can be produced in a year.
Impacts. Bed bugs are not known to transmit diseases to humans. In minor infestations,
they are chiefly an annoyance and embarrassment. Their bites cause itchiness and inflamma-
tion in some people. Scratching the bites might result in secondary infections.
The occurrence of bed bugs can result in lawsuits against landlords and other property
owners obligated to provide safe and habitable conditions for tenants.
Management. Eradication, especially of major infestations in multiple-unit buildings, is
difficult and requires the assistance of experienced, professional pest control companies in
administrating an integrated pest management program. Although the media has recently
tried to blame EPA and its ban of persistent pesticides for the current resurgence in bed
bugs, many populations had already developed resistance to DDT and other organophos-
phates in the 1950s and 1960s as well as to the pyrethroids used in their place. Today,
few over-the-counter insecticides registered for use in homes are effective against bed bugs,
but licensed pest control operators do have some options available. Furthermore, some com-
panies are using specially trained dogs to help locate infestations.
Several nonchemical treatments exist that help control minor infestations. Careful and
repeated vacuuming of places where bed bugs like to hide can physically remove bugs from
sites that a high suction wand can reach. Heat (temperatures above 120°F or 49°C) kills
eggs, nymphs, and adults. Clothes and bedding can be washed in hot water. Backpacks,
toys, or shoes and the like—dry or wet—can treated in a clothes dryer set at medium-to-
high heat for 20 minutes. Larger items such as suitcases can be put in plastic bags and set
in the sun or in a closed car parked in the sun for a day. Freezing temperatures will also kill
bed bugs, but items must be left outside in winter or in a freezer for several days for the cold
to be effective.
Preventing an infestation is paramount in the fight against bed bugs. Bed bugs are trans-
ported from place to place on luggage, clothing, boxes, secondhand furniture and mat-
tresses. They spread through buildings by crawling out entry ways and through holes in
walls and ceilings. Caulking cracks and crevices and sealing or removing loose wallpaper
will help deter them, but careful inspection of items brought into the house is also impor-
tant. This is especially true for purchasers of secondhand beds and sofas and frequent trav-
elers. In hotels, keep suitcases off the floor and on hard surfaces such as luggage stands and
table tops. Put clothing in disposable plastic bags and directly into a washing machine and/
or dryer upon arrival home. Remove clutter from floors and monitor or eliminate any other
potential bed bug harborages. Keeping sheets and blankets from contacting the floor by
tucking them in and placing the legs of a bed frame in a small dish of mineral oil will keep
bed bugs from crawling up into a bed. Usually a combination of techniques is required, as
is the cooperation of all inhabitants of a building. Frequent inspection of one’s surroundings
will at the very least allow early intervention by a professional.
Bed bug bites may be avoided by wearing pajamas that cover arms and legs. Insect repel-
lents have not proven very effective deterrents, although mosquito netting impregnated with
permethrin may help—until bed bugs develop resistance to that pesticide.
Today’s rapid increase in bed bug populations is making them a major public health con-
cern and instigating more research on ways to control and eradicate this long-neglected
insect. New products such as the insecticide chlorofenpyr and the insect growth regulator
hydroprene, though slow acting, seem to be effective. Still, no chemical means of control
by itself can eliminate an infestation entirely.
 EMERALD ASH BORER n 127

Anyone with a suspected infestation should contact a licensed pest control operator for
confirmation and notify the public health department. Do not apply any insecticide or pesti-
cide directly to a mattress or other surface that comes into direct contact with people or pets.

Selected References
Anderson A., and K. Leffler. “Bed Bug Infestation in the News: A Picture of an Emerging Public Health
Problem in the United States.” Journal of Environmental Health 70(9): 24–27, 2008.
Brooks, Shawn E. “Bed Bug, Cimex lectularius Linneaus (Insecta: Hemiptera: Cimicidae).” Document
EENY-140 (IN297), Featured Creatures, Entomology and Nematology Department, Florida
Cooperative Extension Service, Institute of Food and Agricultural Sciences, University of Florida,
2009. https://edis.ifas.ufl.edu/in297.
Centers for Disease Control and Prevention, U.S. Environmental Protection Agency. “Joint Statement
on Bed Bug Control in the United States from the U.S. Centers for Disease Control and
Prevention (CDC) and the U.S. Environmental Protection Agency (EPA),” 2010. http://
www.cdc.gov/nceh/ehs/publications/Bed_Bugs_CDC-EPA_Statement.htm.
Pollack, Richard, and Gary Alpert. “Bed Bugs.” Harvard School of Public Health, 2005. http://
www.hsph.harvard.edu/bedbugs/.
Potter, Michael F. “Bed Bugs.” ENTFACT-636, Department of Entomology, University of Kentucky
College of Agriculture, 2010. http://www.ca.uky.edu/entomology/entfacts/ef636.asp.
Staab, Tina. “The History of Bed Bugs.” eHow.com, 2009. http://www.ehow.com/about_5376204
_history-bed-bugs.html#ixzz0y7tJ0BxF.

n Emerald Ash Borer

Also known as: EAB
Scientific name: Agrilus planipennis
Order: Coleoptera
Family: Buprestidae
Native Range. Emerald ash borers are native to northeastern China and adjacent areas of
Mongolia and Russia, Korea, Japan, and Taiwan, where forests are composed of broadleaf
trees closely related to those in the eastern United States.
Distribution in the United States. This beetle currently occurs in Illinois, Indiana,
Kentucky, Maryland, Michigan, Minnesota, Missouri, New York, Ohio, Pennsylvania,
Virginia, West Virginia, and Wisconsin.
Description. Adult emerald ash borers are small, metallic-green beetles. The slender,
elongate bodies are about 0.5 in. (7.5–13.5 mm) long, males being somewhat smaller than
females. When open, the wing covers expose a metallic-purplish-red abdomen. The segment
just behind the head, to which the first pair of legs is attached, is wider than the head. Larvae
are whitish with a brown head. Their abdomens have 10 segments, are flattened dorso-
ventrally, and end with a pair of brown pincers. Late stage larvae are about 1.0 in.
(26–32 mm) long.
Adults are active and seen in the open only from mid-May to September and usually only
in the afternoons of warm, sunny days, making infestations difficult to detect. Early signs and
symptoms that emerald ash borers have invaded a tree include jagged orange scars in the bark
made by woodpeckers on the upper trunk and branches and a top-down thinning of the can-
opy and yellowing of leaves. Bark may split vertically where larval feeding galleries have been
excavated beneath it. If the bark of infested ash trees is cut away, serpentine tunnels filled with
128 n INVERTEBRATES (INSECTS)

a fine sawdust-like frass are the

obvious signs that the beetles
are present. With heavy infesta-
tions, as trees begin to die, they
resprout from the roots.
Related or Similar Species.
A number of ash borers are
native to the United States but
are not similar in color, body
shape, or shape of the exit hole.
However, several metallic
wood-boring beetles native to
American forests are likely to
be confused with emerald ash
borers. Among them are the
bronze birch-borer (Agrilus anx-
ius) and the two-lined chestnut
borer (Agrilus bilineatus), both
of similar size and shape but
not green; and the six-spotted
tiger beetle (Cinindela sexgut-
tata) and the caterpillar hunter
(Calosoma scrutator), both of
which are green but have dis-
tinctly different body shapes.
Introduction History. The
emerald ash borer was first con-
firmed in the United States in
2002 in Canton, Michigan. It
probably had arrived at least
15 years earlier in wood pack-
Top: Emerald ash borers are native to the broadleaf forests of northeastern ing materials coming from
China, Korea, Japan, and Taiwan. (Adapted from “Native range of China. From Michigan, the bee-
emerald ash borer in Asia.” USDA Forest Service, Northern Research tle migrated to Ohio (2003) and
Station. http://www.nrs.fs.fed.us/disturbance/invasive_species/eab/local Minnesota (2009). In 2003, it
-resources/images/native_range.gif.) Bottom: The emerald ash borer was carried illegally to Prince
currently affects ash trees in cities, suburbs, and forests in 13 states.
George, Maryland, in nursery
(Adapted from map by Cooperative Ash Borer Project, USDA Forest
Service, 2010. http://www.emeraldashborer.info/files/MultiState stock originating in Michigan
_EABpos.pdf.) and sold in Maryland and
Fairfax County, Virginia. The
emerald ash borer continues to
be spread to other parts of infested states and to new states in live trees, green lumber, fire-
wood, wood chips, and debris. It had arrived in Indiana by 2004; Upper Peninsula,
Michigan, by 2005; Cook County, Illinois, by 2006; several counties in Pennsylvania by
2007, and at New River Gorge in Fayette County, West Virginia, by 2007. It was confirmed
in Missouri and Wisconsin in 2008 and in Kentucky, Minnesota, and New York in 2009.
Habitat. Emerald ash borers require ash trees. They survive in urban and suburban
parks, yards, and roadways, woodlots, and natural broadleaf deciduous forests.
 EMERALD ASH BORER n 129

A. Adult emerald ash borer. (Pennsylvania Department of Conservation and Natural Resources–Forestry Archive,
Bugwood.org.) B. Emerald ash borer larva. (Pennsylvania Department of Conservation and Natural Resources–
Forestry Archive, Bugwood.org.) C. Exit holes made by emerald ash borer. (Joseph O’Brien, USDA Forest
Service, Bugwood.org.) D. Twisting tunnels or galleries made by emerald ash borer larvae. (Art Wagner, USDA
APHIS PPQ, Bugwood.org.)

Diet. Emerald ash borers feed exclusively on trees of the genus Fraxinus. Adult beetles eat
along the edges of leaves. Larvae eat phloem and xylem in the outer sapwood. They tend to
prefer the upper trunks and branches of large trees. In the United States, green ash (F. penn-
sylvanica) and black ash (F. nigra) are more vulnerable to attack than white ash (F. ameri-
cana) and blue ash (F. quadrangulata), but all ash species are at risk.
Life History. Females deposit their eggs one by one in crevices in the bark or under flaps
of bark on the trunk and branches of ash trees. Seven to 10 days later, the eggs hatch, and
130 n INVERTEBRATES (INSECTS)

the larvae bore through the bark into the phloem to begin feeding. They feed for several
weeks, creating ever wider, s-shaped tunnels in the outer sapwood as they grow. A single
feeding gallery will range in length from 4 to 12 in. (20–50 cm). Feeding ends in the fall,
and the prepupal larvae overwinter in shallow chambers about 0.5 in. (1 cm) deep in the
outer sapwood or bark. The pupal stage begins in late April or May, and adults begin to
emerge in late May, with peak emergence occurring from early to mid-June. Adults are able
to fly immediately upon emerging and, although strong fliers, usually move less than 0.5 mi.
(0.8 km) to find a mate and a new host tree. They will feed on the foliage for a few days while
they mature, and then they mate. Females begin egg-laying after another week or two of
feeding. Adult males survive about one month after emergence, females for about two
months.
Research in Michigan indicates that the life cycle is longer in healthy trees than stressed
ones. In newly infested trees, many larvae overwinter in the earliest stage of development
and feed during a second summer. In stressed trees, nearly all larvae overwinter in the late
prepupal stage, and then pupate and emerge as adults the next summer.
Impacts. Emerald ash borers kill all trees that they infest. Adult beetles do relatively little
damage when they feed on the foliage. It is the larvae that are so destructive: they feed on
phloem and interrupt the flow of nutrients through the tree, so the tree weakens and dies.
The deaths of ash trees affect city and suburban landscapes, where they are planted as shade
trees and ornamentals in parks and yards. In forests, their deaths mean a loss of browse,
shelter, and seed that may be consumed by birds, small mammals, and insects. Although
many dead or dying trees produce sprouts at the base, these too are attacked once they attain
diameters of about 1 in. (0.5 cm). The demise of white ash across large swaths of eastern
North America could result in the loss of forest biodiversity and have widespread impacts
on habitat and watershed quality. In economic terms, trees valuable in the timber industry
are being lost, and livelihoods based upon the sale of wood products such as lumber, mulch,
and firewood are threatened. A special concern exists among some Native American groups
who view black ash as a cultural resource because they use it in their basket making.
Management. Containment rather than eradication is the main goal of management. The
federal government (APHIS) and infected states set up quarantine zones in efforts to prevent
the further spread of the emerald ash borer. Within each quarantine area, the movement of
firewood, green lumber, ash nursery stock, and ash debris as well as all wood chips is
restricted. As of March 2010, federal quarantine areas cover the entire states of Illinois,
Indiana, and Ohio; all of the Lower Peninsula of Michigan; and parts of Kentucky,
Maryland, Minnesota, New York, Pennsylvania, Virginia, West Virginia, and Wisconsin.
Interstate movement of all firewood is prohibited, since it is difficult to distinguish ash from
other hardwoods. Surveys of areas surrounding infected counties are conducted frequently
to detect new infestations. Monitoring tools such as prism purple panel traps, baited with
a lure and a nontoxic glue on the outer surfaces to catch any adult beetles attracted to the
trap, are used.
Experiments to see if a parasitic wasp can help control emerald ash borers are focusing on
three tiny stingless wasps that lay their eggs in beetle larvae or eggs. In their native range in
China, emerald ash borers are forest pests but do not have devastating effects, in part
because Chinese ashes have evolved resistance to their attacks. Asian trees survive in
Michigan, a state with a large infestation. It may be possible to hybridize American ashes
and Chinese species and select for resistance as has been done with the American chestnut
(Castanea dentata; see Fungi, Chestnut Blight Fungus) and American elm (Ulmus americana;
see Fungi, Dutch Elm Disease).
 FORMOSAN SUBTERRANEAN TERMITE n 131

Selected References
Bauer, Leah S., Therese M. Poland, and Deborah L. Miller. “Emerald Ash Borer.” Forest Disturbance
Processes. U.S. Forest Service, Northern Research Station, 2010. http://nrs.fs.fed.us/disturbance/
invasive_species/eab/biology_ecology/planipennis/.
“Emerald Ash Borer.” Pennsylvania Department of Conservation and Natural Resources, 2009. http://
www.dcnr.state.pa.us/forestry/fpm_invasives_EAB.aspx.
“Emerald Ash Borer.” Wikpedia, 2010. http://en.wikipedia.org/wiki/Emerald_ash_borer.
“Emerald Ash Borer Information.” Wisconsin Department of Agriculture, Trade and Consumer
Protection, n.d. http://datcp.wi.gov/Environment/Emerald_Ash_Borer/index.aspx.
McCullough, Deborah G., Noel F. Schneeberger, and Steven A. Katovich. “Emerald Ash Borer. Pest
Alert.” USDA Forest Service, Northeastern Area State and Private Forestry, 2008. http://
www.na.fs.fed.us/spfo/pubs/pest_al/eab/eab.pdf.

n Formosan Subterranean Termite

Also known as: FST
Scientific name: Coptotermes formosanus
Order: Isoptera
Family: Rhinotermitidae
Native Range. The Formosan subterranean termite is probably native to southern China.
It was first described from the island of Taiwan (Formosa) and hence its name. It may have
been transported to Japan before 1600.
Distribution in the United States. As of 2010, it was established in 10 states: Alabama,
California, Florida, Georgia, Hawai’i, Louisiana, Mississippi, North Carolina, South
Carolina, Tennessee, and Texas. Isolated colonies have been reported in Arizona,
Arkansas, New Mexico, and Virginia.
Description. Like other termites, the Formosan subterranean termite is a social insect that
lives in colonies composed of three castes: workers, soldiers, and the reproductives (alates or
swarmers, queen, king, and immature alates). The majority are white to off-white workers
that resemble most other termites. Identification of the species relies upon distinguishing
soldiers and alates. Soldiers have large teardrop-shaped heads that are orange-brown.
Their mandibles are black and sickle-shaped and, when crossed, form an X. Their
yellowish-white bodies are about 0.25 in. (6.4 mm) long. Soldiers are aggressive and, when
attacked, produce drops of a milky-white, glue-like fluid from a pore (fontanelle) on the
front of the head. Soldiers comprise 10–15 percent of the colony.
Alates are the winged reproductives; they swarm at night. Since they are attracted to light,
they are often found at windows or around light fixtures. They are yellowish brown and
have two pairs of wings covered with tiny hairs, visible under the low-power magnification
of a hand lens. The wings are clear and of equal length. Two thick veins occur on the leading
edge of each wing. Total length is about 0.5 in. (12–15 mm).
Since these termites spend much of their lives underground, one is more apt to see signs
of their presence rather than the insects themselves. A good indication that Formosan sub-
terranean termites have invaded is the occurrence of shelter tubes made of mud on hard sur-
faces such as on tree bark, up the side of foundations, and along concrete slabs. Nests made
of carton may become exposed on door frames, ceilings, stairs, and near the base of trees.
These nests may also be constructed in elevated locations without a connection to the
ground. Damage to windows, doors, and floors as well as utility poles, fences, and landscape
timbers are other signs that large termite colonies are nearby.
132 n INVERTEBRATES (INSECTS)

Related or Similar Species.

No other termites in the United
States build nests of carton, a
mixture of termite excrement,
chewed wood, saliva, and soil.
The feeding galleries of Form-
osan subterranean termites are
essentially free of soil, whereas
native subterranean termites fill
their galleries with soil and fecal
material. Infested timbers often
have layers of moist soil in areas
where termites are active. The
alates of native subterranean ter-
mites (Reticulitermes spp.) are
smaller, their wings have no
hairs, and they swarm in day-
light. Among native subterranean
termites, the proportion of
soldiers to workers is usually
only 1–2 percent, and the sol-
diers do not emit a white sub-
stance from the fontanelle.
Native soldiers’ heads are oblong,
and the mandibles do not cross.
Introduction History. The
Formosan subterranean termite
may have come to Hawai’i as
early as 1869, but it was not
officially reported there until
1913. Its first record in the
Top: The Formosan subterranean termite is believed to be native to continental United States is offi-
southern China. It was first described from the island of Formosa, now cially 1965, when it was discov-
Taiwan. Bottom: The Formosan subterranean termite was established in ered at a shipyard in Houston,
10 states by 2010 and reported in 4 others. (Adapted from “Just the Texas. It probably arrived in
Facts . . . Formosan Subterranean Termite,” 2006.) crated and palleted military
supplies on ships returning
from the Pacific after World War II. Indeed, the termite seems to have spread around the
world on ships; the earliest sites of infestations are at ports. In 1966, it was reported in
Louisiana (New Orleans and Lake Charles). The first official report of the termite in South
Carolina (Charleston) was made in 1967, but a search of earlier collections revealed its pres-
ence by 1957. First reports of the Formosan subterranean termite in Florida date to 1980–
1983; Alabama, 1985–1987; Tennessee (Memphis), 1985; North Carolina, 1990;
California (San Diego County), 1991; and Georgia (Atlanta), 1993. The termites are poor
flyers and do not spread rapidly on their own. After the initial introductions, the termites
were likely accidentally transported in contaminated building and plant materials brought
in from previously infested areas. They are commonly associated with used railroad ties, a
popular landscape timber. In general they have remained confined to the southeastern
 FORMOSAN SUBTERRANEAN TERMITE n 133

United States at latitudes south

of 32°300 N. However, the avail-
ability of centrally heated build-
ings may allow them to extend
their range farther north.
Habitat. Natural forests,
planted stands of trees, and
urban areas. Formosan subterra-
nean termites live underground
in moist areas with moderate
temperatures. Their nest will be
below frost level, but above the
water table. They also build aer-
ial nests on structures where
they can find moisture. Such
sites include boats, porches, flat
roofs where water pools, roof-
tops with vegetation, and gut- Formosan subterranean termite worker (top) and soldier (bottom).
ters. They will exploit live and (Gerald J. Lenhard, Louisiana State University, Bugwood.org.)
dead trees as nesting sites. Often
the nest is outdoors, and a man-made wooden structure is used only as food. The nest is gen-
erally accessed from the ground. Mud galleries or shelter tubes connect nests to food sources.
Diet. These termites consume cellulose from wood, cardboard, or paper. Bacteria and
other microorganisms in the gut digest the cellulose, releasing nutrients and energy for the
termite. They usually attack wood in contact with the ground and hollow out galleries by
feeding between the growth rings. Although they will chew through metal sheeting, asphalt,
some plastics, foam insulation, and plaster, they do not eat these materials. It is often
reported that they can hew through concrete, but they actually need preexisting cracks,
which they then enlarge. The termites feed over a foraging territory that may be several thou-
sand square feet in area.
Life History. Major swarms of Formosan subterranean termites begin in late spring and
continue through the summer. They occur on humid, still evenings, usually at dusk. A single
colony may release 70,000 or more winged reproductives. They fly only a short distance
(60–150 ft. or 20–50 m) and then shed their wings. The female searches for a nesting site
in moist crevices near a good supply of wood and the male follows her. Together, they hol-
low out a royal chamber. The pair, now king and queen, mate, and in a few days, the queen
produces 15–30 eggs. The eggs hatch in 2–4 weeks and are cared for by the king and queen
until they become third instars. These young termites will care for the next batch of larvae,
which will hatch from the second laying of the queen. The queen will continue to lay
2,000 eggs a day. It may take 3–5 years before the colony is large enough to produce alates
or to cause noticeable damage to trees and structures. Mature colonies will have as many as
10 million workers, soldiers, a primary queen, and several secondary reproductives.
Workers forage and take care of eggs and larvae and feed the larvae, soldiers, and reproduc-
tives. Soldiers defend the colony. The secondary reproductives take over egg production if
the queen or king dies.
Impacts. Termites feed on wood used in construction as well as on living trees. They hol-
low out the structure they are feeding upon, leaving a papery-thin covering behind. This
activity weakens and destroys beams, floors, sills, and the like and kills trees, including
134 n INVERTEBRATES (INSECTS)

those of aesthetic and commercial value. While individual Formosan subterranean termites
do not eat more wood than native species, the colonies are so much larger that more
damage results more quickly. Reportedly, in Hawai’i where a house was built on top of a
termite nest, the house was almost completely destroyed in two years. In Hawai’i, it is the
single most economically important insect pest in the state. Before Hurricane Katrina, the
Formosan subterranean termite cost New Orleans $300 million a year for control, repairs,
and replacement of utility poles due to this species. After hurricanes Rita and Katrina, the
highest concentrations of this pest in the continental United States were in the flooded cities
of New Orleans and Lake Charles, Louisiana. Cleanup of soggy debris and downed trees
only spread the termite, which was also attracted to brown rot fungi growing on wet wood
and wallboard. The IUCN has nominated this insect to be among “100 of the ‘World’s
Worst’ invaders.”
Management. Preventive measures include the use of pressure-treated wood wherever
timbers come in contact with the ground. It is also important to prevent the build-up of
moisture from leaky pipes, lawn irrigation sprinklers, clogged gutters, air conditioning con-
densate, and rainwater. Chemical barriers can be placed outside a structure both during and
after construction. Baits that use a chitin synthesis inhibitor are effective in eliminating an
entire colony. General groundskeeping is also a good preventive practice. Remove dead trees
and stumps and scrap wood piles from the property and reduce or do away with the use of
mulch near a building’s foundation. Once a population has become well established, it may
impossible to eradicate it.

Selected References
Carlson, Elizabeth. “The Formosan Subterranean Termite.” BugwoodWiki. Center for Invasive Species
and Ecosystem Health, University of Georgia, 2008. http://wiki.bugwood.org/The_Formosan
_Subterranean_Termite.
Hu, Xing Ping. “Formosan Subterranean Termites.” Alabama Cooperative Extension System, Alabama
A&M University and Auburn University, 2003. http://www.aces.edu/pubs/docs/A/ANR-1035/.
“Just the Facts . . . Formosan Subterranean Termites.” U.S. Army Center for Health Promotion and
Preventive Medicine, Entomological Sciences Program, 2006. http://phc.amedd.army.mil/PHC
%20Resource%20Library/FormosansubterraneantermitesJan2010.pdf.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Coptotermes formosanus (Insect).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?si=61&fr=1&sts=sss.
Su, Non-Yao, and Rudolf H. Scheffrahn. “Featured Creatures: Formosan Subterranean Termite.”
Entomology Department, University of Florida, 2000. http://www.entnemdept.ufl.edu/creatures/
urban/termites/formosan_termite.htm.

n Glassy-Winged Sharpshooter
Also known as: GWSS
Scientific name: Homalodisca vitripennis
Synonym: Homalodisca coagulata
Order: Hemiptera
Family: Cicadomorpha
Native Range. Glassy-winged sharpshooters are native to the southeastern United States,
where it occurs from eastern Texas to southern North Carolina.
 GLASSY-WINGED SHARPSHOOTER n 135

Distribution in the United

States. This leafhopper is a
native transplant to California.
Description. The glassy-
winged sharpshooter is a rela-
tively large leafhopper, measur-
ing from 0.4–0.5 in. (1.1–
1.4 cm) long. Females are
slightly larger than males. The
back is generally brown; small
ivory to yellow spots dot the
head and thorax. The underside
of the abdomen is white; face
and legs are yellow-orange.
Wings are transparent with red
veins. Some females have a
large white spot on the middle
of each wing composed of a
powdery material (brocho-
somes) that is secreted by the
insect. Nymphs look like adults
except that they are gray and
wingless.
Related or Similar Species.
The glassy-winged sharpshooter
can be told from almost all other
sharpshooters by its large size. It
is most similar to the native
smoke tree sharpshooter (Homa-
lodisca liturata), a close relative.
The smoke tree has wavy mark-
ings on its body instead of the Top: The glassy-winged sharpshooter is native to the southeastern United
spots characteristic of the glassy- States. (Adapted from Conklin and Mizell 2009.) Bottom: Glassy-winged
winged sharpshooter. sharpshooters infest several counties in southern California and threaten
Introduction History. Glassy- agricultural areas is the Central Valley and northern parts of the state.
winged sharpshooters were (Adapted from “Glassy-Winged Sharpshooter in California.” California
first collected near Irvine, Department of Food and Agriculture, 2010. http://www.cdfa.ca.gov/
pdcp/Maps/GWSS_Distribution2010.jpg.)
California, in 1989, although it
was mistakenly identified as the
native smoke tree sharpshooter at the time and not recognized as an introduced species until
1994, when it was properly identified in Ventura County. It had probably arrived as egg
masses on plants imported from the southeastern United States. It first became abundant in
commercial groves of citrus and avocado and on some woody ornamentals such as crape myr-
tle. Its presence was visible when “leafhopper rain” evaporated and whitened leaves. Toward
the middle of the 1990s, the glassy-winged sharpshooter moved inland in Riverside and San
Diego counties. By the end of the 1990s, large populations occurred in southern Kern County
in citrus groves and vineyards. It is expected that it will become a permanent resident in suit-
able habitats in the Central Valley and throughout northern California.
136 n INVERTEBRATES (INSECTS)

Habitat. In California, it pre-

fers riparian woodlands in
coastal and foothill areas. In its
native range, it occupies the for-
est edge. It also feeds and repro-
duces on woody ornamental
trees, vines, and annuals. Crape
myrtle and sumac seem to be
preferred in its native range; in
California, it inhabits eucalypts
and coast live oaks, and it is also
found on grapevines and citrus,
avocado, and macadamia trees.
Diet. Adult glassy-winged
sharpshooters suck the sap from
the xylem—the tissues that dis-
tribute water and dissolved
nutrients from the roots—of a
wide variety of host plants.
Feeding times are synchronized
with peak nutrient content in
the host plant. The insects insert
mouthparts that serve as straws
into the xylem. Nutrients in
xylem fluid are very diluted, so
large volumes of fluid must be
processed. The result is that
glassy-winged sharpshooters
produce large amounts of watery
wastes (“leafhopper rain”) that
Adult glassy-winged sharpshooter. (Russ Ottens, University of Georgia, can coat the canopies of infected
Bugwood.org.)
trees with a white residue as
the excreta evaporates. Adult
glassy-winged sharpshooters are able to feed on older wood than other sharpshooters because
of their larger mouthparts, and this also allows them to feed on dormant trees and vines during
the winter. They tend to feed closer to the base of grapevines and other plants than do other
sharpshooters. Young nymphs feed on the stems of the plant on which they hatch.
Life History. In California, mating occurs in spring and summer. Two generations are
produced each year. Initial egg-laying continues from late February through May. Each
female produces 10–12 eggs at a time and deposits them underneath the lower epidermis
of leaves of selected host plants. As she lays the eggs, she covers them with white scrapings
of brochosomes from her wings. Two weeks later, the eggs hatch into small nymphs that will
undergo five molting stages before they are mature. The first generation matures and, from
June through September, lays eggs for the second generation of the year. Adults live about
two months between generations. The adults from the second generation will produce
young the following year. Populations peak in summer and begin to decline in late
August. With the approach of winter, adults move into forested areas, where they go into
 GLASSY-WINGED SHARPSHOOTER n 137

semi-hibernation in the leaf litter until warmer temperatures return and they can begin to
mate.
Impacts. The glassy-winged sharpshooter, throughout its life cycle, is a vector for plant
diseases associated with strains of the bacterium Xylella fastidiosa. As such, it is a major threat
to California’s billion-dollar grape industry, where it is already implicated in an increase in
Pierce’s disease. Pierce’s disease clogs the xylem and destroys a plant’s ability to draw water
and nutrients from the soil to its leaves. At first, the tips of leaves turn brown and die, and in
1–2 years, the entire infected vine dies. Pierce’s disease itself is not new to California and has
probably been there for at least 100 years. The problem is that the glassy-winged sharp-
shooter is a more effective transmitter of the bacterium than native sharpshooters because
it is more of a generalist in terms of hosts, has greater mobility (up to 0.25 mi. per day),
and has larger mouthparts. While other species feed on the parts of the vine that get pruned
away, the glassy-winged sharpshooter feeds at the basal stems, and the infection can become
systemic or chronic, spread to other vines, and wipe out whole vineyards where table
grapes, wine, or raisins are produced.
Different strains of the X. fastidiosa bacterium cause diseases in other valuable plant spe-
cies, and the glassy-winged sharpshooter could become a vector for almond leaf scorch,
phony peach disease, alfalfa dwarf, and citrus variegated chlorisis, which affects orange
trees. It is already involved in the spread of oleander leaf scorch. Oleanders are widely
planted as ornamental shrubs in southern California, particularly in freeway medians, but
also in parks and suburban yards. The cost to the California Department of Transportation
should it have to remove diseased oleanders and replant medians with other ornamentals
would be well over $50 million. The wholesale nursery trade is also economically affected,
as ornamental plants require inspection and treatment with insecticides before transport to
other parts of California or neighboring states.
Management. Management is aimed at containing the invasion and preventing outbreaks
of Pierce’s disease and related plant diseases in new areas. Better early detection methods are
needed. Short-term strategies to slow the spread of the disease include the use of systemic
insecticides and behavior modifiers that would disrupt the transmission of the bacteria.
Biological control of the glassy-winged sharpshooter may be possible by using a small wasp
from Texas and northern Mexico, Gonatocerus truguttatus, that is an egg parasite of the
glassy-winged sharpshooter; or by inserting genes conferring resistance to X. fastidiosa into
host plants or rootstocks. Other bacteria or fungi may be found that are restricted to xylem
tissues and interfere with X. fastidiosa’s reproduction or transmission. One candidate is the
fungus Hirsutella sp. that affects glassy-winged sharpshooters in their native range.

Selected References
Cimino, Andria. “Glassy-Winged Sharpshooter, Xylophagous Leafhopper (Homalodisca coagulata).”
Introduced Species Summary Project, Columbia University, 2002. http://www.columbia.edu/itc/
cerc/danoff-burg/invasion_bio/inv_spp_summ/Homalodisca_coagulata.html.
Conklin, Tracy, and Russell F. Mizell III. “Featured Creatures: Glassy-Winged Sharpshooter.”
University of Florida, Institute of Food and Agricultural Sciences, 2009. http://entomology
.ifas.ufl.edu/creatures/fruit/glassywinged_sharpshooter.htm.
Garrison, Rosser W. “New Agricultural Pest for Southern California: Glassy-Winged Sharpshooter
(Homalodisca coagulata).” County of Los Angeles, Agricultural Commissioner/Weights and
Measures Department, 2001. http://acwm.co.la.ca.us/pdf/GWSSeng_pdf.pdf.
138 n INVERTEBRATES (INSECTS)

Hoddle, Mark S. “The Glassy-Winged Sharpshooter.” Center for Invasive Species Research, University
of California, Riverside, 2003. http://cisr.ucr.edu/glassy_winged_sharpshooter.html.
National Biological Information Infrastructure and IUCN/SSC Invasive Species Specialist Group.
“Homalodisca vitripennis (Insect).” ISSG Database, 2006. http://www.issg.org/database/species/
ecology.asp?si=240&fr=1&sts.
Pierce’s Disease Research and Emergency Response Task Force. “Glassy-Winged Sharpshooter
(Homalodisca coagulata).” University of California, Agriculture and Natural Resources, n.d. http://
news.ucanr.org/speeches/glassywinged.html.

n Gypsy Moth
Also known as: European gypsy moth
Scientific name: Lymantria dispar
Order: Lepidoptera
Family: Lymantridae
Native Range. The gypsy moth comes from the temperate broadleaf forests of Eurasia and
northern Africa. European and Asian subspecies have evolved. The gypsy moth that is now
invasive across the eastern United States originated in Europe. The first egg masses were
brought from France.
Distribution in the United States. Gypsy moths are established in many northeastern
states and continue to expand their distribution area westward and southward. Currently
they occur from northern North Carolina to Maine and west through Ohio and northern
Indiana into Wisconsin.
Description. Gypsy moths are sexually dimorphic. The female is white with dark chev-
rons across the forewings. She has a wing span of about 2 in. (80 mm) but is flightless.
Her body is covered with hairs, and her antennae are slender. The male is darker and
smaller. His wings are dark brown with black banding, and he has feathery antennae.
A strong flier, the male has a wingspan of about 1.5 in. (60 mm). The abdomen of the male
is narrower than that of the female.
Eggs are deposited in oval masses that are covered with soft, buff-colored hairs from the
female’s abdomen. The velvety masses are about 1.5 in. (60 mm) long and 0.75 in. (30 mm)
wide.
Newly hatched larvae are about 0.125 in. (3 mm) long and initially tan, but become black
within four hours. Younger caterpillars (first to fourth instars) are brown to black and have
long body hairs. Late instars are black with 11 pairs of bumps (tubercles) along the back.
The forward five pairs are blue; the rear six pairs are red. Each tubercle has a tuft of yellow
or brown hairs. A single yellow line runs the length of the back of younger caterpillars
from the head to the last segment. Additional yellow lines adorn the head of fourth
through sixth instars. True legs are dark red. At maximum growth, the caterpillars are 2–3 in.
(80–120 mm) long.
Pupae are dark red-brown and teardrop-shaped, rounded in the front, and tapered
toward the rear. Each pupa attaches to a substrate by means of a few strands of silk. Male
pupae are 0.75–1.5 in. (30–60 mm) long; female pupae are up to 2.5 in. (100 mm) long.
Related or Similar Species. The Asian gypsy moth is a subspecies of Lymantria dispar that
is occasionally intercepted at western ports. The adult moths look very similar to the
European variant that has invaded the northeastern United States, but the female is a strong
flier. Asian caterpillars vary more in color than European ones. It is safe to assume, for now,
that any flying white moth is not a gypsy moth.
 GYPSY MOTH n 139

Gypsy moth caterpillars

might be mistaken for some
native tent caterpillars; however,
gypsy moths never make tents
or webs. The eastern tent
caterpillar (Malacosoma ameri-
canum) has a white line down
its back and light blue and black
spots on the sides. The forest
tent caterpillar (Malacosoma diss-
tria) has a line of white blotches
the length of its back, and light
blue stripes on the sides. The fall
web worm (Hyphantria cunea) is
greenish or yellow with long
white hairs and has a black stripe
down the back and a yellow
stripe on each side.
Introduction History. The
introduction of the gypsy moth
can be traced to a single person
and a specific address. A
Frenchman and amateur ento-
mologist, Etienne Leopold Tro-
uvelet, had come to Medford,
Massachusetts, with his family
in 1852. He was interested in
identifying American silkworms
for possible use in producing
silk. For unknown reasons, in
1868 or 1869, after a visit to
France, he brought some gypsy Top: The gypsy moth’s native range coincides with the temperate
moth egg masses back to his broadleaf forests of Eurasia and North Africa. (Adapted from “Gypsy
house at 27 Myrtle Street. It Moth around the World.” USDA Forest Service. http://www.fs.fed.us/ne/
seems he let the eggs develop morgantown/4557/gmoth/world/.) Bottom: The gypsy moth continues
on trees in the backyard. When to spread southward and westward from established populations in the
Northeast and Great Lakes region. (Adapted from “Gypsy Moth
some caterpillars escaped, he
Quarantine Map.” USDA Forest Service, 2007. http://www.fs.fed.us/ne/
notified local authorities, but morgantown/4557/gmoth/atlas/q2007.gif.)
no effort was made to eradicate
them. In 1882, a gypsy moth
outbreak on Myrtle Street marked the first evidence of an emerging problem. The gypsy
moth population continued to grow, and in 1889, the first program to eradicate the moth
began. The Massachusetts State Board of Agriculture implemented manual removal of egg
masses from trees and structures, applied pesticides, and burned infested forests, all to no
avail. Eradication efforts ceased in 1900. The historical rate of spread between 1900 and
1915 is estimated to have averaged about 6 mi./year (9.5 km/year). By 1934, gypsy moths
had spread north through Vermont, New Hampshire, and Maine and south into
Connecticut and Rhode Island. Expansion rates had slowed to less than 2 mi./year
140 n INVERTEBRATES (INSECTS)

A. Gypsy moth caterpillar. (John H. Ghent, USDA Forest Service, Bugwood.org). B. Adult moths, male (left) and
female (right). (USDA Forest Service Archive, USDA Forest Service, Bugwood.org.)

(2.8 km/year) between 1916 and 1965, so that by 1965, only eastern New York State, including
Long Island, was added to the distribution area. After 1965, the rate of spread again increased,
as did the range of the gypsy moth. At an estimated spread rate of nearly 13 mi./year (21 km/
year), the gypsy moth moved south through Pennsylvania and New Jersey into Maryland,
West Virginia, and Virginia and west across northern Ohio, into Michigan, and eventually
Wisconsin and northeastern Minnesota. Nothing suggests that this spread has stopped.
Newly hatched (first instar) gypsy moth caterpillars sometimes move down from the tree
canopy on silken threads that break in strong winds, carrying the larvae out of the canopy.
Updrafts can carry them as much as 12 mi. (nearly 20 km) from the tree where they hatched.
However, most dispersal to new areas is probably achieved by unintentional human trans-
port. Gypsy moth egg cases and larvae can hitchhike on firewood, nursery stock, vehicles,
and outdoor equipment. The typical pattern of spread is one of isolated populations appear-
ing ahead of the expanding boundary of the distribution area and then coalescing to form a
continuous, new area of infestation.
Habitat. Temperate broadleaf deciduous forests are preferred habitat. Populations
become densest and resulting defoliation most intense on dry ridges and other areas with
shallow soils or excessively drained soils. Coniferous forests, urban and suburban land-
scapes, and agricultural environments also have habitat value for gypsy moths.
Diet. Gypsy moth caterpillars feed on foliage. Adult moths do not have mouthparts and
thus do not eat at all. Host plants are extremely varied; over 500 different types can be con-
sumed. Suitability of plant species for forage depends on the developmental stage of larvae.
In general, trees under stress are more vulnerable to attack than healthy specimens. In the
northeastern United States, oaks are the most favored hosts, but maples, hickories, and
many other hardwoods are also eaten. Trees that are avoided include American holly, ash
(Fraxinus spp.), black walnut (Juglans nigra), butternut (Juglans cinerea), flowering dogwood
(Cornus florida), mountain laurel (Kalmia latifolia), balsam fir (Abies balsamea), and arborvi-
tae (Thuja occidentalis).
Life History. Moths emerge from pupae in summer, males usually 1–2 days ahead of
females. A few hours after emergence, females release a pheromone that attracts males,
 GYPSY MOTH n 141

which fly as much as 0.5 mi. (0.8 km) upwind to mate with them. Females deposit eggs
within 24 hours of mating. Egg masses are placed on tree trunks and the undersides of
branches as well as in crevices in the bark and under loose bark. They also may be laid on
camping gear and in vehicles or anywhere else to which females can crawl. The eggs are
covered with hairs from the female’s abdomen that probably provide some protection from
predators and cold temperatures. Although development from embryo to larva is completed
within a month, larvae enter diapause and overwinter inside the egg. Eggs hatch the follow-
ing spring at about the time leaf buds open on oak trees.
Most larvae in a given egg mass hatch within seven days of each other. In cool or rainy
weather, they stay near the remains of the egg mass, but once the sun shines, they move
upward toward the light into the canopy of the tree. Larvae feed first on new leaves and,
when not feeding, stay attached by silk threads on the undersides of leaves. The caterpillars
undergo molts about once a week to accommodate their growth. Males have four molts;
females five. The larval period lasts about 40 days, after which time each caterpillar finds a
pupation site, wraps itself in a thin silk net, and rests for 1–2 days before becoming a pupa.
The pupa erupts from the larval skin and becomes dark reddish-brown; it stays in the silk
net for the next two weeks while development into an adult moth is completed. When the
adult moth emerges, it takes a few hours to expand its wings. The males then fly away in
search of females. Adult moths live about a week.
Gypsy moth populations are cyclical. A low-level, innocuous phase may last several years,
to be followed by a release phase of 1–2 years’ duration that sees an increase in population
density of several orders of magnitude. The next phase is the outbreak phase, when popula-
tion density is great enough that defoliation is evident over a considerable area of forest.
Outbreaks rarely last more than one or two years before starvation and disease cause the
population to crash in what is known as the decline phase. The foliage of infested trees may
respond by increasing levels of toxic chemicals and decreasing the nutritive value of leaves,
contributing to the die-off. Naturally occurring pathogens such as the fungus Em
(Entomophaga maimaiga)—originally introduced from Asia when Massachusetts was
attempting to stamp out gypsy moths in the early 1900s—and NPV (nucleopolyhedrosis
virus) commonly contribute to population declines. Population cycles are synchronous over
large distances, but they proceed aperiodically.
Impacts. Gypsy moths, especially during outbreaks, defoliate hardwood and sometimes
softwood trees. This can weaken the tree, making it more susceptible to drought, diseases,
and other pests such as the shoestring fungus (Armillaria mellea) and the two-lined chestnut
borer (Agrilus bilineatus). Defoliated forests open the forest floor to sunlight, causing acceler-
ated drying of leaf litter and increased forest-fire risk. Most trees can recover from a single
defoliation event, but several defoliations in successive years can kill them. In the Northeast,
an estimated 20 percent of forest trees die after heavy infestations. How this will affect species
composition and ecological dominance is as yet unknown, but in time, ecologically dominant
oaks could be replaced by less palatable tree species, which would have ripple effects
throughout the forest community. The loss of trees and the unsightly mess of hundreds of cat-
erpillars and a constant rain of their feces from the treetops during an outbreak can negatively
affect not only timber production, but also tourism and recreation. Other economic costs are
incurred in inspection and other control measures. Automobile and train wrecks on roads
and tracks slippery with caterpillars have been reported during some severe outbreaks.
Management. Gypsy moth control is aimed at suppressing existing populations, eradica-
ting pioneer populations, and slowing the spread of this invasive moth. Homeowners can
control small local infestations by manually scraping and destroying egg masses from trees
142 n INVERTEBRATES (INSECTS)

in their yards or attaching sticky barrier bands to tree trunks to prevent the migration of
caterpillars. More widespread suppression of populations is achieved by spraying areas with
Bt (Bacillus thuringiensis), a bacterium that produces a chemical toxic to moth and butterfly
larvae.
The National Gypsy Moth Slow the Spread (STS) program, a joint local, state, and federal
initiative, was implemented in 1999 to monitor recently established and still-isolated low-
level populations in zones between infested areas and non-infested areas. Along the front
line of range expansion, gypsy moth population monitoring is conducted with pheromone
traps to capture males, and several types of barriers are placed on tree trunks to collect lar-
vae. Direct counts of egg masses are also employed. Once new infestations are identified,
Integrated Pest Management (IPM) and other controls are initiated to eradicate or suppress
the population. APHIS has established quarantine areas in much of the infested region.
Nursery stock, firewood, vehicles, outdoor equipment, and household items moving out
of these areas into non-infested ones must be inspected by certified inspectors.

Selected References
Baczynski, Tracy. “European Gypsy Moth (Lymantria dispar).” Introduced Species Summary Project,
Columbia University, 2002. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/
inv_spp_summ/Lymantria_dispar.htm.
“Gypsy Moth.” Animal and Plant Health Inspection Service, U.S. Department of Agriculture, 2010.
http://www.aphis.usda.gov/plant_health/plant_pest_info/gypsy_moth/index.shtml.
“Gypsy Moth Pest Profile.” California Department of Food and Agriculture, State of California, 2010.
http://www.cdfa.ca.gov/phpps/pdep/target_pest_disease_profiles/gypsy_moth_profile.html.
“Integrated Pest Management Manual: Gypsy Moth.” National Park Service, U.S. Department of the
Interior, 2010. http://www.nature.nps.gov/biology/ipm/manual/gypsymth.cfm.
Liebold, Sandy. “Gypsy Moth in North America.” Forestry Sciences Laboratory, Forest Service,
Northeastern Research Station, U.S. Department of Agriculture, 2003. http://www.fs.fed.us/ne/
morgantown/4557/gmoth/.

n Hemlock Woolly Adelgid

Also known as: Woolly
Scientific name: Adelges tsugae
Order: Homoptera
Family: Adelgidae
Native Range. Hemlock woolly adelgids are native to Asia. They are known to be from
Japan, Taiwan, southwestern China, and India. Genetic analysis has determined that those
in the eastern United States came from southern Japan. In their native range, these adelgids
utilize both hemlocks and spruces and undergo both asexual and sexual reproduction. Asian
hemlocks appear to have developed resistance to adelgid attacks.
Distribution in the United States. The hemlock woolly adelgid is invasive in 16 eastern
states from southeastern Maine to northern Georgia and west to eastern Kentucky,
Tennessee, and West Virginia. Its distribution area continues to expand; and it may soon
become established in Ohio, where isolated collections have occurred. In the western United
States, where it is not a pest, it occurs from northern California to southeastern Alaska.
Description. The hemlock woolly adelgid is a tiny, aphid-like insect less than 1/16 in.
(1.5 mm) long. They are oval and blackish gray in color. Newly hatched nymphs (first
 HEMLOCK WOOLLY ADELGID n 143

instars) are about the same size,

but reddish brown. The insect
probably would not be noticed
were it not for the fact that
nymphs covers themselves in
white wax-like filaments and
adults secrete a white “woolly”
ovisac in which to lay their
eggs. The woolly or cottony cas-
ings are about 1/8 in. (3 mm) in
diameter and quite visible at the
base of needles on the under-
sides of the outermost tips of
hemlock branches from late fall
to early summer.
Introduction History. The
hemlock woolly adelgid was
first identified on the West
Coast in 1924. First reports on
the East Coast came from
Richmond, Virginia, in 1951. It
may have been accidentally
imported on nursery stock from
Japan. Today, it continues to
spread on ornamental hemlocks
via the nursery trade, but early-
stage larvae and eggs can be
transported by birds and mam-
mals moving through hemlock
forests. The wind can disperse
infested twigs as well as the
mobile crawlers. Also, humans Top: The hemlock woolly adelgid is native to coniferous forests from
can unintentionally transport India to Japan. (Adapted from Reardon, R., et al. “Biological Control of
eggs, nymphs, and adults on Hemlock Woolly Adelgid.” U.S. Department of Agriculture, Forest
live ornamental trees and with Service. Forest Health Technology Enterprise Team. Burlington, VT.
debris from dead and dying FHTET-2004-04. http://wiki.bugwood.org/Archive:HWA/Introduction.)
Bottom: The hemlock woolly adelgid is currently invasive in 16 eastern
hemlocks. The hemlock woolly
states, and it continues to expand its range. In the west it is not
adelgid appeared in Shenan- considered a pest. (Adapted from “Counties with Established HWA
doah National Park in Virginia Populations 2009.” USDA Forest Service. Http://na/fs/fed/us/fnp/hwa/
in the 1980s, moved southward maps/2009.pdf.)
on the Blue Ridge Parkway in
the 1990s, and was in Great Smoky Mountains National Park by 2002. In 2007, it was
spreading at a rate of nearly 10 mi./year (15.6 km/year) in the southern part of its range
and 5.0 mi./year (8.13 km/year) in the northern part. By some estimates, it could kill all
the hemlocks in the southern Appalachians by 2020.
Habitat. In the United States, these aphid relatives are restricted to hemlocks, although in
their native range, certain spruces are hosts during the sexually reproducing stages of the life
cycle. In the eastern United States, this adelgid infests both eastern hemlock (Tsuga
144 n INVERTEBRATES (INSECTS)

A. Woolly casings on a hemlock twig infested with hemlock woolly adelgids. (Connecticut Agricultural
Experiment Station Archive, Connecticut Agricultural Experiment Station, Bugwood.org). B. The aphid-like
adult adelgid. (Michael Montgomery, USDA Forest Service, Bugwood.org.)

canadensis) and Carolina hemlock (T. caroliniana). In the West, the woolly hemlock adelgid
occurs on both western hemlock (T. heterophylla) and mountain hemlock (T. mertensia),
but these trees are resistant, and the adelgid is not considered a pest.
Diet. Nymphs and adults suck sap from the twigs of hemlocks. They feed at the base of
new needles by extracting starch-rich fluids from the tissues in the xylem that manufacture
and store the plant’s food. At the same time, they inject toxins in their saliva into the plant.
Life History. Hemlock woolly adelgids have both sexual and asexual (parthenogenic)
stages in the life cycle, but U.S. populations only multiply asexually; all are females. Six
stages of development define the life cycle: egg, four instars of nymphs, and adult. Two gen-
erations are produced each year. In March, females that had overwintered lay 20–75 eggs
and cover them in cottony material excreted by the adult. Larvae, known as crawlers, emerge
in April and May and move to feeding places at the bases of hemlock needles. They may
hitchhike to nearby hemlocks on birds and mammals or be blown there by the wind.
When the crawlers settle, they become immobile nymphs. By June, they have matured into
wingless and winged adults. The winged females fly off looking for spruce trees, but since
no suitable host spruces exist in the United States, most of them presumably die without
reproducing. If there were such spruces, these females would reproduce sexually. The wing-
less females reproduce asexually and lay 100–300 eggs that will become the second genera-
tion of the year. The larvae hatch in early July, settle at the bases of hemlock needles, and
then go into dormancy through the hot summer months. Nymphs resume development in
October and November and feed throughout the winter, maturing into adult females early
the next spring.
Impacts. Hemlock woolly adelgids weaken and kill eastern and Carolina hemlocks,
including those in forests and those cultivars used as ornamental trees and hedges. To date,
the most severe impacts have occurred in parts of Virginia, New Jersey, Pennsylvania, and
Connecticut. In the Northeast, hemlock decline and death takes 4–10 years; in the southern
Appalachians, the pace is more rapid, and trees may die in only 3–6 years. Infestations often
start in large, mature hemlocks, which may be 150 ft. (45 m) tall and more than 500 years
old. Infested trees can be identified first by their grayish-green needles that contrast with
 HEMLOCK WOOLLY ADELGID n 145

the glossy green of healthy hemlocks. Defoliation follows, progressing from the lower
branches upward and ultimately causing the death of the tree. The rate and extent of hem-
lock decline are accelerated by other stress factors, such as drought, poor soil conditions,
and other pests and diseases. Hemlocks form important habitat for numerous birds from
warblers to turkeys, and for small mammals such as snowshoe hares and rabbits; they pro-
duce a deep shade along mountain streams that maintains the cool water temperatures
needed by trout and other fish. The loss of the hemlock canopy increases light penetration
to the forest floor, drying and warming riparian and riverine habitats in the summer and
reducing shelter in upland habitats in winter. Research suggests that nitrogen cycles are
altered when hemlock stands perish. The demise of hemlocks could have cascading effects
that would alter forest ecosystems in much of the eastern United States.
Management. Few environmentally sound methods for controlling hemlock woolly
adelgids in forests are available. Natural predators and pathogens exist, but fail to lower pop-
ulation levels enough to prevent tree deaths. Biological control by Asian beetles, such as the
ladybird beetle (Pseudoscymnus tsugae), that limit their attacks to adelgids and control them
in Japan, hold some promise; several states have released them experimentally. DNA
research suggests that Asian hemlocks have evolved a degree of resistance to adelgid infesta-
tions. Crossing eastern hemlocks with Asian hemlocks has successfully maintained the
appearance of the eastern hemlock but produced a tree that discourages adelgid settlement
and slows the growth of those nymphs that do feed upon it.
Isolated trees in yards or parks can be treated with insecticidal soaps, or adelgids can be
physically removed with strong sprays of water; but these treatments must be repeated often.
Removal of infected branches will slow the decline of the tree. In Great Smoky Mountains
National Park, treatments with systemic insecticides that are applied at the roots or injected
into the trunk of trees in campgrounds have reversed some of the impacts of the infestation
and remained effective for several years.
Preventing or slowing the spread of the adelgid can be aided by cultural control methods
such as not moving live plants, logs, firewood, or bark chips from infested areas. Selective
removal of heavily infested trees reduces that likelihood that wind, birds, and other wildlife
will disperse eggs and nymphs. When replanting landscape trees, the use of adelgid-resistant
hemlock species, such as T. diversifolia and T. sieboldii from Japan or western and mountain
hemlocks from the Pacific Northwest, instead of eastern hemlock will limit the availability of
suitable host trees and may help control adelgid populations.

Selected References
Chowdhury, Shahrina. “Hemlock Woolly Adelgid (Adelges tsugae).” Introduced Species Summary
Project, Columbia University, 2002. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/
inv_spp_summ/Adelges_tsugae.html.
“Hemlock Woolly Adelgid.” Pest Alert, State and Private Forestry NA-PR-09-05. U.S. Department of
Agriculture, Forest Service, Northeastern Area, 2005. http://na.fs.fed.us/spfo/pubs/pest_al/
hemlock/hwa05.htm.
McClure, Mark S. “Hemlock Woolly Adelgid, Adelges tsugae (Annand).” Connecticut Agricultural
Experiment Station, 1998. http://www.ct.gov/caes/cwp/view.asp?a=2815&q=376706.
Pennsylvania Department of Conservation and Natural Resources. “Hemlock Woolly Adelgid.”
Commonwealth of Pennsylvania, 2009. http://www.dcnr.state.pa.us/forestry/woollyadelgid/
index.aspx.
146 n INVERTEBRATES (INSECTS)

n Japanese Beetle
Scientific name: Popillia japonica
Order: Coleoptera
Family: Scarabaeidae
Native Range. Japan.
Distribution in the United States. The Japanese beetle is established in all states east of
the Mississippi River except Florida. Noncontinuous infestations occur west of the
Mississippi in Arkansas, Colorado, Iowa, Kansas, Minnesota, Missouri, Nebraska, and
Oklahoma. Far-western states are protected by quarantine, and any beetles arriving have,
so far, been eradicated.
Description. The Japanese
beetle is an oval-shaped, metal-
lic-green insect about 0.5 in.
(10–13 mm) long and 0.25 in.
(6–7 mm) wide. The wing cov-
ers are bronze-colored and
extend almost the full length of
the abdomen. Five small white
tufts of hair line each side of the
body and two others lie just pos-
terior to the wing covers. Males
are usually slightly smaller than
females. Japanese beetles travel
and feed in groups.
Eggs are spherical and al-
most translucent. They swell to
a diameter of about 0.08 in.
(2 mm) before hatching. The
grub or larvae is white with a
reddish-brown head. It has
three pairs of legs and lies
curled in the form of a C. Full-
grown larvae are about 1.0 in.
(26 mm) long. The pupa looks
like a cream-colored or tan
adult with the legs folded close
to the body.
Related or Similar Species.
Japanese beetle larvae look like
those of most beetles and can
be distinguished by a V-shaped
row of spines beneath the
Top: The Japanese beetle is native to Japan. Bottom: In the United States,
abdomen.
the Japanese beetle occurs in all states east of the Mississippi River except Introduction History. The
Florida. Isolated populations occur as far west as Colorado. (Adapted Japanese beetle was first discov-
from APHIS, 2004.) ered in a nursery in Riverton,
 JAPANESE BEETLE n 147

A. Adult Japanese beetle. (David Cappaert, Michigan State University, Bugwood.org.) B. Japanese beetle larvae
(grubs) are commonly found in the soil beneath lawns. (David Cappaert, Michigan State University,
Bugwood.org.)

New Jersey, in 1916. It is believed that grubs contaminated shipments of iris bulbs sometime
before 1912, when inspections of imports began in the United States. In the next 60 years, it
spread throughout 22 states east of the Mississippi River. The climate is favorable to the
insect, large expanses of turf in lawns and golf courses provide excellent habitat for repro-
duction, and abundant foliage is available to support adults. The beetle can be spread during
any stage of its life cycle in plant materials, soil, and sod. It has now moved into states west
of the Mississippi, but for the most part, populations have remained isolated. Hitchhikers on
aircraft are continually intercepted at airports in western states, where it could become a
major pest in orchards and truck farms.
Habitat. Open woods, meadows, farms, gardens, and lawns.
Diet. Adult Japanese beetles eat the leaves, flowers, and fruits of hundreds of different
plants. Their hosts include trees, shrubs, vines, and perennial and annual herbs, including
crops and ornamentals. Preference is for young leaves, which they skeletonize by consuming
the green tissue between the veins. Grubs feed primarily on the roots of grasses, but will also
consume the roots of annual fruit and vegetable plants.
Life History. Adults begin to emerge from pupae in early summer and congregate
on plants to feed and mate. Each female leaves the plant upon which she is feeding
again and again to deposit 1–4 eggs at a time. In the course of the summer, she will lay
40–60 eggs total. She deposits her eggs in soil at depths of 3–4 in. (1.2–1.6 cm), and prefer-
ably beneath turf. The larvae hatch out about two weeks later and begin to feed on plant
roots and grow. With the approach of cold weather, the grubs move deeper into the soil to
overwinter. When soil temperatures warm in the spring, the grubs migrate back up toward
the surface and resume feeding. They pupate and, in 8–20 days emerge as the next genera-
tion of adults. The life cycle usually takes a year. Ten months of the cycle are spent in the
larval stage.
Impacts. Leaves of heavily infested ornamental, truck, and garden plants will turn brown
and die. Soft fruits such as grapes and berries may be eaten completely. Grapes injured by
beetles become vulnerable to attack by native green June beetles (Cotinus mitida), which
are unable to bite into intact grapes themselves. Corn is damaged when Japanese beetles
148 n INVERTEBRATES (INSECTS)

eat the silk and prevent the formation of kernels. The larvae damage turf, turning patches of
lawns brown when numbers are high.
Management. Biological controls can be effective in controlling grubs and hence adult
beetles. Applications of milky spore (the bacterium Bacillus popillae) to turf can reduce pop-
ulations for decades if properly used. Nematodes (Heterorhabditis spp.) also work well in
destroying grubs. Commercially available traps lure Japanese beetles away from plants, but
may also attract beetles into a yard. Landscape plantings of non-palatable plants such as for-
sythia, holly, juniper, arborvitae, boxwood, spruce, and yew may deter beetles from massing
in suburban properties.

Selected References
APHIS. “Managing the Japanese Beetle: A Homeowner’s Handbook.” Animal and Plant Health
Inspection Service (APHIS), U.S. Department of Agriculture, 2004. http://www.aphis.usda.gov/
lpa/pubs/pub_phjbeetle04.pdf.
Bilberry, S. “Popillia japonica.” Animal Diversity Web, 2001. http://animaldiversity.ummz.umich.edu/
site/accounts/information/Popillia_japonica.html.
Day, Eric, Pete Schultz, Doug Pfeiffer, and Rod Youngman. “Japanese Beetle.” Virginia Cooperative
Extension, Virginia Tech, and Virginia State University, 2009. http://pubs.ext.vt.edu/2902/2902
-1101/2902-1101.html.
“Japanese Beetle Pest Profile.” California Department of Food and Agriculture, 2009. http://
www.cdfa.ca.gov/phpps/pdep/target_pest_disease_profiles/japanese_beetle_profile.html.
“Japanese Beetles.” Forest Insect and Disease Newsletter. Minnesota Department of Natural Resources,
2002. http://www.dnr.state.mn.us/fid/november02/japanese.html.

n Multicolored Asian Lady Beetle

Also known as: Japanese ladybug, Asian lady beetle, Halloween lady beetle
Scientific name: Harmonia axyridis
Order: Coleoptera
Family: Coccinellidae
Native Range. Asia. The native range stretches from the Altai Mountains east to the
Pacific Ocean and Japan and from southern Siberia south to southern China. The distribu-
tion area includes Kazakhstan, Uzbekistan, Russia, Mongolia, China, Korea, Taiwan, and
Japan. This lady beetle has at various times been imported to the United States from Japan,
Korea, and Russia.
Distribution in the United States. Well-established populations exist in many parts of the
Midwest, Northeast, South, and Northwest.
Description. Like most other lady beetles, the adult multicolored Asian lady beetle has a
domed, oval shape. Populations in the United States contain individuals in a mix of colors,
from tan or pale yellow-orange to bright red-orange. They may or may not have black spots
on the wing covers. Those that do have up to 10 spots on each wing cover. The middle body
segment (pronotum) is white to straw-yellow and usually marked with a distinct black M.
Adults are relatively large, measuring about 0.3 in. (7–8 mm) long.
Larvae are elongate and flattened; their bodies are covered with flexible spines. Late
(fourth) instars are bluish black with a yellow-orange patch on each side of the abdomen.
Eggs are bright yellow.
 MULTICOLORED ASIAN LADY BEETLE n 149

Related or Similar Species.

Many native lady beetles occur
in the United States. The most
common is the convergent lady
beetle (Hippodamia convergens).
Adults are somewhat elongate
in shape compared to the multi-
colored Asian lady beetle.
Convergent lady beetles have
black spots on red wing covers.
Behind the head, white lines
converge on a black back-
ground. They range in size from
0.16 to 0.28 in. (4–7 mm).
Native lady beetles do not over-
winter indoors.
Another introduced lady
beetle, the seven-spotted lady
beetle (Coccinella septempunc-
tata), from Europe, is estab-
lished in some northeastern
and north central states. It is
about the same size as the
multicolored Asian lady beetle
but has a white spot on either
side of its black head. The wing
covers are red or orange with 1–
5 spots on each one. It overwin-
ters in sheltered areas outdoors
near the fields in which they
feed.
Introduction History. Multi- Top: The multicolored lady beetle occurs across a broad swath of Asia
colored Asian lady beetles have from Kazakhstan to Japan. Bottom: This alien lady beetle is invasive in
been deliberately introduced to most of the lower 48 states. (Adapted from “Multicoloured Asian Lady
the United States numerous Beetle.” Project UFO. http://www.projectufo.ca/drupal/multicoloured
times as agents for the bio- _asian_lady_beetle.)
logical control of aphids and
aphid-like insects. They were first released by the U.S. Department of Agriculture in
California in 1916 and again in the mid-1960s to control pecan aphids. From 1978 to
1982, additional releases were made in Connecticut, Delaware, Georgia, Louisiana, Maine,
Maryland, Mississippi, Ohio, Pennsylvania, and Washington. Interestingly, none of these
efforts seem to have given rise to established populations.
In 1988, a population was discovered north of New Orleans in Louisiana. This may have
derived from an accidental introduction. From this point of origin, multicolored lady beetles
spread quickly through southern and midwestern states. By 1994, they were reported in
Alabama, Florida, Georgia, North Carolina, South Carolina, Ohio, and Minnesota. They
were also established in the Northeast by that time, perhaps as a result of other, independent
150 n INVERTEBRATES (INSECTS)

A. Adult multicolored Asian lady beetle. (Gerald J. Lenhard, Louisiana State University, Bugwood.org.) B. Larva.
(Gerald J. Lenhard, Louisiana State University, Bugwood.org.) C. Pupa. (John Ruberson, University of Georgia,
Bugwood.org)

introductions rather than dispersal from the South or Midwest. It is known, for example,
that they arrived at ports in South Carolina and Delaware as accidental hitchhikers on
imported nursery stock.
Habitat. Multicolored Asian lady beetles live in cultivated fields, orchards, vineyards,
nurseries, and gardens. They usually overwinter in buildings.
Diet. Aphids and similar soft-bodied insects such as scales and psyllids comprise the
main food of both adult and larval multicolored Asian lady beetles. An adult can devour
90–270 aphids a day, and a single larva may consume 600–1,200 while it develops. These
lady beetles also feed on the larvae of butterflies and of other beetles as well as on injured
fruits such as grapes.
Life History. Mating takes place in early spring after adults emerge from overwintering
sites. Females lay clumps of roughly 20 eggs on the undersides of leaves. The eggs hatch
in 3–5 days, and the fast-moving larvae forage on the host plant for aphids and scale insects.
The larvae will molt four times as they grow larger. After the fourth molt, they enter an
 MULTICOLORED ASIAN LADY BEETLE n 151

immobile pupa stage. The adult beetle emerges from the pupa 5 or 6 days later. The com-
plete life cycle from egg to adult takes 15–25 days. Before the first killing frost of autumn,
as prey become less abundant, adults seek shelter indoors. They seem to be attracted by
bright reflective surfaces such as the south-facing sides of light-colored buildings. They enter
homes and other buildings in huge numbers though cracks and other poorly sealed open-
ings and occupy cool places inside walls, floors, and attics. There they become dormant.
Winter aggregations may number in the tens of thousands. On warm, sunny days, they wake
up and move to the interior of the building, where they seek the light at windows. With the
warming temperatures of springtime, they exit their winter shelters and begin to mate. They
do no physical damage to structures and do not reproduce indoors. They may return to the
same overwintering sites every year. Adults can live 2–3 years.
Impacts. The introduction of the multicolored Asian lady beetle as a biological control
agent on shrubs, trees, and a variety of crops has been largely successful. Its consumption
of large numbers of aphids, scales, and thrips has reduced the need for chemical pesticides.
In California, it has nearly eradicated pecan aphids, and elsewhere significantly reduced soy-
bean aphids after their recent introduction from China.
Multicolored Asian lady beetles are a minor, but perhaps increasing, agricultural pest.
They appear to displace native lady beetles and other beneficial insects through competition
for food and predation. This lady beetle may also depress numbers of the exotic seven-
spotted lady beetle (Coccinella septempunctata). In autumn, the beetles congregate and feed
on ripening fruits such as pears, apples, and grapes—especially if the fruits have already
been damaged by birds or other insects. This is particularly troublesome in vineyards, where
it is difficult to remove them from the grapes being harvested for winemaking. Lady beetles
get crushed along with the grapes and taint the flavor of the wine.
The multicolored Asian lady beetle may be a threat to eggs and caterpillars of the mon-
arch butterfly (Danaus plexippus). Its real infamy, however, derives from its status as an
annoying overwintering invader of homes. Numbers can be so great that homeowners can
hear them moving in the walls. If a lady beetle is frightened or crushed, it exudes an unpleas-
ant odor and secretes, via reflex bleeding, a yellowish liquid from its leg joints that will stain
drapes, carpets, and other light-colored surfaces. They are more a nuisance than a threat,
however. They do not destroy wood or textiles or otherwise damage human property.
They do not carry diseases. They do not sting, although they can bite. Sensitive people
may have allergic reactions to their presence.
Management. Preventing multicolored Asian beetles from entering homes is the best way
to deal with them. All possible entries should sealed; door and window screens, as well as
those in all vents, should be kept in a state of repair; and door sweeps should be installed
on exterior doors. Once inside, it is difficult to remove them. The most recommended
method is by vacuuming them and employing a means to discard the catch before survivors
escape, and before the smell of dead beetles becomes too strong. Swatting them or sweeping
them up can lead to reflex bleeding and bad odors. Do not use bug bombs. They may have
little effect and will only attract other scavenger insects.

Selected References
“Biological Control: A Guide to Natural Enemies in North America: Harmonia axyridis.” Cornell
University, College of Agriculture and Life Sciences, Department of Entomology, n.d. http://
www.nysaes.cornell.edu/ent/biocontrol/predators/Harmonia.html.
152 n INVERTEBRATES (INSECTS)

Hahn, Jeffrey. “Multicolored Asian Lady Beetles.” University of Minnesota Extension Service, 2004.
http://www.extension.umn.edu/distribution/housingandclothing/m1176.html.
Jones, Susan C., and Joe Boggs. “Multicolored Asian Lady Beetle.” Ohio State University Extension Fact
Sheet, HSE-1030-01.Ohio State University, n.d. http://ohioline.osu.edu/hse-fact/1030.html.
Koch, R. L. “The Multicolored Asian Lady Beetle, Harmonia axyridis: A Review of Its Biology, Uses in
Biological Control, and Non-Target Impacts.” Journal of Insect Science 3: 32, 2003. Published online,
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC524671/.

n Red Imported Fire Ant

Also known as: RIFA
Scientific name: Solenopsis invicta
Order: Hymenoptera
Family: Formicidae
Native Range. South America. Red imported fire ants come from a narrow strip extend-
ing from Porto Velho, Brazil, northwestward to Santa Fe, Argentina. Genetic studies suggest
northeastern Argentina as the most likely source area of fire ants in the United States.
Distribution in the United States. Imported red fire ants can be found across the
southeastern United States. They occur from southeastern Virginia and eastern and southern
North Carolina across the Gulf states and eastern Texas. Isolated populations occur in parts
of Maryland, Tennessee, Arkansas, Oklahoma, New Mexico, and California. They are likely
to spread through southwestern Texas to southern Arizona and from southern California
north along the Pacific coastal region into northern California and Oregon. The ants also
occur in Puerto Rico and the U.S. Virgin Islands.
Description. Red imported fire ant colonies produce three types of workers, all sterile
females and distinguished from other ants by their very aggressive behavior. The smallest,
the so-called “minor workers,” are about 0.1 in. (1.6 mm) long and the largest, comprising
about 35 percent of a mature colony, the “major workers,” are about 0.25 in. (6 mm) long.
Intermediate in size are the many “media workers.” These ants are reddish to dark brown.
The waist (pedicel) has two segments, and all workers possess a stinger at the end of their
black bulbous abdomens (gaster). Winged females and males, known as “reproductives,”
are both about 0.34 in. (8.8 mm) long. Each colony will have one or more queens, also about
0.34 in. (8.5 mm) long.
Red imported fire ant mounds occur in open areas and are rarely more than 18 in.
(45 cm) in diameter or 16 in. (40 cm) high. They are hard and have no visible entrances.
Those built in clay soils are typically symmetrical domes, while those constructed in sandy
soils are irregularly shaped. However, in some circumstances, no mounds are evident. In
urban and suburban areas, they may nest under concrete slabs, in the walls of buildings,
or in electrical equipment.
Related or Similar Species. Among the many fire ant species that live in the United
States, three other members of the genus Solenopsis are also pests. Separating one from
another is difficult and requires a sample size of 40 or more workers for proper identification
even by experts. The black imported fire ant (S. richteri) is limited in its distribution, found
only in a small part of northern Mississippi and Alabama. It is displaced by and may actually
be a subspecies or regional variant of the red imported fire ant. The southern fire ant (S.
xyloni) is native to the Southeast. Nests are often built under stones or boards or at the bases
of plants. The nest usually appears as loose soil with many craters spaced over an area of 2–4
sq. ft (0.18–0.37 m2). The tropical fire ant (S. geminata) is another native species of the
 RED IMPORTED FIRE ANT n 153

southeastern United States.

Tropical fire ant workers have
square heads that are large in
proportion to their bodies.
Their mounds are often built
around clumps of vegetation.
Introduction History. Im-
ported red fire ants were intro-
duced to the United States
through the port of Mobile,
Alabama, in the mid-1930s.
Genetic studies trace the origins
to 9–20 queens and (presum-
ably) their workers. A secondary
introduction may have occurred
60 miles to the west of Mobile.
From these pioneers, popula-
tions spread to other south-
eastern states. They arrived in
Virginia in 1989, and were dis-
covered in Los Angeles, Orange
and Riverside counties, Cali-
fornia, in 1998. In single-queen
colonies, growth is outward at a
rate no more than 40 m per year.
In multi-queen colonies, disper-
sal of winged reproductives cov-
ers greater distances. Most new
colonies form within a mile of
the reproductives’ birthplace,
but some may be 10 miles or
more distant. Nonetheless, most Top: The red imported fire ant is native to South America. Bottom: The
of the range expansion in the red imported fire is invasive in the Gulf States and east Texas and
United States is probably the expected to expand its range in coming years. It has also been
result of human actions. Ants introduced into Puerto Rico. (Adapted from USDA Agricultural Research
are spread when mated females Service map, 2007. http://www.ars.usda.gov/fireant/Imported.htm.)
are transported in sod, hay, the
root balls of ornamental plants, and on earth-moving equipment.
Habitat. Red imported fire ants are most often associated with disturbed habitats such as
agricultural fields, pastures, lawns, and other open, sunny sites. Rarely do they occur in natural
forests or similar well-shaded locations. These ants are limited by cold temperatures and cli-
mates receiving less than 20 in. (510 mm) of precipitation a year. In dry regions, however, they
can exist by colonizing areas near permanent sources of water or areas regularly irrigated. Red
imported fire ants can regulate their microclimate to a degree by moving their broods to higher
and lower levels of their mounds, which may extend as much as 4 ft. (1.2 m) below the ground.
Diet. Red imported fire ants are omnivores. They are voracious predators of other arthro-
pods, but also consume emerging seedlings of field and truck crops such as soybeans, egg-
plant, cucumbers, corn, okra, and a host of others. They are known to chew the bark of
154 n INVERTEBRATES (INSECTS)

citrus trees and eat fruits. In

homes, they prefer oily and
greasy foods high in protein.
These ants also tend aphids and
consume honeydew. Worker
ants can only swallow liquids.
Solids are cut to a manageable
size and carried back to the nest
to be fed to larvae. The larvae
digest the food and regurgitate a
protein-rich brew that feeds the
workers and queen. The workers
also regurgitate what they have
swallowed so that their nest
mates can lick or suck it up.
Life History. Biological pat-
terns for fire ants must be
The waist of the red imported fire ant has two distinct segments. All
described in terms of both the
workers have a stinger at the end of the abdomen. (Eli Sarnat.)
individual and the colony, since
the latter is the essential social
unit. The individual’s life cycle begins as an egg. A grub-like legless larva hatches in
8–10 days and begins the major growth stage of the individual. It periodically sheds its
skin as it grows larger, and becomes a pupa with legs in 6–12 days. The pupal phase lasts
another 9–16 days, after which time adult sterile workers emerge and begin foraging or
reproductives emerge, take flight, and mate some 300–800 ft. (90–240 m) up in the air.
The complete transition from egg to adult takes 22–38 days.
A colony begins life when a mated female descends to the earth, breaks off her wings, and
digs a founding nest 2–5 in. (5–12 cm) straight down into the ground. She seals herself off and
lays 12–14 worker eggs. It takes about a month for these eggs to develop into adults. During
this period, the queen does not feed, but she nourishes the larvae with sterile eggs, regurgi-
tated oils, and salivary secretions. These workers will be the smallest in the colony cycle and
are called “minims.” They open the nest, forage for food, rear a new group of workers, and
feed the queen. The queen becomes essentially an egg-making machine and can produce
200 eggs a day. As the colony grows, more foragers (10–20% of total workers) mean more
food, and a larger proportion of larvae become majors. At the end of its second year, a colony
matures and produces winged reproductives. The colony at that time averages 27,000 work-
ers. Very large mature mounds may house 240,000 workers. Minor workers live 1–2 months,
media workers 2–3 months, and major workers 3–6 months. Queens live 2–6 years.
Impacts. Red imported fire ants are mainly feared for their bites. Their venom creates a
burning pain in the victim and, within 24 hours, raises itching pustules on the skin.
These aggressive ants swarm over anything disturbing their nests, so bites are multiple.
For sensitive people, bites can result in anaphylactic shock and, rarely, death.
Increasingly, red imported fire ants are invading homes, outdoor electric meters and air
conditioners, traffic control boxes, and airport runway lights, increasing opportunities for
human-ant interaction, interfering with switching mechanisms, and causing short circuits
and shutdowns. Infestations of red imported fire ants also pepper lawns and parks with
unsightly mounds.
 RED IMPORTED FIRE ANT n 155

Fire Ants Leave the United States for Distant Shores

S ince 2000, red imported fire ants have invaded China, Taiwan, and Australia.
The southern United States appears to be the source of these invasions. Recent
genetic studies indicate two or more introductions to each area have occurred.
The ants that arrived in Taiwan stem directly from populations in California; how-
ever, California received its fire ants from southern U.S. states. American popula-
tions carry distinct combinations of genes (haplotypes) that are extremely rare in
South America but common in the United States, Asia, and Australia and allow
the invasion history of the ants to be tracked. Hitchhikers in cargo, fire ants appear
to be following the world’s major trade routes to new locales.
Source: Ascunce, Marina S., Chin-Cheng Yang, Jane Oakey, Luis Calcaterra, Wen-Jer Wu,
Cheng-Jen Shih, Jérôme Goudet, Kenneth Ross, and DeWayne Shoemaker. “Global Invasion
History of the Fire Ant Solenopsis invicta.” Science 331: 1066–68, 2011.

In agricultural fields, mounds can damage farm equipment. Ants feed on young shoots,
protect aphids that feed on plants, and can interfere with root growth; many crops suffer sig-
nificant damage where ant colonies occur. On the other hand, red imported fire ants are
major predators of pests such as boll weevils, sugarcane borer, and ticks. It is credited with
reducing the range of the lone star tick, which targets livestock.
In more natural settings, the red imported fire ant is a strong competitor of native ants,
especially the tropical fire ant; and displaces the invasive Argentine ant (Linepithema humile).
It has been implicated in the decline of the bobwhite (Colinus virginianus) in the southeastern
United States and has negatively impacted other ground-nesting birds and reptiles through
bites, predation, and competition for space and food.
Management. Poisonous baits have proven most effective in reducing or eliminating col-
onies, since red imported fire ants carry food back to their nests. Boiling water poured on
individual mounds may help reduce populations, but area flooding only causes the ants to
link together in rafts and disperse. Biological controls hold promise, particularly the use of
the nematode Neoaplectana carpocapsae and parasitic phorid flies that lay their eggs in ants.
Spread of the ant is being slowed with the establishment of quarantine areas.
Ant-proofing structures can prevent the entry of ants to buildings. Caulking and sealing
all cracks and crevices keep ants out, since they usually nest outdoors and enter homes only
to feed. Generally sanitizing outdoor areas through the frequent emptying of trash cans and
dumpsters also discourages foraging ants.

Selected References
Apperson, Charles, and Michael Waldvogel. “Red Imported Fire Ant in North Carolina.” Insect
Note_ENT/rsc-35, Department of Entomology, North Carolina Cooperative Extension, n.d. http://
www.ces.ncsu.edu/depts/ent/notes/Urban/ifa.htm.
Collins, Laura, and Rudolf H. Scheffrahn. “Red Imported Fire Ant.” Featured Creatures, Department of
Entomology and Nematology, University of Florida, Institute of Food and Agricultural Sciences,
2008. http://entnemdept.ufl.edu/creatures/urban/ants/red_imported_fire_ant.htm.
156 n INVERTEBRATES (INSECTS)

“Integrated Pest Management Manual: Fire Ants.” National Park Service, U.S. Department of the
Interior, 2010. http://www.nature.nps.gov/biology/ipm/manual/fireants.cfm.
IUCN/SSC Invasive Species Specialist Group. “Solenopsis invicta (Insect).” ISSG Database, 2006. http://
www.issg.org/database/species/distribution.asp?si=77&fr=1&sts=&lang=EN.
“Red Imported Fire Ant Pest Profile.” California Department of Food and Agriculture, 2010. http://
www.cdfa.ca.gov/phpps/pdep/target_pest_disease_profiles/rifa_profile.html.
n Vertebrates
n Fish
n Alewife
Also known as: Big eye herring, freshwater herring, gray herring, kyack, sawbelly, white
herring, branch herring, river herring, glut herring, mulhaden, golden shad
Scientific name: Alosa pseudoharengus
Synonym: Pomolobus pseudoharengus
Family: Clupeidae
Native Range. Native to the Atlantic Ocean from Red Bay, Labrador, to South Carolina;
spawning in estuaries and Atlantic Slope rivers of the eastern United States as much as
100 miles inland.
Distribution in the United States. As a native transplant, it is established in all five Great
Lakes and also reported in streams, landlocked lakes, and reservoirs in Colorado (head-
waters of the Colorado River basin), Georgia (Savannah River), Kentucky (Ohio River),
Indiana (Bass Lake), Maine (Belgrade Lakes), Nebraska (Merritt Reservoir, Ainsworth
Canal), New York (Lake Otsego, Cayuga, Upper Saranac, Big Moose, Woodhull, Saratoga,
and Seneca lakes; St. Regis headwaters, mountain lakes in Adirondacks), Pennsylvania
(Delaware Gap National Recreation Area), Tennessee (Dale Hollow Reservoir, Watauga
Reservoir), Vermont (Lake St. Catherine), Virginia (Claytor Lake, John W. Flanagan
Reservoir, Lake Chesdin, Leesville Reservoir, Smith Mountain Lake), West Virginia
(Bluestone Reservoir, New and Kanawha rivers), Wisconsin (Kangaroo Lake, Pigeon River,
Pigeon Lake, East Twin River, Sheyboygan River, Green Bay, St. Louis River estuary, Sauk
Creek, and Milwaukee River).
Description. The transplanted, landlocked alewife is smaller than the native anadromous
alewife and typically reaches a length of 6–10 in. (15–25 cm) and weight of less than 4 oz.
(0.11 kg). The body is laterally compressed and relatively deep. It has a silvery color with
a darker greenish sheen on the back. A distinctive black dot occurs on the body behind
the eye. Scales merge along the belly to form scutes that create a serrated edge, the reason
for one of the fish’s common names, sawbelly. The tail or caudal fin is forked. Alewives move
in large schools.
Introduction History. Alewives were first reported from Lake Ontario in 1873. Some have
suggested they are actually native to the lake, having migrated up the St. Lawrence River
from the Atlantic Ocean at some earlier time in geologic history. Another possible route for
entry to the Great Lakes was the Erie Canal, opened in 1825 to connect Lake Erie with the
Hudson River and the Atlantic Ocean. The alewife was first recorded from Lake Erie in
1831. The fish slowly dispersed upstream, perhaps aided by the Welland Canal, which con-
nects Lakes Ontario and Erie. Populations were established in Lake Huron by 1933, Lake
Erie by 1940, Lake Michigan by 1949, and Lake Superior by 1954. Genetic information sug-
gests that alewives arrived via the Erie Canal and are not descendants of a population native
158 n VERTEBRATES (FISH)

to Lake Ontario. The fish is now

abundant in Lake Huron and
dominant in Lake Michigan. Its
numbers are held in check in
Lake Superior by cold water
and in Lake Erie by water too
shallow to provide many over-
wintering refuges.
Invasion of the Great Lakes
was aided by prior overfishing
of its major predators, Atlantic
salmon and lake trout. The
introduction of the sea lamprey
(see Fish, Sea Lamprey) may
also have reduced the popula-
tion sizes of competitors.
Equally important, alewife phy-
siology was such that they
could thrive in the landlocked
freshwater habitats to which
they were introduced without
the need to return to the sea
after spawning, part of the natu-
ral life cycle of populations in
their native range.
In water bodies other than
the Great Lakes, all introduc-
tions were intentional, through
both legal (e.g., in Virginia and
West Virginia) and illegal (e.g.,
Lake Otsego and Adirondack
Top: The alewife is native to the western Atlantic Ocean, from Labrador lakes in New York and Lake St.
south to South Carolina. In its native range it is anadromous. Bottom: Catherine in Vermont) stocking
As a native transplant, the alewife is established in all of the Great Lakes of streams and lakes with forage
and in streams, lakes, and reservoirs in several states. (Both maps fish.
adapted from Fuller, Maynard, and Raikow 2009.) Habitat. Temperate lakes,
reservoirs, and rivers. They pre-
fer the deeper waters of lakes by day and move into shallower waters near shore at night.
Alewives spawn in shallow waters and tributary streams in the spring. They overwinter in
the deepest parts of lakes, but extremely cold winters may cause die-offs.
Diet. Primarily a filter-feeder, the alewife consumes zooplankters such as copepods.
However, they are generalists and also feed on insect larvae, fish eggs, and small fish fry.
Life History. Alewives spawn in the spring during the night. Each female produces
12,000–50,000 eggs, each about 0.9 mm in diameter; they are broadcast as the males release
sperm. Eggs are deposited over all types of substrate. No parental care is invested in eggs or
larvae; the adults leave the spawning grounds as soon as spawning is completed. The larvae
hatch after 3–6 days, depending upon water temperature; and 3–5 days after that they begin
to feed. Larvae transform into juvenile fish slowly. Landlocked fishes mature in 1–3 years,
 ALEWIFE n 159

faster than native anadromous

members of the species. They
live fewer than 10 years. Mass
die-offs occur periodically, usu-
ally in the spring.
Alewives are prey for many
predators, including native lake
trout (Salvelinus namaycush),
eels (Anguillidae), bigmouth
bass (Micropterus salmoides),
and whitefish (Coregonus spp.).
Herons and other fish-eating
birds take alewives, as do semi-
aquatic mammals such as otter
and mink.
Impacts. Alewives trans-
formed the Great Lakes ecosys-
tem. In some parts of the lakes, Transplanted alewives inhabit landlocked lakes, reservoirs, and streams.
(Eric Engbretson.)
they are now a keystone spe-
cies. In Lake Michigan, 70–90 percent of the fish, by weight, are alewives. They outcom-
peted native zooplankton-feeders and are blamed for the demise of the lake trout in Lake
Michigan, once a mainstay of the lake’s commercial fishery. Their preying on fish fry is
implicated in declines of emerald shiner (Notropis atherinoides), yellow perch (Perca flaves-
cens), deepwater sculpin (Myoxocephalus thompsoni) and burbot (Lota lota). In smaller bodies
of water, they compete with yellow perch, rainbow smelt (Osmerus mordax), and young bass,
all of which survive on zooplankton. Alewives may also interfere with reproduction in land-
locked Atlantic salmon (Salmo salar) and lake trout in areas where alewives are the main prey
species: Alewives have a high amount of the enzyme thiaminase, so that the eggs of their
predators become deficient in thiamine, resulting in high mortality rates among the fry.
Alewives are so well established in the Great Lakes ecosystem that their removal at this
point would be disruptive. Instead, they are seen as a boon to efforts to reestablish an impor-
tant native game fish in the lakes, the Atlantic salmon, for which they serve as an important
forage species. Pacific Chinook salmon (Oncorhynchus tshawytscha) and coho salmon
(Oncorhynchus kisutch) have been introduced to Lake Michigan to help control alewives
and to provide a sport fishery.
Mass die-offs, such as occurred during the 1960s in the Great Lakes, polluted the beaches
with rotting fish and were both nuisances and health hazards.
In other waters, where alewives are more recent invaders, they have the potential to alter
the food web and reduce biodiversity.
Management. Population reduction is the only option available in the Great Lakes, where
the alewife is a permanent addition to the fish fauna, and eradication might actually cause
more damage to the ecosystem. Reestablishment of the Atlantic salmon and the stocking of
other predatory fish such as Pacific salmon and brown trout (Salmo trutta: see Fish, Brown
Trout), itself an exotic species, may help control alewife numbers. Elsewhere, culling extant
populations and the prevention of new introductions are essential to reduce threats to native
ecosystems. Legislation and enforcement of laws that make transport of alewives illegal are
important. Construction of net barriers to prevent downstream dispersal out of lakes has
been suggested. In some instances, fishing of alewives and collecting them when they
160 n VERTEBRATES (FISH)

congregate on spawning or overwintering grounds have been recommended. Alewives are

sometimes used for bait and for pet food.

Selected References
Bean, Tim. “Alewife (Alosa pseudoharengus).” Introduced Species Summary Project, Columbia
University, 2002. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/
alewife.html.
Fuller, Pam, Erynn Maynard, and David Raikow. “Alosa pseudoharengus.” USGS Nonindigenous
Aquatic Species Database, Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/FactSheet.asp
?speciesID=490.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Alosa pseudoharengus (Fish).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=625&fr=1&sts=&lang=EN.
Tobias, V., and W. Fink. “Alosa pseudoharengus.” Animal Diversity Web, University of Michigan
Museum of Zoology, 2004. http://animaldiversity.ummz.umich.edu/site/accounts/information/
Alosa_pseudoharengus.html.

n Asian Swamp Eel

Also known as: Rice eel, rice-paddy eel, belut, white ricefield eel, yellow eel.
Scientific name: Monopterus albus
Synonym: Fluta alba
Family: Synbranchidae
Native Range. Indian subcontinent, Southeast Asia, East Asia. It is also native to Central
and South America and may be native to Australia.
Distribution in the United States. Florida (North Miami area, Little Manatee River and
Bullfrog Creek drainages near Tampa, and a canal system near Homestead, South Miami–
Dade County); Georgia (Chatahoochee Nature Center, Roswell, Fulton County;
Chattahoochee National Recreation Area, Gwinnett County); Hawai’i (O’ahu), and New
Jersey (vicinity Silver Lake, Gibbsboro, Camden County).
Description. Asian swamp eels have elongated bodies and a compressed, tapering tail.
They lack scales and fins and are covered with mucous. A single V-shaped gill opens on
the underside just behind the head. The nose is blunt, and the eyes are small and dark.
The upper lip is thick and covers part of the lower lip. Teeth are bristle-like (villiform).
Body color ranges from olive green to brown; some are spotted with flecks of gold, yellow,
or black. Total length is about 3 ft. (1 m); adults weigh about 1 lb. (0.5 kg).
Related or Similar Species. Asian swamp eels are not true eels, but they may be mistaken
for native American eels (Anguilla rostrata). True eels have scales and fins and are anadro-
mous. Asian swamp eels also resemble sea lampreys (see Fish, Sea Lamprey), which have
obvious dorsal and caudal fins and seven gill openings on each side of the head. Lampreys
lack jaws and have oval-shaped mouths. In Florida, Asian swamp eels might be confused
with two large native aquatic amphibians, the two-toed amphiuma (Amphiuma means) and
the greater siren (Siren lacertina). The former have four tiny legs and the latter have small
front legs and bushy external gills.
Introduction History. Asian swamp eels were in Hawai’i by 1900. They were likely deliber-
ately introduced for food. Swamp eels are a common food fish in China and other parts of
Asia, where they are sold live. It is likely that immigrants brought them to Hawai’i. The first
 ASIAN SWAMP EEL n 161

reports of swamp eels occurring

in Florida and Georgia stem
from the 1990s; these may have
been accidentally released from
fish farms or intentionally
“freed” by aquaria owners. In
Florida, two populations (North
Miami and Tampa) can be genet-
ically traced to China, and the
third (South Miami–Dade
County) to Southeast Asia. The
Georgia population stems from
Japan or Korea, which may be
why it is cold-tolerant.
Habitat. Swamp eels are
freshwater creatures and prefer
shallow (less than 10 ft. or 3 m
deep) sluggish, even stagnant
waters. They are found in
ponds, reservoirs, wetlands,
streams, canals, and ditches.
Swamp eels can survive rela-
tively wide temperature ranges
and tolerate cold, even freezing,
temperatures. They also are
able to withstand low oxygen
levels in water since they can
“breathe” through their skin.
During periods of drought, they
can remain burrowed in damp
mud for weeks without eating.
Their adaptability to a wide Top: The so-called Asian swamp eel is native not only to Asia, but to
range of ecological conditions Central and South America as well. It may also be native to Australia.
extends to brackish and saline Bottom: The Asian swamp eel is established in several widely separated
water, and they can even crawl locations in the United States, where it has apparently been introduced
across land if moist enough. as a food fish. (Adapted from Nico and Fuller 2009.)
During the day, Asian swamp
eels burrow in wet mud or hide in crevices or beneath dense vegetation.
Diet. Nocturnal predators, swamp eels consume a variety of animals, including oligo-
chaete worms, aquatic insects, amphipods, crayfish, tadpoles, frogs, turtle eggs, fish eggs,
and other fish. They also eat detritus.
Life History. Asian swamp eels go through their complete life cycle in freshwater.
Reproduction has been reported throughout the year. Eggs are laid in floating bubble nests
near the mouths of burrows and are guarded by males. All young hatch as females, but some
mature females are transsexual and transform into males after a yearlong nonfunctional
stage. The males are larger than the females.
Impacts. To date, negative impacts on native ecosystems are more of a threat than a real-
ity. Their broad range of tolerances, diet, and ability to move over land give Asian swamp
162 n VERTEBRATES (FISH)

A. Adult Asian swamp eel. (U.S. Geological Survey.) B. Close-up of head. (U.S. Geological Survey.)

eels the potential to be aggressive invasive organisms. Furthermore, they have no known
predators in the United States. Their burrowing habits and nocturnal activity periods could
allow them to spread without being detected. They might reduce populations of native prey
species as well as native predators such as large fish, frogs, turtles, and wading birds that
compete for the same food source. A major concern is that they will invade Everglades
National Park. The Homestead, Florida, collection site is only 0.5 mi. (0.8 km) from the
park.
Management. Because of their ability to breathe air, control by usual fish eradication
methods of poisoning water is difficult. Electrical barriers may prevent movement to new
areas. Removal of vegetation could help. Electro-fishing devices are the only way to capture
and detect Asian swamp eels for research and monitoring. The best practice is prevention:
Do not stock these fish; do not release pet aquarium eels (or any other organisms) into local
waters; and do not transport them as bait, forage, or aquarium pets.

Selected References
Bricking E. M. “Asian Swamp Eel.” Introduced Species Summary Project, Columbia University,
2002. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/Monopterus
_albus.html.
Hamilton, H. Frequently Asked Questions about the Asian Swamp Eel. Florida Integrated Science Center,
Gainesville, FL: USGS, 2006. http://fl.biology.usgs.gov/Nonindigenous_Species/Swamp_eel_FAQs/
swamp_eel_faqs.html.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Ecology of Monopterus albus.” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=446&fr=1&sts.
Nico, Leo, and Pam Fuller. “Monopterus albus.” USGS Nonindigenous Aquatic Species Database,
Gainesville, FL, 2009. Revised November 17, 2008. http://nas.er.usgs.gov/queries/FactSheet.asp
?speciesID=974.
Rotham, Carly J. “Asian Swamp Eel Threatens N.J. Wildlife,” New Jersey Real-Time News, 2008. http://
www.nj.com/news/index.ssf/2008/09/asian_swamp_eel_threatens_nati.html.
 BIGHEAD CARP n 163

n Bighead Carp
Scientific name: Hypophthalmichthys nobilis
Synonym: Aristichthys nobilis
Family: Cyprinidae
Native Range. China, where it is native to lowland rivers of the North China Plain and
South China, including the Yangtze (Chiang Jiang), Pearl, and West Xi Kiang.
Distribution in the United States. Established in the middle and lower Mississippi and
Missouri rivers, in Illinois and Missouri. It may also be established in the Tallapossa
Drainage in Sougahatchee Creek and Yates Reservoir, Alabama; in the Big Muddy, Cache,
and Kaskaskia rivers in Illinois;
and in a backwater outlet of
the Black River, Louisiana. It
has been reported in Arizona,
Arkansas, California, Colorado,
Florida, Hawai’i, Indiana, Iowa,
Kansas, Kentucky, Louisiana,
Mississippi, Nebraska, Okla-
homa, South Dakota, Tennessee,
Texas, Virginia, and West
Virginia.
Description. Bighead carp
are deep-bodied, laterally com-
pressed fish with, as their name
implies, very large heads. A com-
plete lateral line arcs ventrally in
the anterior part of the body.
Scales on the body are tiny; the
head and opercle are scaleless.
Body color is dark gray on top
and off-white on sides and belly.
Mature specimens have dark
grayish blotches on the top of
the body. Young up to eight
weeks old are silvery. On the
underside, a distinct, smooth
keel runs from near the base of
the pectoral fins to the vent.
The large upturned mouth has
bony, rigid lips without barbels;
the lower jaw protrudes slightly
beyond the upper one. There
are no teeth in the jaws. The eyes
are close to the mouth and lie on
Top: The bighead carp is native to the lowland rivers of eastern China.
the body midline. Fins of small Bottom: Bighead carp are invasive in the Mississippi-Missouri River sys-
individuals lack spines, but large tems and are reported in a number of other waterways. (Adapted from
specimens have a heavy, stiff, Nico and Fuller 2009.)
164 n VERTEBRATES (FISH)

non-serrated spine at the origin

of the dorsal fin, which has 8–9
soft rays. A slightly stiffened
spine appears at the origin of the
anal fin, which is hooked and
has 13–14 soft rays. Pectoral fins
on large males have sharp, non-
serrated ridges along several of
the anterior rays. The gill rakers
are long and closely spaced, but
are not fused together. The pha-
ryngeal tooth count is 4–4.
Large individuals may be more
than 4 ft. (1.2 m) long and weigh
over 100 lbs. (45 kg). Males are
larger than females.
Related or Similar Species.
Silver carp (Hypophthalmichthys
molitrix), another invasive
Asian carp (see Fish, Silver
Carp), is a close relative and
occurs in many of the same pla-
ces as bigheads. The silver carp
has a longer keel that runs from
the base of the anal fin to the
isthmus at the base of the gills.
Its body color is greenish dor-
sally and silvery below the mid-
line; it lacks the blotches of
mature bigheads. The anal fin
has 12–13 rays, and pectoral
fins have 15–18 rays and a hard
spine with serrated rear margin.
Bighead carp can exceed 4 ft. in length and weigh more than 100 lb. (U.S. Gill rakers are fused into a plate
Geological Survey.)
and covered with a net-like,
porous matrix with which they strain the smaller phytoplankton from the water. Silver carp
are somewhat smaller than bighead carp, attaining a length of 3 ft. (1.0 m) and a weight of
60 lbs. (27 kg). Silver carp are known for leaping high into the air when disturbed.
Juvenile bigheads can be confused with gizzard shad (Dorosoma cepedianum; see Fish,
Gizzard Shad), often used as bait fish.
Introduction History. Both bighead and silver carp were imported into Arkansas in 1972
by a catfish farmer interested in their potential for improving water quality in his fish ponds
by their ability to consume huge amounts of algae. The fish spread to other aquaculture oper-
ations for the same purpose. Carp first appeared in the open waters of the Ohio and
Mississippi rivers in the early 1980s; they probably escaped from culture ponds. Thousands
of bighead carp made their way into the Osage River, Missouri, when a fish hatchery near
Lake of the Ozarks was inundated by floodwaters in April 1994. Fish that had escaped into
the Missouri River multiplied and spread into the lower Kansas River in Kansas after 1990.
 BIGHEAD CARP n 165

Hojo-e

B ighead carp are a food fish for some Asian Americans and are usually pur-
chased live at markets in large cities such as Chicago and New York. Cultural
practices among Chinese, Vietnamese, and other Asian immigrant groups could
provide a potential pathway for the fish to enter the Great Lakes. As part of a
Buddhist ceremony known as hojo-e, releasing captive animals secures merits for
the afterlife and lengthens the life of the practitioner. The ritual commonly takes
place at Buddhist temples under the guidance of monks. In the United States, gold-
fish, turtles, and birds are usually released, but the practice may explain the
appearance of Asian carp in public ponds and lagoons in the Great Lakes region.
Chicago and New York now both have regulations requiring that Asian carp be
killed before they are sold.
Source: Higbee, E., S. Fellow, and K. G. Shwayder. “The Live Food Fish Industry: New
Challenges in Preventing the Introduction and Spread of Aquatic Invasive Species.” Great
Lakes Panel on Aquatic Nuisance Species, ANS Update 10(2): Fall–Winter 2004. Online at
http://www.glc.org/ans/ansupdate/pdf/2004/ANSUpdateFW.pdf.

Illegal introductions by commercial fish farmers in the late 1980s are responsible for carp
in Grand River, Oklahoma; California; and Cherry Creek Reservoir, Colorado.
Habitat. Bighead carp are native to subtropical and temperate freshwater habitats, prefer-
ring large rivers and the lakes connected to them. They usually are found in the upper or
euphotic layer of the water column, where food is most abundant. They may migrate up
streams from lakes to spawn, since they require a current for their eggs to float and develop
properly.
Diet. Bighead carp filter relatively large particles (10–100 μm) from the water. As plank-
ton feeders, their diet changes as they grow. Larvae consume mostly small phytoplankters
such as protococcaceans and cyanobacteria; but large individuals specialize on larger par-
ticles, including zooplankters such as cladocerans and midge larvae and algae such as dia-
toms and colonial phytoplankters. They are opportunistic feeders, however, and what they
eat depends upon the type of suspended materials that are most abundant. They have been
said to consume their weight in plankton each day. They also feed on detritus.
Life History. Males and females mature in 2–4 years in warmer, subtropical waters and
5–7 years in cooler, temperate waters. They are prolific breeders, with females producing
up to a million eggs during their lifetimes. Eggs have diameters near 0.2 in. (4.5–5 mm).
Spawning takes place in the spring as water levels in rivers rise. Adults may migrate
100 miles or more upstream to breed. Eggs and larvae then float downstream to the lower
reaches of rivers and to lakes. Intolerant of brackish water, carp spend their entire lives in
freshwater.
Impacts. As voracious plankton and detritus feeders, bighead carp can outcompete native
filter-feeding organisms such as mussels, fish larvae, and some adult fishes. The adult fish
most likely to suffer from carp invasions are paddlefish (Polydon spathula), bigmouth buffalo-
fish (Ictiobus cyprinellus) and gizzard shad (Dorosoma petense). Bighead carp have the poten-
tial of disrupting entire aquatic food webs. Their introduction into the Great Lakes system is
particularly feared for this reason.
166 n VERTEBRATES (FISH)

Management. The main focus of management is preventing the introduction of bighead

carp into the Great Lakes. They already inhabit the Illinois River, which connects to Lake
Michigan via the Chicago Sanitary and Ship Canal. By 2011, the U.S. Fish and Wildlife
Service, the EPA, the U.S. Army Corps of Engineers, the State of Illinois, the International
Joint Commission, and the Great Lakes Fishery Commission had completed a
multimillion-dollar series of three electric dispersal barriers on the canal.

Selected References
“Asian Carp: Key to Identification.” U.S. Fish and Wildlife Service, 2002. http://www.fws.gov/
Midwest/fisheries/library/broch-asiancarpkey.pdf.
Bighead and Silver Carp (Hypophthalmichthys nobilis and H. molitrix). Wisconsin Department of Natural
Resources, 2004. http://dnr.wi.gov/invasives/fact/asian_carp.htm.
Nico, Leo, and Pam Fuller. “Hypophthalmichthys nobilis.” USGS Nonindigenous Aquatic Species
Database, Gainesville, FL, 2009. Revised November 10, 2010. http://nas.er.usgs.gov/queries/
FactSheet.asp?speciesID=551.
U.S. Army Corps of Engineers, Chicago Sanitary and Ship Canal Aquatic Nuisance Species Dispersal
Barrier System, 2009. http://www.lrc.usace.army.mil/AsianCarp/BarriersFactSheet.pdf.

n Brown Trout
Also known as: German trout
Scientific name: Salmo trutta
Synonyms: Salmo fario, Fario argenteus
Family: Salmonidae
Native Range. Eurasia. In Europe, this anadromous fish is native to Atlantic, Baltic, and
Black Sea and Caspian Sea drainages. In western Asia, it is reported as native to
Afghanistan, Armenia, and Turkey. It is also considered native in parts of North Africa.
The fish has been introduced into areas where it is not native on every continent except
Antarctica. Those imported into the United States came from Germany.
Distribution in the United States. The brown trout has been introduced into almost all
50 states and the Commonwealth of Puerto Rico. The only states without brown trout are
Alaska, Louisiana, and Mississippi. The fish may not be breeding in most states, but it is con-
tinually restocked for recreational fishing.
Description. Brown trout are so named because of their brown to golden-brown color.
The sides are yellow or silvery and bellies white to yellow. Red spots with blue halos and
black spots adorn the sides of stream-dwelling browns, but are faint on lake-dwelling indi-
viduals. The lateral line is iridescent when light hits it from the right angles. There are two
dorsal fins, the rear one a small fatty (or adipose) fin with a reddish color. The anal fin has
9–10 rays. The tail is square. Adults in the United States can be 13–16 in. (33–40 cm) long
and weigh up to 10 lbs. (4.5 kg).
Related or Similar Species. Atlantic salmon (Salmo salar) are close relatives. They have no
red on the adipose fin and the tail is slightly forked. Rainbow trout (Onchorhynchus mykiss)
have lines of black spots on the tail (see Fish, Rainbow Trout, for a fuller description).
Introduction History. Brown trout were first imported from German hatcheries into the
United States in 1883, when a shipment of 80,000 eggs landed at Cold Spring Harbor, New
York. These eggs were distributed by the U.S. Fish Commission to the Caledonia Fish
 BROWN TROUT n 167

hatchery in New York and the

Norville Hatchery in Michigan.
In 1886, Pennsylvania began
stocking brown trout in streams
where native brook trout popu-
lations had been extirpated or
reduced by practices such as
logging, farming, dam construc-
tion, and industrial discharges,
all of which had led to warmer
water temperatures and siltation.
Since then, brown trout have
been stocked by state and local
agencies across the country for
sport fishing.
Habitat. Although anadro-
mous in its native range, brown
trout in the United States are
freshwater fish and prefer
streams and lakes. They hide
during the day in shallow beds
of aquatic vegetation, in shallow
rock-strewn areas, under sub-
merged logs, or in deep pools.
They are most active at dawn
and dusk and when the water
temperature is near 55°F
(12.8°C). They generally prefer
water temperatures between
65° and 75°F (18–24° C) and
tolerate warmer water tempera-
tures than do native brook trout. Top: The brown trout is native to Europe and western Asia. It has been
Diet. Smaller individuals widely introduced throughout Eurasia, making delineation of the actual
feed on insects such as mayflies, native range difficult. Bottom: Brown trout have been introduced for
caddisflies, and midges that the sport-fishing in all states except Alaska, Louisiana, and Mississippi.
stream carries to them. Larger (Adapted from Fuller, P. “Salmo trutta.” USGS Nonindigenous Aquatic
brown trout have a broader diet Species Database, Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/
speciesmap.aspx?speciesID=931.)
that includes large aquatic
insects, mayflies, caddisflies,
crustaceans, snails, amphibians, and small fish. They actively hunt their prey primarily at
night. Both small and large trout also consume detritus washed from the shore.
Life History. Brown trout spawn from October to December in shallow headwater streams,
ideally where the water is about 1 ft. (30 cm) deep, the current is about 7 in./second (18 cm/
second), and substrate particles are small (diameters of ca. 0.5 in. or 1.27 cm). The female hol-
lows out a nest, or redd, in which to release her eggs. As she releases her eggs, the male releases
milt to fertilize them. The female covers the fertilized eggs with sand and fine gravel, and both
adults leave. The larvae stay in the redd for 2–3 weeks until they are about 0.1 in. (25 mm)
168 n VERTEBRATES (FISH)

long. They then move down-

stream or into lakes for the first
two years of their life. Brown
trout begin to establish territo-
ries when they are juveniles.
They mature at 3–4 years of age.
Many populations in the
United States are maintained
through restocking efforts rather
than natural reproduction.
Impacts. Brown trout may
compete with native fish, espe-
cially other salmonids (trout
and salmon) for food. They
Red spots with blue halos and black spots appear on brown trout but are grow larger and more rapidly
faint on lake-dwelling individuals. (Tramper/Shutterstock.) than native species. They
reportedly have also reduced
populations of native trout through predation. As aggressive defenders of territories, they
can displace native fishes from prime habitat. They have been implicated in population
reductions of brook trout (Salvelinus fontinalis) and Modoc sucker (Catostomus microps), an
endangered fish in California. Although it rarely happens, brown trout can hybridize with
native trout. Brown trout has been nominated as one of “100 of the ‘World’s Worst’ invasive
alien species” by the IUCN and Global Invasive Species Programme.
Management. Brown trout are regularly stocked into most waters. Sport fishing often
reduces numbers so that annual restocking is necessary. In New York, programs aimed at
improving water quality for native brook trout become control methods for brown trout.
In California, attempts are being made to eliminate brown trout where they may compete
with golden trout (Oncorhynchus mykiss aguabonita).

Selected References
“Brown Trout in Pennsylvania.” Pennsylvania Council of Trout Unlimited, n.d. http://www.patrout.org/
Documents/Reference/brown.pdf.
Idema, A. “Salmo trutta.” Animal Diversity Web, Museum of Zoology, University of Michigan, 1999.
http://animaldiversity.ummz.umich.edu/site/accounts/information/Salmo_trutta.html.
Lauterbach, Sandra. “Brown Trout (Salmo trutta).” Introduced Species Summary Project, Columbia
University, 2006. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/
Salmo_trutta.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialty Group
(ISSG). “Salmo trutta (fish).” ISSG Global Invasive Species Database, 2006. http://www.issg.org/
database/species/ecology.asp?si=78&fr=1&sts=sss&lang=EN.

n Gizzard Shad
Also known as: Hickory shad, mud shad, nanny shad, skipjack, winter shad
Scientific name: Dorosoma cepedianum
Family: Clupeidae
 GIZZARD SHAD n 169

Native Range. Native to

eastern and central North
America in the Mississippi,
Atlantic and Gulf coast drainage
systems. Probably native in the
Arkansas, South Platte, and
Republican drainage systems in
eastern Colorado. Possibly
native to the St. Lawrence River
and the Great Lakes.
Distribution in the United
States. The gizzard shad has
been introduced to reservoirs
and natural bodies of water
both within and peripheral to
its natural range. It is nonnative
but established in the Colorado
and Salt rivers and their con-
nected reservoirs in Arizona; in
the upper Colorado River dra-
inage system in Arizona,
Colorado, and Utah; and in
Wyoming. Within its native
range, its distribution has been
extended by stocking in Colo-
rado, Illinois, Indiana, Kansas,
Kentucky, Minnesota, Nebra-
ska, Pennsylvania, Utah, and
Virginia. They may or may not
be native to the Great Lakes,
where they are found in all
lakes except Lake Superior. Top: The gizzard shad is native to the eastern and central United States.
Relatively recent natural range Bottom: Gizzard shad have been introduced to rivers, lakes, and
expansion probably explains reservoirs within and beyond the borders of its native range. (Both maps
their occurrence in Connect- adapted from Fuller 2009.)
icut (Connecticut River), Maine
(lower Saco and Kennebec rivers), Massachusetts (Connecticut and Merrimack rivers), and
Vermont (Lake Champlain and Connecticut River).
Description. This is a deep-bodied, moderately compressed herring that reaches lengths
of 22 in. (56 cm) and weights of up to 7 lbs. (3.2 kg). It has a dark-blue or gray back, silvery
sides that may reflect various colors, and a white belly. A dark, purplish blotch occurs high
on the side behind the opercle in young and small specimens; it is faint or absent in older,
larger individuals. It has no lateral line. The small mouth has a deep notch in the center of
the upper jaw, which protrudes slightly beyond the lower jaw. The snout is blunt, and the
eye large. The short, soft-rayed dorsal fin is centered on the back and has 10–12 rays; the last
ray is a long filament that is distinctive for the species. The tail (caudal fin) is deeply forked.
Scute-like scales form a distinct saw-toothed keel along the belly. Gill rakers are fine and
170 n VERTEBRATES (FISH)

number more than 400. The stomach is thick-walled and acts like a gizzard, giving the fish
its common name.
Related or Similar Species. Threadfin shad (D. petense) is a smaller fish, rarely more than
8 in. (20 cm) long. The ray extending from its dorsal fin is much longer than that of gizzard
shad and almost reaches the tail. The mouth especially distinguishes the two: in the thread-
fin, the lower jaw extends beyond the upper jaw—the reverse of the situation in gizzard
shad. Fins of threadfin shad tend to be yellowish, whereas those of gizzard shad are grayish.
Introduction History. In most cases, gizzard shad were intentionally introduced to
ponds, lakes, and reservoirs as forage fish, especially in recreational sport fisheries. From
their sites of entry, however, they have dispersed rapidly into connecting bodies of water,
often assisted by man-made facilities. Gizzard shad in the Upper Colorado River basin of
Arizona, Colorado, and Utah may trace their origins to an accidental introduction in a con-
taminated stocking of largemouth bass from a hatchery in Texas into Morgan Lake, New
Mexico, near the San Juan River around 1996. Gizzards were first reported in the San Juan
arm of Lake Powell in 2002 and are now found throughout the lake. From Lake Powell, they
dispersed downstream to Lake Mead and, by 2008, were found in the Grand Canyon stretch
of the Colorado River. They also dispersed upstream into the headwaters of the Colorado: in
2006 they were collected in the Gunnison and Middle Green rivers. Gizzard shad were
stocked in Lake Havasu in the lower Colorado River drainage and, from there, have dis-
persed upstream as far as Davis Dam and downstream to the Mexican border and into the
Salton Sea.
In Connecticut, Massachusetts, and Vermont, gizzard shad have expanded their range
upstream in the Connecticut River. They were first recorded at the mouth of the river around
1980, part of what seems to be a natural range expansion northward along coastal rivers,
which has also brought them into the Merrimack River in Massachusetts and lower Saco
and Kennebec rivers in Maine. By 1986, they had appeared at Holyoke Dam and now occur
in the mainstem of the Connecticut River as far north as the Bellows Falls dam. Fish “lad-
ders” built to let American shad (Alosa sapidissima) and Atlantic salmon (Salmo salar) circum-
vent the dams on their annual spawning runs have aided this movement. Gizzards entered
Lake Champlain, Vermont, through the Hudson Barge Canal.
Some believe gizzard shad may be native to the Great Lakes and St. Lawrence River, but it
is also possible that they entered at Green Bay, Wisconsin, from the Mississippi River via the
Fox-Wisconsin Canal or via the Chicago River Canal. Entry into Lake Erie was facilitated by
the Ohio Canal.
Gizzards in Wyoming stem from intentional introductions made in Nebraska.
Habitat. Freshwater lakes and reservoirs, slow-moving rivers, and pools of smaller
streams are preferred habitat. Gizzard shad can also be found in the brackish waters of
estuaries. Native populations in the Mid-Atlantic states are anadromous. They often feed in
schools over mucky and sandy bottoms.
Diet. Gizzard shad are filter-feeders and, as adults, strain algae and detritus from bottom
sands and muds. They also ingest sand to aid in grinding food in their gizzard-like stomachs.
Young shad capture zooplankters (copepods and cladocerans) from the water column. Shad
feed in large schools and characteristically leap from the water and flip onto their sides, a
behavior that gave them the common name of skipjack.
Life History. Gizzard shads spawn near shore during spring nights as water temperatures
rise near 70°F (21°C). Spawning may continue for several weeks, after which adults return
 GIZZARD SHAD n 171

to deeper water. A single female

may produce 300,000 or more
eggs. Eggs and milt are broad-
cast near the surface in shallow
water, and fertilized eggs settle
down through the water and
stick on roots, plants, and
debris. They hatch in 2–3 days.
Larvae grow rapidly and,
during their first year, young
shad may reach lengths of 4–7
in.—too large to be prey for all
but the largest predatory fish.
Most are sexually mature at age
2. Life span is 2–3 years.
Adults experience die-offs in The gizzard shad, a member of the herring family, can attain a length of
winter and after spawning. 22 in. and weigh as much as 7 lb. (Eric Engbretson.)
Impacts. As larvae, gizzard shad may compete with the young of other, more desirable fish
for zooplankton. Not only do shad populations grow rapidly, but they also spawn earlier than
some other fish and thus may deplete the food resource enough to negatively affect growth
and survival of sport and other fishes. They are usually stocked as a forage fish, but their rapid
growth rate soon makes them too large to be taken by most predatory fishes. However, while
they are small, they can be important prey for striped bass, largemouth bass, crappie, walleye,
and other sport fishes. The closely related threadfin shad, also introduced to reservoirs as for-
age and baitfish, do not grow so large and so are favored by sport fisheries managers, but they
too may be outcompeted by gizzard shad. In the Colorado River, gizzard shad are viewed as
one more threat to endangered endemic fishes such as Colorado River pike minnow
(Ptychocheilus lucius), bonnytail chub (Gila elegans), humpback chub (Gila cypha), and razor-
back sucker (Xyrauchen texanus) as well as to a number of other sensitive species.
Management. Little to none.

Selected References
Finney, Sam T., and Mark H. Fuller. “Gizzard Shad (Dorosoma cepedianum) Expansion
and Reproduction in the Upper Colorado River Basin.” Western North American Naturalist 68(4):
524–25, 2008.
“Fish Facts—Gizzard Shad.” U.S. Fish and Wildlife Service, Connecticut River Coordinator’s Office,
n.d. http://www.fws.gov/r5crc/Fish/zc_doce.html.
Fuller, Pam. Dorosoma cepedianum. USGS Nonindigenous Aquatic Species Database, Gainesville, FL,
2009. Revised March 10, 2008. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=492.
“Gizzard Shad Dorosoma cepedianum,” Utah Division of Wildlife Resources, n.d. http://
wildlife.utah.gov/pdf/AIS_plans_2010/AIS_12nGizzardShad-Dan-final.pdf.
“Shad Species Used in Striped Bass Fishing.” Ben Sanders’ ArkansasStripers.com, n.d. http://
www.arkansasstripers.com/shad_species.htm.
Steiner, Linda “Herrings,” Chapter 10 in Pennsylvania Fishes. Pennsylvania Fish and Boat Commission,
2000. http://www.fish.state.pa.us/pafish/fishhtms/chap10.htm.
172 n VERTEBRATES (FISH)

n Grass Carp
Also known as: White amur
Scientific name: Ctenopharyngodon idella
Family: Cyprinidae
Native Range. Native to eastern Asia, from the Amur River, Russia, to the West River in
southern China. They have been widely distributed throughout the world as a food fish.
The first imported into the United States reportedly came from Taiwan and Malaysia.
Distribution in the United States. Grass carp have been widely used to control aquatic
vegetation and have been reported from 45 states. States without records of grass carp are
Alaska, Maine, Montana, Rhode Island, and Vermont. The fish is established in parts of
the Mississippi River drainage, including the Mississippi River itself, in Arkansas,
Kentucky, Illinois, Louisiana, Montana, and Tennessee; the Illinois River in Missouri, the
lower Missouri River in Montana; the Ohio River in Illinois; and the Trinity River in Texas.
Description. The grass carp is a large fish, often attaining a total length greater than 3 ft.
(1.0 m) and a weight of 40 lbs. (18 kg) or more. It has a thick, oblong body with a rounded
belly and broad head. The anal fin is set close to the caudal fin, which is forked. The dorsal
fin is short, with 7–8 rays and set over the pelvic fins. All fins are soft-rayed and usually
green-gray to dull silver. The lateral line is complete and slightly down-curved. Eyes are
positioned at or slightly above midline of body. Body color is silver to pale gray, darker on
the back and brassy on the sides. The large scales on the back and sides have distinct dark
edges that give a characteristic cross-hatched pattern to the body. The pharyngeal teeth are
long and serrated, with deep, parallel grooves. Gill rakers are short, unfused, and widely
spaced.
Related or Similar Species. Black carp (Mylopharyngodon piceus), another introduced
Asian carp but one of much more restricted distribution in the United States, is darker in
color, although not black. Its pharyngeal teeth are smooth and resemble human molars. It
is a bottom-dwelling species that feeds on snails and other mollusks. All native cyprinids
(minnows) have anal fins positioned more anteriorly than grass carp.
Introduction History. Grass carp first entered the United States in 1963, when they were
imported by the aquaculture program at Auburn University, Alabama, and the Fish Farming
Experimental Station in Stuttgart, Arkansas. Fish escaping from the latter facility in 1966
were the first released into the White River, but later intentional stockings in Arkansas took
place in lakes and reservoirs with free passage to river systems. Grass carp were reported in
the Illinois section of the Mississippi River in 1971. Soon thereafter, grass carp were being
commonly reported in the Mississippi and Missouri rivers. The fish has been widely dis-
persed since then through legal and illegal interstate transport and release by private individ-
uals and organizations, escapes from farm ponds and aquaculture facilities, and stockings by
federal, state, and local government agencies, among them the U.S. Fish and Wildlife
Service, the Tennessee Valley Authority, and state fish and game agencies in Arkansas,
Delaware, Florida, Iowa, New Mexico, and Texas.
Habitat. Grass carp prefer shallow, quiet waters in lakes, pools, and the backwaters of
large rivers that have abundant aquatic vegetation. They undertake long spawning migra-
tions into faster-moving rivers.
Diet. A herbivore, the grass carp feeds on algae, submerged and floating aquatic macro-
phytes, and even overhanging terrestrial plants. Although they are known to consume hun-
dreds of different kinds of aquatic plants, they show distinct preferences. Young fish select
 GRASS CARP n 173

soft, succulent species such as

filamentous algae, pondweeds,
elodea, duckweeds, chara, and
the soft, growing tips of sub-
merged plants. As they grow
older and larger, they add more
fibrous, less succulent plants
and tend to avoid filamentous
algae. Adults will eat such inva-
sive plants as waterhyacinth
(see Volume 2 Aquatic Plants,
Waterhyacinth, Eichhornia cras-
sipes), hydrilla (see Volume 2
Aquatic Plants, Hydrilla, Hy-
drilla verticillata), and Eurasian
watermilfoil (see Volume 2,
Aquatic Plants, Eurasian water-
milfoil, Myriophyllum spicatum),
especially if more palatable spe-
cies are not present. They may
also take insects and other
invertebrates and detritus, par-
ticularly if it is attached to
aquatic plants.
Food consumption rates
vary with water temperature,
age and size of fish, and plant
species available. Optimal con-
sumption occurs when water
temperatures are between 70°
and 86°F (21–30°C). Carp feed
only intermittently when tem- Top: The grass carp is native to eastern Asia. Bottom: Grass carp,
peratures fall into the 30s introduced to control aquatic plants, have been reported from 45 states.
(below 0°C). Essentially a (Adapted from Nico, Fuller, and Schofield 2009.)
warm-water fish, they are dor-
mant during the cold winters of temperate regions.
Life History. Grass carp make long annual migrations to spawn in fast-moving large riv-
ers. They often congregate in areas of turbulent water such as in rapids or at falls or the base
of dams. Spawning occurs in the spring as water temperatures rise into the 60s and 70s (15–
low 20s°C). The eggs must stay afloat in order to develop, which they do as they drift down-
stream, perhaps as much as 110 miles. Eggs hatch in 2–3 days, and larvae shelter in the veg-
etated areas of floodplains and lakes. They may winter in deep pools in rivers. Grass carp
become sexually mature between two and five years of age in the subtropics and four to
seven years or longer in cooler regions. Females usually mature a year later than males.
Females produce 500,000–700,000 eggs, giving carp enormous reproductive potential.
Impacts. Grass carp are stocked to remove aquatic weeds, many of which are also classi-
fied as invasive. This may be advantageous for some game fish, but overconsumption by
174 n VERTEBRATES (FISH)

carp can also modify habitats

and alter the species composi-
tion and trophic structure of
ecosystems by eliminating the
aquatic vegetation that provides
food, breeding sites, and cover
for native invertebrates, fishes,
and birds. They may destroy
The large, black-edged scales on the back and sides of grass carp give spawning substrates, disturb
them a distinctive appearance. (Oleg_Z/Shutterstock.) sediments and increase water
turbidity, increase nutrient levels
in the water (their short gut makes their digestive system very inefficient, so large volumes of
waste are produced), promote algal blooms, and decrease oxygen levels. Reportedly, they have
contributed to reductions in populations of bluegill (Lepomis macrochirus), sunfish, smelt, and
pike. Grass carp also have been implicated in the declining numbers of certain waterfowl such
as Gadwall (Anas stepera), American Wigeon (Anas americana), and American Coot (Fulica
americana) in areas to which the fish has been introduced.
Carp may transmit several parasites and diseases to native fish. They are probably the
source of the Asian tapeworm (Bothriocephalus opsarichthydis), which has infected some
native cyprinids, including the endangered woundfin (Plagopterus argentissimus), a small
minnow endemic to the Virgin River, a tributary of the Colorado River.
Although considered an invasive species, grass carp is still stocked and even recom-
mended as a biocontrol method for noxious weeds (see sidebar on Triploid Grass Carp).
Management. All states have restrictions on the importation of grass carp for aquatic
weed control, but most allow stocking with proper permitting. Because of the reproductive
potential and propensity to disperse, and because of environmental concerns, sterile triploid
fish (see sidebar on Triploid Grass Carp) are widely used today for weed control. Fish
screens may be required at all inlets and outlets to a water body to prevent grass carp from
migrating out of lakes and into rivers. Carp may or may not have the desired effect of remov-
ing aquatic weeds. They seem to be an all-or-nothing proposition and should be stocked
only where complete elimination of submerged vegetation is desired or can be tolerated.

Triploid Grass Carp

S cientists have sought ways to reduce the likelihood of reproduction in grass

carp and thereby control their spread into open waters. Crossing female grass
carp with male bighead carp produced 100 percent triploid hybrids that were ster-
ile. Triploid means each fish has three copies of each chromosome instead of the
normal two (diploid). Later, it was discovered that shocking the eggs (in fish hatch-
eries) with cold or heat under high hydrostatic pressure also produced triploids,
and this became the standard method of producing sterile fish. Since triploids have
larger red blood cells and nuclei than diploids, the two are readily separated in a
simple blood test. The U.S. Fish and Wildlife Service now has an established testing
and inspection protocol to screen all fish leaving hatcheries and to guarantee that
only triploids are imported into those states that do not permit the introduction of
diploid grass carp.
 LIONFISH n 175

Selected References
“Grass Carp, Ctenopharyngodon idella (Valenciennes 1844).” Florida Integrated Science Center, USGS,
2005. http://fl.biology.usgs.gov/Carp_ID/html/ctenopharyngodon_idella.html l.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Ctenopharyngodon idella (Fish).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=369&fr=1&sts=sss&lang=EN.
Nico, L. G., P. L. Fuller, and P. J. Schofield. “Ctenopharyngodon idella.” USGS Nonindigenous
Aquatic Species Database, Gainesville, FL, 2006. http://nas.er.usgs.gov/queries/FactSheet.asp
?speciesID=514.

n Lionfish
Also known as: Red lionfish, turkeyfish, devil firefish, zebrafish
Scientific name: Pterois volitans/P. miles
Family: Scorpaenidae (Scorpionfishes)
Native Range. The Indo-Pacific Ocean region, from southern Japan to the Philippines to
Micronesia and the east coast of Australia through western Polynesia to the Marquesas Island
and Oeno in the Pitcairn islands. The red lionfish (P. volitans) is mostly a Pacific Ocean fish;
its range extends eastward through Oceania; the devil firefish (P. miles) occurs primarily in
the Indian Ocean, from Sumatra northwestward into the Red Sea. Both fishes are found off
Sumatra. The devil firefish has passed through the Suez Canal into the Mediterranean Sea.
Whether or not these are two true species or simply regional variants of the same species
has not been conclusively determined. Genetic studies indicate that both now occur as
exotic taxa in the Atlantic Ocean off the U.S. coast, but the Atlantic population consists pri-
marily of P. volitans. Their likely source area is the Philippines.
Distribution in the United States. Apparently established and reproducing from the
Florida Keys to Cape Hatteras, North Carolina. Divers have reported sightings of juveniles
as far north as Long Island, New York, New Jersey, and Rhode Island; but it is assumed they
were carried north in the Gulf Stream as the young of the year and will either migrate south
or die in winter. Winter bottom-water temperatures may create a northern limit to their dis-
tribution area as it does for other tropical fishes whose young are frequently encountered off
the coast of the northeastern United States in summer.
Description. Lionfish are unmistakable. The red lionfish has long, separated dorsal fin
spines. The appearance of the dorsal spines together with the elaborate fan-like pectoral fins
resulted in the common name, turkeyfish. The body is white to cream-colored with red-to-
maroon vertical stripes that gave it another of its common names, zebrafish. The stripes have
a regular, alternating pattern between wide and narrow bands and sometimes converge in a
V on the sides. Fleshy tabs or tentacles occur over the eyes and above the mouth.
The dorsal, anal, and pelvic spines of lionfish are highly venomous. A pair of venom
glands is found on each spine, which is covered by a sheath. When a spine is pushed into
flesh, the sheath presses down on the glands, and venom is released into a groove that runs
along the spine, delivering the poison to the wound. The resulting sting is extremely painful,
but not fatal to humans.
The largest lionfish collected off the East Coast of the United States was about 17 in.
(43 cm) long and weighed about 2.5 lbs. (1.1 kg).
Related or Similar Species. The devil firefish and the red lionfish are very similar in
appearance. They are distinguished from each other by the number of rays in the dorsal
176 n VERTEBRATES (FISH)

and anal fins and by genetic

assays. The lionfish usually has
11 dorsal-fin rays and 7 anal
fin rays, whereas the devil fire-
fish has 2 and 6, respectively.
The two fishes are very closely
related and may actually be the
same species. A 2007 study
showed that the composition of
the Atlantic population was
97 percent lionfish and 7 per-
cent devil firefish.
Introduction History. The
first known occurrence of lion-
fish in the Atlantic off the
United States was a specimen
collected near Dania, Florida,
in October 1985. Later, six fish
escaped into Biscayne Bay from
a large private aquarium that
was destroyed during Hurri-
cane Andrew in 1992. These
fish were seen alive in the bay
a few days later. Sporadic or
undocumented sightings of red
lionfish were reported from
Florida’s east coast between
1993 and 2002. In February
and March 2002, three speci-
mens were taken off northeast
Florida near St. Augustine,
Top: Lionfish are native to the Indo-Pacific region. Bottom: Lionfish Jacksonville, and Amelia Island.
appear to be established along the Atlantic coast of the United States from Lionfish were first recorded
the Florida Keys north to Cape Hatteras, North Carolina. Juveniles have off Georgia in 2001, when a sin-
been reported as far north as Rhode Island. (Adapted from Schofield, gle adult was collected. The same
Morris, Langston, and Fuller 2009.) year, two juveniles were taken
off Long Island, New York.
Small groups of lionfish were observed during research dives in submersibles off South
Carolina and North Carolina in 2002. This was the first, though circumstantial, evidence
that the Atlantic population was reproducing.
Although water ballast is a possible route of entry for the fish, it is highly unlikely. The red
lionfish is a very popular marine aquarium species, and the aquarium trade is the most prob-
able pathway by which the fish came to the Atlantic Ocean. The color patterns of Atlantic lion-
fish are quite similar to those from the Philippines, where many are collected for the aquarium
trade. Either accidentally or deliberately, whether they had grown too large or for whatever rea-
son, these (and other) aquaria pets are being dumped into the open waters of the ocean.
The wild population is growing rapidly. That juvenile fish smaller than those sold for
aquaria are showing up off the North Carolina coast and elsewhere is a strong indication that
 LIONFISH n 177

the species is reproducing. This

is the first time a western Pacific
fish has become established in
the Atlantic Ocean off the
United States.
Habitat. Lionfish are tropical
marine fish associated with the
continental shelf biome. They
inhabit reefs at depths of 30 to
575 ft. (9–175 m), where they
can hide in crevices by day.
Along the southeastern coast of
the United States, they have
been sighted on reefs and other
hard substrates, including ship-
wrecks. In their native distribu-
tion areas, they are found in The lionfish, with its long dorsal spines and fan-like pectoral fins, is
lagoons and turbid inshore unmistakable. (Albert Kok.)
waters, including harbors, in
addition to offshore coral reefs and rocky outcrops.
Diet. Lionfish are carnivores that prey on crustaceans and small fish, including young
lionfish and juveniles of some commercially important species such as grouper and snapper.
They focus increasingly on fish as they age. Nocturnal hunters, lionfish move into deeper
waters at night to forage. They are ambush predators and sweep up and trap their prey in
their extended pectoral fins. They then quickly pounce and ingest it.
Life History. Lionfish are solitary and territorial most of the year, only forming small
groups during mating, when males join with several females. Males use their spines and fins
in competitive visual displays. Females release a pair of mucous-encapsulated clusters of
2,000–15,000 eggs that are externally fertilized by the males. The egg mass floats, and as
microbes decompose the mucous, eggs are released into the water column. Early life-
history stages are poorly known. Presumably, lionfish larvae have a pelagic stage during
which they are dispersed by ocean currents. The larval stage lasts an estimated 25–40 days.
Age at sexual maturity and average lifespan are unknown. There are few known predators of
lionfish, although off the Bahamas, lionfish have been found in the stomachs of native
groupers.
Impacts. Recent (2008) studies in the Bahamas have shown that lionfish may have nega-
tive impacts on native Atlantic reef-fishes. Reefs with lionfish had 79 percent fewer juvenile
reef fish compared to reefs lacking lionfish. Prey items found in lionfish stomachs included
fairy basslet (Gramma loreto), bridled cardinal fish (Apogon aurolineatus), white grunt
(Haemulon plumierii), bicolor damselfish (Stegastes pertitus), several wrasses, striped parrot-
fish (Scarus iserti), and dusky blenny (Malacoctenus gilli). Juvenile spiny lobster (Panulirus
argus) is possibly a food item. These findings suggest that lionfish affect recruitment of coral
fishes and may thereby compete with native piscivores. They could potentially reduce the
number of ecologically important species such as parrotfish and other herbivores that pre-
vent macroalgae from taking over the reef.
Management. Very little can be done in a marine environment to control invasive species.
Environmental changes occurring on the southeastern-U.S. continental shelf favor the con-
tinued expansion of lionfish in these waters. Many important native reef fish predators have
178 n VERTEBRATES (FISH)

Tournament Held to Help Control Lionfish Population

in the Florida Keys National Marine Sanctuary

I n September 2010, the Reef Environmental Education Foundation (REEF) and the
Florida Keys National Marine Sanctuary sponsored a lionfish hunt with cash and
other prizes for teams of divers that would catch invasive lionfish. The 27 partici-
pating teams collected a total of 534 lionfish during the weekend tournament,
the winning team catching 111 in one day. The largest was 11 in. (27.9 cm) long,
but most were too small to filet, so the planned follow-up feast was a failure.
Nonetheless, sanctuary superintendent Sean Morton said, “The sanctuary is
thrilled by the response from the dive community. The volume of fish caught
during this single-day event demonstrates that dedicated diver removal efforts
can be effective at helping keep this invasive [species] at bay.”
Source: Frink, Stephen. “The Tipping Point(s).” Alert Diver, Fall 2010: 8.

been overfished, but even if they were at past densities, it is unknown if native predators
could eventually control lionfish numbers. None of them have prior experience with a prey
fish with venomous spines. The fish fauna in general has become more tropical during the
last decades of the twentieth century, so it appears conditions increasingly favor the breed-
ing and recruitment of lionfish.

Selected References
Hare, J. A., and P. E. Whitfield. “An Integrated Assessment of the Introduction of Lionfish (Pterois vol-
itans/miles Complex) to the Western Atlantic Ocean.” NOAA Technical Memorandum NOS NCCOS
2, 21 pp., 2003. http://coastalscience.noaa.gov/documents/lionfish_ia.pdf.
“Invasive Lionfish Threaten Native Fish and the Environment in U.S. Atlantic Coastal Waters.”
National Ocean Service, NOAA, 2009. http://oceanservice.noaa.gov/facts/lionfish.html.
Masterson, J. “Pterois volitans.” Smithsonian Marine Station at Fort Pierce, 2007. http://www.sms.si.edu/
IRLSpec/Pterois_volitans.htm.
Schofield, P. J., J. A. Morris Jr., J. N. Langston, and P. L. Fuller. “Pterois volitans/miles.” USGS
Nonindigenous Aquatic Species Database, Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/
FactSheet.asp?speciesID=963.

n Mosquitofish
Scientific names: Gambusia affinis (western mosquitofish)
Gambusia holbrooki (eastern mosquitofish)
Family: Poeciliidae
Native Range. The western mosquitofish is native to the south-central United States,
from Illinois and Indiana south into Mexico, west to New Mexico, and east into the Mobile
River drainage system. Also native in the Chattahoochee and Savanna rivers. Apparently,
most introductions to other states (and worldwide) stem from a few populations from
Georgia, Illinois, Tennessee, and Texas.
The eastern mosquitofish is native to the Atlantic and Gulf coast drainages from southern
New Jersey south and west to southern Alabama.
 MOSQUITOFISH n 179

Distribution in the United

States. The western mosquito-
fish has been widely introduced
into most western states. The
eastern mosquitofish has likely
been transplanted outside its
native range only in eastern
states. It is known to have been
introduced into New Jersey and
Tennessee as well as Alabama.
Sometimes they were moved to
other regions within the same
state to which they are native.
Such is the case in Virginia, for
example. The two mosquitofish
species are so similar in appear-
ance and so closely related, and
early records often so imprecise,
that it is difficult to always be
sure which species was trans-
planted where.
Description. These two little
fishes are very similar in appear-
ance. They have arched backs
and deep bellies. The head is
large, its upper surface flattened.
The eyes are very large relative
to body size. The small mouth
is upturned and protrusible to
allow feeding from the surface.
Dorsal and anal fins are rou-
nded; small black spots appear Top: The western mosquitofish is native to water bodies in the south
on the dorsal fin. The western central United States. Bottom: Gambusia affinis has been introduced into
mosquitofish usually has six dor- most western states. (Both maps adapted from Nico, Fuller, and Jacobs
sal rays, and the eastern mos- 2009.)
quitofish seven.
The head and much of the body are covered with large scales outlined by dark pigments
to give a characteristic diamond pattern on the body. The back is greenish or brown, the
sides gray-blue, and the belly silvery-white. In mature females, a black patch appears on
the rear of the abdomen above and forward of the vent. Anal, pelvic, and pectoral fins are
a pale, translucent amber.
Mosquitofish are sexually dimorphic; the females are much larger than the males. Adult
females may be as much as 2–2.75 in. (6–7 cm) long, adult males only about 1.5 in. (4 cm).
Adult males possess a gonopodium, a long tube-like structure composed of fused rays of the
anal fin and used to transfer sperm into the vent of females during mating. The gonopodium
is usually held back along the belly, but during mating, it is turned down and forward.
Related or Similar Species. The two mosquitofish are easily confused with each other. In
Arizona, the Sonoran topminnow (Poeciliopsis occidentalis) could be mistaken for a
180 n VERTEBRATES (FISH)

mosquitofish. The gonopodia of

the two are distinct, that of the
topminnow is asymmetrical to
the left, and that of the mos-
quitofish is symmetrical. In
addition, the pelvic fins of male
Gambusia are modified with a
fleshy appendage. In Florida,
the sailfin molly (Poecilia lati-
pinna) looks superficially like a
mosquitofish, but lacks the
net-like scale pattern, and the
position of the dorsal fin differs.
Introduction History. The
first known introduction of
mosquitofish in the United
States occurred in the early
1890s. The western mosquito-
fish was taken from Texas to
Hawai’i to test their effective-
ness in consuming mosquito
larvae in 1905. The same year,
eastern mosquitofish from
North Carolina were released
into New Jersey waters to con-
trol mosquitoes. For several
decades afterward, mosquito-
fish were routinely and widely
introduced by government
agencies; for example, the
former U.S. Public Health
Top: The eastern mosquitofish is native to drainages of the Atlantic Slope Service introduced it as a cost-
south of New Jersey and of the Gulf of Mexico. Bottom: Gambusia holbrooki effective way to combat malaria.
has probably been transplanted only into bodies of water in the eastern Gambusia continue to be pro-
United States. (Both maps adapted from Nico and Fuller 2009.) moted and intentionally stock-
ed by local, state, and federal
agencies for mosquito control and are viewed as attractive alternatives to the use of chemical
insecticides. In some areas, range expansions have occurred once introduced populations
became established.
Habitat. Mosquitofish can be found in slow-moving streams, ponds, and wetlands and in
all sorts of artificial habitats. They thrive in shallow, often stagnant, ponds and at the edges
of streams and lakes where vegetation is dense and the water shallow and warm. They will
also live in slightly brackish backwaters. Mosquitofish tend to be much more tolerant of pol-
luted water than many native fishes because they can survive at low oxygen levels. Generally
subtropical to warm-temperate in distribution, they do not tolerate extremely cold temper-
atures, which may limit their success in northern states.
Diet. Best described as omnivores, mosquitofish prefer taking zooplankters and small
aquatic invertebrates near the water surface. They also consume small fish, fish eggs, and
 MOSQUITOFISH n 181

the larvae of amphibians. When

preferred items are in short
supply, they consume diatoms
and other algae. Although their
name implies they are proficient
feeders of mosquito larvae, they
seem to be no more effective in
mosquito control than some
native fish. Indeed, in a labora-
tory experiment, mosquitofish The two species of mosquitofish are very similar in appearance. Their
small, protrusible mouths are upturned so they can feed at the surface
fared poorly on a diet of mos-
of the water. (Gualtiero Boffi/Shutterstock.)
quito larvae.
Life History. Mosquitofish are ovoviviparous, undergoing internal fertilization of the
eggs, which hatch in the mother’s body. Thus, they are livebearers and reproduce rapidly.
Females can be sexually mature at six weeks of age, males at four weeks. When mating,
the male swings its gonopodium forward and inserts it into the female’s vent to transfer
sperm. The female is able to store sperm for several months and fertilize several broods from
one copulation. Gestation lasts 3–4 weeks. When the young are born, they are little more
than 0.25 in. (6 mm) long. Females can produce 4–5 broods a year, with 1–300 young in
each. At birth, the sex ratio is 1:1, but in the adult population, there are fewer males than
females. The typical lifespan is 15 months.
Impacts. Outside their native range, mosquitofish are notoriously destructive, contribut-
ing to population declines among native fishes and amphibians. Adults are very aggressive
and attack other fish, shredding their fins and sometimes killing them. They compete with
or displace indigenous fishes. They also alter zooplankton communities, as well as insect
and crustacean communities, by their selective predation on members of each. In some
circumstances, they may actually benefit mosquitoes by eating predatory invertebrates.
Mosquitofish have had especially deleterious effects in western states, where they are
implicated in the elimination or decline of endemic and federally endangered and threat-
ened fish. They have displaced the plains topminnow (Fundulus sciadicus) from preferred
habitat in shallow clear streams. They have caused the threatened Railroad Valley springfish
(Crenichthys baileyi) to shift its habitat and decline in numbers in springs in Nevada. In
Arizona, they are responsible for the local extirpation of the endangered Sonoran top-
minnow (Poeciliopsis occidentalis). Populations of the Gila topminnow (P. o. occidentalis) only
survive where Gambusia is absent. Another subspecies, the Yaqui topminnow (P. o. sonrien-
sis), is threatened with a similar fate. Mosquitofish are also responsible for the demise of
the least chub (Itichthys phlegethontis) in parts of Utah. Desert pupfish (Cyprinodon spp.) pop-
ulations are threatened by the introduction of mosquitofish in Nevada springs. Mosquitofish
are at least partially responsible for population declines in the Chiricahuan leopard frog
(Rana chiricahuensis) in southeastern Arizona, in the California newt (Taricha torosa) in
California, and in native damselflies on O’ahu, Hawai’i.
Mosquitofish have posed problems for native species wherever in the world they have been
introduced. This fish is on the IUCN’s list of “100 of the World’s Worst Invasive Alien Species.”
Management. It may be possible to eradicate some populations of mosquitofish. More
likely to be successful are efforts to prevent the further spread of the species. Application
of rotenone to waters can be a means of eliminating mosquitofish from small areas of perma-
nent water. The poison deprives the fish of oxygen, so they come to the surface for air and
can be mechanically removed. Rotenone is a broad-spectrum piscicide, so nontarget species
182 n VERTEBRATES (FISH)

are also affected. Barrier construction is feasible in some areas to stop range expansion.
Regulations to halt the transport and stocking of all nonnative fishes need to be drafted or
enforced to prevent future introductions.

Selected References
Aarn, and Peter Unmack. “Gambusia control homepage.” 1998. http://www.gambusia.net/.
IUCN/SSC Invasive Species Specialist Group (ISSG). “Gambusia affinis (Fish).” ISSG Global Invasive
Species Database, 2006. http://www.issg.org/database/species/ecology.asp?fr=1&si=126.
Masterson, J. “Gambusia affinis. Mosquitofish.” Smithsonian Marine Station at Fort Pierce, 2008. http://
www.sms.si.edu/IRLSpec/Gambusia_affinis.htm.
Myers, G. S. “Gambusia, the fish destroyer,” Oecologia 141: 713–21, 1965. Appendix A-73. Available
online at http://wildlife.utah.gov/pdf/AIS_plans_2010/AIS_12oWesternMosquitofish-Jenny
-final.pdf.
NatureServe. “Gambusia affinis (Baird and Girard, 1853): Western Mosquitofish.” NatureServe
Explorer: An Online Encyclopedia of Life, Version 7.1, 2009. NatureServe, Arlington, VA. http://
natureserve.org/explorer.
Nico, Leo, and Pam Fuller. “Gambusia holbrooki.” USGS Nonindigenous Aquatic Species Database,
Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=849.
Nico, Leo, Pam Fuller, and Greg Jacobs. “Gambusia affinis.” USGS Nonindigenous Aquatic Species
Database, Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=846.

n Northern Snakehead
Scientific name: Channa argus
Synonym: Ophicephaslus argus
Family: Channidae (Snakeheads)
Native Range. China (Amur, Hwang, and Yangtze river systems), North Korea, and
Russia (Amur River drainage system)
Distribution in the United States. Established in Maryland (Lower and Middle Potomac
River and tributaries), New York (Meadow Lake, in Queens), Pennsylvania (Edgewood
Lake, Philadelphia), and Virginia (Fairfax County in tributaries of the Middle Potomac
River such as Dogue Creek, Kanes Creek, Little Hunting Creek, Massey Creek, Occoquan
River; also Mulligan Pond and in Pohick Bay, Fort Belvoir; Dyke Marsh, Alexandria to
Mason Neck National Wildlife Refuge; King George County in Upper Machodoc Creek). It
has been collected in California, Florida, Massachusetts, and North Carolina, but is not
established in these states. A 2008 capture in Arkansas suggests snakeheads may be estab-
lished there.
Description. The northern snakehead has an elongated body with a long dorsal fin run-
ning almost the full length of the back and a long anal fin as well. The pelvic fins are close
to the gills and set almost directly below the pectoral fins. The body is a golden tan with large
dark irregular blotches on the flanks. The head is flattened and bears large scales resembling
those of a snake. The eyes are set forward on the head, and nostrils are tubular. The mouth is
large; the lower jaw is toothed and protrudes beyond the upper jaw; the roof of the mouth
(i.e., palatine and prevomer bones) is also toothed. Some teeth look like the canines of mam-
mals. Total length can reach about 40 in. (102 cm); adults may weight up to 15 lbs. (6.8 kg).
Related or Similar Species. Native bowfin (Amia calva) and burbot (Lota lota), both of
which have long bodies and long dorsal fins, might be mistaken for snakeheads. Bowfin
 NORTHERN SNAKEHEAD n 183

are distinguished by a short

anal fin, pelvic fins set well
behind the gills and pectoral
fins, and a rounded tail. A black
spot appears at the base of the
tail in males and juveniles.
Barbots display a split dorsal fin
(a shorter segment lies in front
of a longer one); and a single bar-
bel or whisker hanging below
the lower jaw. Other species
superficially resembling snake-
heads include the American eel
(Anguilla rostrata) and the sea
lamprey (Petromyzon marinus).
Eels lack pelvic fins and their
dorsal and anal fins converge
with the caudal fin so that it
looks like they have a single con-
tinuous fin. Sea lampreys have a
round mouth and reddish eyes
(see Fish, Sea Lamprey).
Introduction History. The
snakehead is a popular fish in
Asian food markets. Most intro-
ductions were probably deliber-
ate efforts to raise the fish in
local waters. The northern
snakehead is not part of the
aquarium trade, although other
species in the family are. The
first report of the species in the Top: The northern snakehead is native to parts of China, North Korea,
United States was in 1977. and the Amur River system in Russia. Bottom: Northern snakehead
This represented a failed populations are established in Maryland and Virginia. The fish has been
attempt to establish a popula- collected in other states, but with the possible exception of Arkansas,
tion in Silverwood Lake, San does not seem to have established populations in those areas. (Adapted
Bernadino County, California. from Fuller and Benson 2009.)
In 2000, three specimens were
taken from the St. Johns River in Florida. Subsquent collections were made in Burnham
Harbor (Lake Michigan), Chicago, Illinois; two locations in Massachusetts (2001 and
2004); Lake Wylie and South Fork Catawba River, North Carolina; and FDR Pond,
Philadelphia, Pennsylvania. In June 2008, a snakehead was taken from the Schuylkill
River in Philadelphia. None of these captures have been interpreted as evidence of estab-
lished populations.
Discovery of snakeheads in a pond in Crofton, Maryland, in June 2002 created intense
media coverage and brought the problem of invasive fishes to the public’s attention. This
population was subsequently eradicated by state biologists, who applied the pesticide
rotenone to the water.
184 n VERTEBRATES (FISH)

In April 2004, several snake-

heads were collected from the
Potomac River in Maryland and
Virginia. The source of these
fish is unknown. Many others
have since been collected in
the Middle and Lower Potomac
River basin around Dogue
Northern snakehead. The large scales on the head give this fish a snake- and Little Hunting Creeks in
like appearance. (U.S. Geological Survey Archive, U.S. Geological Survey,
Virginia and from the Anacostia
Bugwood.org.)
River in Maryland.
Habitat. The northern snakehead is a purely freshwater fish; it prefers shallow ponds and
sluggish streams with well-vegetated or muddy bottoms. It has also been found in wetlands.
Snakeheads inhabit waters with temperatures ranging from 32°F to 86°F (0–30°C).
Snakeheads are air breathers and, if kept moist, can survive out of water for as long as four
days. This enables them to wriggle across land and disperse to other bodies of water. It also
allows them to live in oxygen-depleted water, since they can gulp air at the surface.
Diet. Snakeheads are top-level carnivores. Juveniles consume zooplankters, insect larvae,
small crustaceans, and the fry of other fish species. Adults prey primarily on other fish, but
will take crustaceans, frogs, small reptiles, and even small birds and mammals.
Life History. These fish reach sexual maturity at 2–3 years, when they are about 12–14
in. (30–36 cm) long. Spawning occurs 1–5 times a year. A breeding pair forms a nest by mak-
ing a circular clearing in aquatic vegetation, sometimes weaving the plants they removed
around the edge to protect a vertical column of water. The female releases 1,300 to 15,000
eggs near the center of the water column. The male wraps his body around hers as he releases
milt and fertilizes the eggs when she releases them. Each egg contains a droplet of oil that lets it
float. The snakehead pair, which remain together throughout the spawning season, guard the
eggs and then the fry when they hatch 28–48 hours later. The fry remain in the nest until they
are about 0.3 in. (8 mm) long, at which time the yolk sac is completely absorbed. Typically,
the young stay together in a pack guarded by an adult until they reach early juvenile stage; they
are then about 0.7 in. (18 mm) long and beginning to eat zooplankton.
Impacts. Unknown. As predators, they are expected to compete with native fishes for
food and habitat.
Management. It is difficult to impossible to eradicate a population from a river system.
Snakeheads can be removed from ponds by applying rotenone, but nontarget fish are killed
too. Preventing new introductions is the main management strategy. Since 2002, federal law
prohibits the introduction and interstate transport of live snakeheads or their eggs without a
permit in the United States, the Commonwealth of Puerto Rico, or any territory or posses-
sion of the United States.

Selected References
“Do You Know the Difference?” Virginia Department of Game and Inland Fisheries, 2004. http://
www.dgif.state.va.us/fishing/snakehead_comparisons_052004.pdf.
Fuller, P. F., and A. J. Benson. “Channa argus (Cantor 1842).” USGS Nonindigenous Aquatic Species
Database, Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=2265.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Channa argus (fish).” ISSG Global Invasive Species Database, 2005. http://www.issg.org/
database/species/ecology.asp?si=380&fr=1&sts.
 RAINBOW TROUT n 185

“Recognizing Northern Snake-

head.” U.S. Fish and Wildlife
Service, 2003. http://www.ans
taskforce.gov/Education/Snake
head_Final%20from%20RO.pdf.
U.S. Fish and Wildlife Service.
“Injurious Wildlife Species:
Snakeheads (Family Chanidae):
Final Rule.” Federal Register 67
(193), Rules and Regulations:
62193–62204, October 4, 2002.

n Rainbow Trout
Also known as: Steelhead
(anadromous form)
Scientific name: Oncorhynchus
mykiss
Synonyms: Salmo gairdnerii,
Fario gairdnereri
Family: Salmonidae
Native Range. North America.
Pacific slope streams from the
Kuskokwim River, Alaska, south
to Rio Santa Domingo, Baja
California, Mexico. Also in the
upper Mackenzie River drainage
system, Alberta and British
Columbia, Canada; and in in-
ternal drainages of southern
Oregon.
Distribution in the United Top: Rainbow trout are native to Pacific Slope streams in North America.
States. Rainbow trout has been Bottom: Rainbow trout have been transplanted into streams in all 50
stocked in all states. states and Puerto Rico. (Both maps adapted from Fuller 2009.)
Description. Rainbow trout
are deep-bodied, compressed
fishes with large heads; mouths extend to the back of the eyes. Coloration is highly variable.
(There are hundreds of varieties and several subspecies of native rainbow trout.) The back is
bluish to dark olive-green. Black spots appear on the back and may extend down the sides as
far as the lateral line. Radiating rows of black spots appear on the unforked tail. The sides are
silver, and the belly and underside of head are white. Often a pink or reddish band runs
along the sides of the body and head. Colors become much more intense when rainbows
are on spawning runs. Resident riverine populations normally exhibit the most intense pink
stripe and heaviest spotting. Rainbows can be positively identified by the 8–12 rays in the
anal fins and lack of teeth at the base of the tongue. A typical four-year-old river rainbow
weighs about 1 lb. (0.5 kg) and is about 13 in. (33 cm) long. Lake-dwelling rainbow trout
will be larger.
Related or Similar Species. Brown trout (see Fish, Brown Trout).
186 n VERTEBRATES (FISH)

Introduction History. Stock-

ing of this fish outside its native
range began in the late 1800s.
Introductions have been pri-
marily into streams for sport-
fishing. Around the world, rain-
bow trout are one of the most
widely introduced fish species
and important in aquaculture,
as they are highly valued food
fish.
Habitat. Rainbow trout is
usually a freshwater species,
although anadromous popula-
tions occur where rivers are
Coloration on rainbow trout is highly variable. Usually a reddish band open to the sea. Such fishes are
runs along the sides of the head and body. (Eric Engbretson.) called steelheads or coastal rain-
bows. Rainbows thrive in cool,
clean, well-oxygenated streams and lakes where temperatures range from 50°F (10°C) to
75°F (24°C). A water temperature around 54°F (12°C) is preferred.
Diet. Rainbow trout consume zooplankters, crustaceans, larval and adult insects both
aquatic and terrestrial, mollusks, fish eggs, and small fishes. Young trout feed on
zooplankters.
Life History. Rainbow trout usually migrate into small tributary streams of lakes to spawn
in the spring. The female uses her tail to excavate a redd or depression 4–12 in. deep and
10–15 in. (25–38 cm) in diameter in the streambed gravel into which she releases 700–
4,000 orange-red eggs. The male then fertilizes the eggs. The female covers successive
groups of fertilized eggs by digging at the upstream edge of the redd. When spawning is fin-
ished, the redd consists of layers of eggs and clean gravel, and the adults leave. Depending
on water temperature, eggs incubate for 1–5 months. Several weeks after hatching, the fry
wriggle through the gravel and swim free. They shelter and feed along the edges of streams
or near lake shores for 2–3 years. In native populations, sexual maturity is usually reached
at age 6–7.
Most introduced populations are bred in hatcheries and released into local streams just
ahead of the fishing season. Fishing pressures typically deplete the population, so they must
be restocked each year. They may also be restocked annually in areas where environmental
conditions are not conducive to self-sustaining populations.
Impacts. A major problem with the introduction of rainbow trout outside their native
range has been their ability to hybridize with other native salmonids. They thereby threaten
the genetic integrity of several rare species and subspecies vulnerable to extinction. In
California, they have crossed with Lahontan cutthroat trout (O. clarki henshawi), redband trout
(O. mykiss subspp.), and California golden trout (O. mykiss aguabonita), the state fish. In the
Lahontan drainage east of the Sierra Nevada and in several Rocky Mountain rivers, hybridiza-
tion with rainbow trout has been a major cause of the decline of native cutthroat trout. In
Nevada, it is implicated in the extinction of the Alvord cutthroat (O. clarki alvordensis), and
in Arizona, it hybridizes with native Gila trout (O. gilae) and Arizona trout (O. gilae apache).
Rainbow trout negatively affect other species through predation and aggressive behavior.
For example, rainbows push the Little Colorado spinedace (Lepidomeda vittata) from the
 ROUND GOBY n 187

undercut banks where it seeks shelter into open water, where it becomes much more vulner-
able to predation. They are also known to drive suckers and squawfish off their feeding ter-
ritories. In the Little Colorado River, rainbow trout prey upon endangered humpback chubs
(Gila cypha). They are at least partly responsible for declines in the Chiricahuan leopard
frog (Rana chiricahuensis) in southeastern Arizona and the Sierra Nevada yellow-legged frog
(R. sierrae). They also compete with native brook trout (Salvelinus fontinalis) in many locations.
Through the stocking of hatchery-bred rainbows in rivers in more than 20 states, whirling
disease has been introduced into open waters, where it threatens native species. Whirling
disease is caused by a parasite (Myxobolus cerebralis) of salmonids. Introduced from Eurasia,
the cnidarians-like organism infects juveniles and affects the nervous and skeletal systems,
resulting in a curvature of the spine that affects the victim’s ability to maintain its orientation
in the stream. Infected fish swim in spirals, chasing their tails. Mortality rates are very high.
Rainbow trout has been nominated as one of “100 of the World’s Worst Invasive Alien
Species” by the IUCN/SSC Invasive Species Speciality group.
Management. Most management practices favor rainbow trout since they are an impor-
tant element in sport fisheries across the country. Many continue to be raised in hatcheries
and used in put-and-take operations.

Selected References
Fuller, Pam. “Oncorhynchus mykiss.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL,
2009. http://nas.er.usgs.gov/queries/factsheet.asp?SpeciesID=910.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSC). “Oncorhynchus mykiss (fish).” ISSC Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?fr=1&si=103.
Root, Laurie. “Rainbow Trout (Oncorhynchus mykiss).” South Dakota Department of Game, Fish &
Parks, 1994 http://www3.northern.edu/natsource/FISH/Rainbo1.htm.
“Steelhead /Rainbow Trout” Alaska Department of Fish and Game, 2011. http://www.adfg.alaska.gov/
index.cfm?adfg=steelhead.main.
Talbot, Ret. “Native California Trout Species and Subspecies: At Risk Rainbow and Cutthroat Trout
Indigenous to California.” Suite 01.com. 2009. http://freshwater-fish.suite101.com/article.cfm/
native_california_trout_species_and_subspecies.

n Round Goby
Scientific name: Neogobius melanostomus
Synonym: Apollonia melanostomus
Family: Gobiidae
Native Range. Eurasia. Native to brackish waters of the Sea of Azov, Black Sea, and
Caspian Sea and their tributary streams, where it prefers brackish waters.
Distribution in the United States. Great Lakes and their tributary streams and connecting
canals.
Description. The round goby is a small, soft-bodied fish with eyes raised above its head as in
frogs. Its lips are thick. Body color is a mottled gray, brown, or greenish color, except in breed-
ing males, which are black. The front dorsal fin usually has a conspicuous black spot. Pelvic fins
are fused to form a suction disk. The front dorsal fin has 5–6 spines, and the posterior dorsal fin
has one spine and 13–16 soft rays. The anal fin has one spine and 11–14 soft rays. Adult gobies
are 4–10 in. (10–25 cm) long. In the Great Lakes, they rarely grow larger than 7 in. (18 cm).
188 n VERTEBRATES (FISH)

Related or Similar Species.

Round gobies are similar in
appearance to native sculpins
(Cottus spp.), which have two
separate pelvic fins instead of
the fused one of gobies. They
also look similar to tubenose
gobies (Proterorhinus marmora-
tus), another exotic fish in the
Great Lakes. The tubenose goby
has nostrils that extend forward
in tubes to or beyond the lower
lip. There is no spot on the
front dorsal fin, but oblique
black lines. It usually is only
about 4.5 in. (11 cm) long.
Smaller and less aggressive than
round gobies, the tubenose has
only been found in Lake St.
Clair, the St. Clair River, the
Detroit River, and western Lake
Ontario, although recently it
appears to be spreading in
Lake Erie.
Introduction History. The
round goby was first collected
in the United States in 1990
from Lake St. Clair. They most
likely were introduced during
ballast water exchange by trans-
Atlantic ships coming from
Top: The round goby is native to fresh and brackish waters of the Sea of the Black Sea region. By 1993,
Azov, Black Sea, and Caspian Sea, as well as their tributaries. (Adapted it had entered Lake Erie. In
from Stepien, Carol, Joshua Brow, and Emily Sopkovich. “Round Goby 1994, it appeared in Lake
Phylogeography,” Lake Erie Center, University of Toledo, 2009. http:// Michigan near Chicago. It has
www.utoledo.edu/as/lec/goby/RoundPhy.html.) Bottom: Round gobies since been collected near the
are invasive in all of the Great Lakes and many of their tributary terminus of the Chicago Sani-
streams. (Adapted from Fuller, Benson, and Maynard 2009.)
tary and Shipping Canal, entry-
way to the Mississippi River
drainage system. In 1995, gobies were collected from Wisconsin waters in Lake Superior
and, in 1996, from Duluth Harbor, Minnesota. The spread to Lake Superior was probably
aided by freighters operating on the lakes. By 1998, the round goby was reported from sev-
eral Michigan sites on Lake Huron. Gobies were reported in the Erie Canal, Buffalo River, St.
Lawrence River, Genessee River, Tonowanda Creek, and Lake Ontario in 2004 and 2005.
Today there are large populations in Lakes Erie, Ontario, and St. Clair. Lake Michigan expe-
rienced a population explosion of gobies in 2008.
Habitat. Freshwater lakes, streams, and canals. These bottom-dwelling fish prefer hard
substrates such as rock, gravel, or sand and areas shallow enough for aquatic macrophytes
 ROUND GOBY n 189

to grow. They hide in crevices.

They tolerate a wide range of
temperatures and are able to
inhabit brackish water and
degraded water quality.
Diet. Round gobies are car-
nivores and feed on zoo-
plankters, mollusks, aquatic
insects and benthic inverte-
brates, fish eggs, and small fish.
They feed voraciously on zebra
mussels (Dreissena polymorpha),
another invasive species in the
Great Lakes (see Mollusks,
Zebra Mussel). Gobies are able
to pluck mussels from their The round goby’s pelvic fins are fused to form a suction disk. In their
attachment sites on the lake introduced range, they are rarely more than 7 in. long. (Eric Engbretson.)
floor and crush them with their
pharyngeal teeth. They spit out the shells before swallowing the soft insides.
Gobies have well-developed lateral lines, which increase their ability to detect water
movement and allow them to be nocturnal feeders.
Life History. Gobies spawn from April to September. Males migrate from wintering sites
in deeper water to spawning areas ahead of the females. They establish territories and attract
mates by releasing pheromones. Females attach 500–3,000 eggs to the underside of rocks, in
pipes, or on other sheltered hard surfaces. They utilize several nests and spawn with several
different males. Males aggressively guard the nests and newly hatched young. Gobies will
spawn every 20 days during the long season. Females are sexually mature at 1–2 years;
males at 3–4 years. Males die after spawning, so population sex ratios are usually skewed
to favor females.
Impacts. Round gobies compete with native fishes for food and space and could poten-
tially have serious impacts on aquatic ecosystems if they were to gain entry to the
Mississippi drainage system. Their ability to feed at night and their ability to cling with their
pelvic fins to rocks in fast currents give them a competitive advantage over native species,
many of which have overlapping food preferences. They compete for small macroinverte-
brates with rainbow darters (Etheostoma caeruleum), logperch (Percina caprodes), and
northern madtoms (Noturus stigmosus). Since gobies were introduced to Lake St. Clair, pop-
ulations of native sculpin and logperch have declined. They are known to eat the eggs and
fry of lake trout and the eggs of lake sturgeon, and to prey on small darters in laboratory
situations.
Since adult round gobies vigorously defend their spawning territories, they become dom-
inant in prime spawning areas and displace natives. Mottled sculpin (Cottus bairdi) is one
species that has been affected in that way by the establishment of round gobies. Large,
mature gobies drive them from spawning sites; and small gobies, focusing on insects, com-
pete with them for food.
Indirectly, gobies have had an impact on smallmouth bass (Micropterus dolomieu) in Lake
Erie. Male bass guard nests and can successfully ward off goby attacks. However, if the males
are removed through fishing, gobies immediately move in and gorge themselves on bass
eggs. In order to preserve the smallmouth bass population, the State of Ohio has had to close
190 n VERTEBRATES (FISH)

the fishery in May and June, while bass are spawning. This has led to a significant economic
loss as anglers traditionally took half the bass catch during those months. Gobies also affect
recreational fisherman by snatching the bait from their hooks.
On the positive side, the diet of large adult round gobies overwhelmingly consists of
zebra mussels. No other fish in the Great Lakes exploits this resource to such an extent.
However, predation of zebra mussels by gobies is not sufficient to stem the spread of those
aquatic invaders. And because zebra mussels are filter-feeders and accumulate contaminants
in their body tissues, fish that eat gobies will biomagnify the concentrations, potentially ren-
dering some sportfishes unfit for human consumption.
Some native fishes, including smallmouth bass and walleye, consume gobies. It is hoped
that Atlantic salmon (Salmo salar), introduced into the Great Lakes in 1966 to feed on ale-
wives (see Fish, Alewife), may switch to gobies as alewife populations decline in Lake
Michigan, threatening the salmon fishery.
Management. Controlling or reducing the goby populations of the Great Lakes is impos-
sible. Preventing the inland spread of the invader is the key goal of management. Electric bar-
riers on tributary streams are under consideration. Good fishing and boating practices need to
be applied and enforced. Dumping of ballast water in waterways needs to be regulated. Live
bait should not be discarded into water. Boats, trailers, and fishing gear should be thoroughly
inspected upon leaving any body of water and all visible organic debris removed. All water
should be drained from the boat before leaving the shoreline. All boats should be washed
and then sun-dried for five days before being put into another body of water.

Selected References
Crosier, Dani, and Dan Malloy. “Round goby (Neogobius melanostomus),” United States Federal Aquatic
Nuisance Species Task Force, 2005. http://www.anstaskforce.gov/spoc/round_goby.php.
Fuller, P., A. Benson, and E. Maynard. “Neogobius melanostomus.” USGS Nonindigenous Aquatic
Species Database, Gainesville, FL, 2011. http://nas.er.usgs.gov/queries/FactSheet.asp
?speciesID=713.
Hayes, R. “Neogobius melanostomus.” Animal Diversity Web, University of Michigan Museum of
Zoology, 2008. http://animaldiversity.ummz.umich.edu/site/accounts/information/Neogobius
_melanostomus.
Morlock, Jerry W. “Invasive Species Round Goby Has Population Explosion in Lake Michigan.”
Michigan Live LLC, 2009. http://www.mlive.com/news/muskegon/index.ssf/2009/01/invasive
_species_round_goby_ha.html.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Neogobius melanostomus (Fish).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?si=657&fr=1&sts.
“Round Goby.” Great Lakes Science Center, USGS, 2008. http://www.glsc.usgs.gov/main.php
?content=research_invasive_goby&title=Invasive%20Fish0&menu=research_invasive_fish.

n Sea Lamprey
Also known as: Lamprey eel, lake lamprey
Scientific name: Petromyzon marinus
Family: Petromyzontidae
Native Range. The Atlantic coast of North America from Labrador to Florida and into the
Gulf of Mexico. This is primarily a marine fish, but it ascends freshwater rivers to spawn. It is
native to Atlantic Slope drainages, the St. Lawrence River, and probably Lake Ontario, and it is
 SEA LAMPREY n 191

probably also native in the Finger

Lakes of New York, and Lake
Champlain, New York/Vermont.
It also occurs naturally along the
Atlantic coast of Europe and in
the western Mediterranean Sea.
Distribution in the United
States. Sea lampreys are nonna-
tive transplants established and
invasive in the four upper
Great Lakes and in cold, clear
streams in the region. Cur-
rently, populations are espe-
cially high in the St. Mary’s
River, which connects Lake
Huron and Lake Michigan.
Description. The sea lamprey
is a thin, eel-like fish without
jaws. A member of a primitive
group of agnathans, its skeleton
is made of cartilage. The body is
smooth and scaleless. It has two
close dorsal fins situated toward
the rear of the body, but no
paired fins, no lateral line, and
no swim bladder. There are seven
gill slits. The back is gray-blue,
with a metallic violet shimmer
on the often mottled sides. The
belly is yellow to silver-white.
The seven gill openings are lined
up behind the eyes, which are Top: The native range of the catadromous sea lamprey encompasses the
red. The mouth is a disk-like Atlantic coasts of North America and Europe. It is probably also native
sucker containing whorls of 100 to Lake Ontario, the Finger Lakes of New York, and Lake Champlain.
or more sharp, curved teeth Bottom: The sea lamprey is invasive in Lakes Superior, Huron,
made of keratin and a toothed Michigan, and Erie and in their clear, cold tributary streams (Both maps
adapted from Fuller, Nico, and Maynard, 2009.)
“tongue.” Total length of adults
ranges between 12 and 20 in.
(30.5–50.8 cm); landlocked fish are usually smaller than native marine-dwelling individuals,
which may attain lengths of 24–30 in. (61–76 cm). Adults weigh 8–13 oz. (0. 23 kg).
Related or Similar Species. There are four other native species of lamprey in the United
States: the silver lamprey (Ichthyomyzon unicuspis), chestnut lamprey (Ichthyomyzon casta-
neus), northern brook lamprey (Ichthyomyzon fossor), and American brook lamprey
(Lampetra appendix). The latter two, unlike the sea lamprey, are not parasitic. All are benign.
Each has a characteristic structure to its mouth and arrangement of the teeth, which differ
from that of the sea lamprey and are diagnostic.
American eels (Anguilla rostrata), true eels, might be mistaken for sea lampreys, but they
have jaws and a sharp, pointed head.
192 n VERTEBRATES (FISH)

A. The sea lamprey is a jawless fish that resembles an eel. (Andrei Nekrassov/Shutterstock.) B. The mouth of the
sea lamprey is a sucker containing whorls of sharp teeth and a toothed “tongue.” (U.S. Environmental Protection
Agency, Great Lakes National Program office.)

Introduction History. Sea lampreys were first reported in Lake Ontario in 1835. It was
not collected from Lake Erie until 1921. It was found in Lake Michigan in 1936, Lake
Huron in 1937, and Lake Superior in 1946. Increasingly, genetic evidence points to the
sea lamprey being native to Lake Ontario and, along with populations in Lake Champlain
and the Finger Lakes, a relict of the last Pleistocene glaciation. Niagara Falls presented a
barrier to dispersal from Lake Ontario into Lake Erie until the Welland Canal was opened
in 1829 to bypass the falls. Still, it seems lampreys were not able to overcome the barrier
until improvements were made on the canal and lock system in 1919. Within two years, it
was in Lake Erie and within 25 years, it had spread to all the other Great Lakes.
Habitat. The larvae inhabit soft sediments at the bottoms of clear streams. Adults are
parasitic and attach to larger fish in temperate cool-water lakes. They tolerate temperatures
ranging from 40°F (5°C) to 68°F (20°C).
Diet. Larvae are filter-feeders, consuming invertebrates and detritus from the water.
Juveniles are external parasites on healthy fish. A sea lamprey fastens onto its victim and uses
its tongue to rasp a hole in the host fish’s body. It then sucks out the blood, body fluids, and
flesh, keeping the wound open for hours or even weeks by means of an anticoagulant in its
saliva. One lamprey is estimated to eat 40 lbs. of fish in its lifetime. Mature adults do not feed.
Life History. In their native habitat, sea lampreys spawn in freshwater streams where the
substrate is stony. They move upriver in late May and early June, by which time the adults
are no longer feeding. The male and female excavate nests, or redds, about 6 in. (15 cm)
deep and 2–3 ft. (60–90 cm) in diameter, removing stones with their sucker mouths, and
piling them downstream. Both attach themselves to a large stone at the upstream edge of
the nest. (This behavior is reflected in their Latin name: petromyzon means “stone sucker.”)
With the male wrapped around the female, the pair stirs up sand and deposits eggs and milt.
The sand grains stick to the fertilized eggs, burying them; the larvae develop in the stream
bottom. Adults die after spawning.
 SEA LAMPREY n 193

The larval (ammocoete) stage lasts 6–8 years. When 4–5 in. long, the larvae undergo an
extreme metamorphosis and transform into adults. It is then that they migrate downstream
into estuaries and the sea, where they remain as parasitic juveniles for 1–2.5 years.
Populations introduced into the Great Lakes or that may be landlocked, ice age relicts in
the Finger Lakes and Lake Champlain, of course, never go to sea; instead, juveniles migrate
from the streams where they hatched and spend their long larval period in freshwater lakes,
where they attach themselves to large lake fish, including lake trout (Salvelinus namaycush),
yellow perch (Perca flavescens), burbot (Lota lota), walleye (Stizostedion vitreum), channel cat-
fish (Ictalurus punctatus), and northern pike (Esox lucius). At the end of the juvenile period,
they become sexually mature adults, cease feeding, and swim upriver to spawn.
Impacts. The introduced sea lamprey is considered to have been a major factor in the col-
lapses of lake trout, whitefish (Coregonus clupeaformis), and chub (deepwater ciscos) fisheries
in the Great Lakes (especially in Lakes Huron and Michigan and eastern Lake Superior)
during the 1940s and 1950s. In its parasitic phase, its attacks and feeding on other fish
result in high mortality among host species. It kills directly by sucking out fluids and tissues,
and it kills indirectly by leaving open wounds vulnerable to infection. Circular scars on sur-
viving host fish attest to many of them having been attacked more than once. The species’
introduction to the Great Lakes coincided with major declines in several large native fish
that had been important in commercial fisheries and sport fisheries. Combined with the
effects of pollution and overfishing, the influx of sea lampreys led to the extinction of three
endemic fish: the longjaw cisco (Coregonus alpenae), deepwater cisco (C. johannae), and
blackfin cisco (C. nigripinnis). Since lamprey had so reduced populations of predatory fish,
populations of the invasive alewife (Alosa pseudoharengus) exploded and affected the species
composition of the native fish fauna (see also Fish, Alewife). Although sea lamprey popula-
tions have been reduced in most parts of the Great Lakes, they are still abundant enough to
hamper restoration efforts directed at native game fish and the introduced Atlantic salmon.
Management. The Great Lakes Fishery Commission, U.S. Fish and Wildlife Service, and
Fisheries and Oceans Canada are all involved in controlling sea lampreys in the Great Lakes.
Mechanical weirs and electrical barrier were the first methods used to control the upstream
migration of spawning sea lampreys. The release of sterile males also was implemented to
reduce successful reproduction. In the late 1950s, an effective lampricide, TFM (trifluoro-
methyl-4-nitrophenol), was developed. This chemical kills lamprey larvae in the stream bot-
tom. It was successful in reducing populations by over 90 percent and allowing the recovery
of some commercial fisheries. TFM must be used repeatedly to remain effective, and some-
times it is harmful to nontarget species such as walleye and native lampreys. Recently, a
controlled-release version of the lampricide Bayluscide was developed at the Upper
Midwest Environmental Sciences Center (USGS); it is supposed to kill sea lamprey larvae
in bottom sediments without affecting other species.

Selected References
Fuller, Pam, Leo Nico, and Erynn Maynard. “Petromyzon marinus.” USGS Nonindigenous Aquatic
Species Database, Gainesville, FL, 2009. http://nas.er.usgs.gov/queries/FactSheet.asp
?SpeciesID=836.
“Sea Lamprey.” Upper Midwest Environmental Sciences Center, USGS, 2007. http://www.umesc
.usgs.gov/invasive_species/sea_lamprey.html.
Summers, Adam P. “Sea Lamprey, Petromyzon marinus.” Department of Biology, University of
Massachuseatts, Amherst, 1997. http://www.bio.umass.edu/biology/conn.river/sealampr.html.
194 n VERTEBRATES (FISH)

n Silver Carp
Also known as: Flying carp
Scientific name: Hypophthalmichthys molitrix
Family: Cyprinidae
Native Range. Large Pacific drainages of eastern Asia. Silver carp are native to major low-
land rivers from the Amur River of far-eastern Russia south to the Pearl River of China, and
possibly into northern Vietnam.
Distribution in the United States. Established in the Mississippi River drainage system in
Illinois and Louisiana. Silver carp have been reported in Alabama, Arizona, Arkansas,
Colorado, Florida, Indiana, Kansas, Kentucky, Mississippi, Missouri, Nebraska, Ohio,
South Dakota, and Tennessee, but these may not be reproducing populations.
Description. The silver carp
is a large, deep-bodied fish with
a laterally compressed body. A
long ventral keel extends for-
ward from the vent almost to
the junction of the gill mem-
branes and is diagnostic. The
lateral line is complete and
bends down toward the belly.
Large eyes are set low on the
head. The mouth is large, tooth-
less, and downturned. There
are no barbels. Gill rakers are
thin, branched, and fused into
a sponge-like structure. The
pharyngeal teeth are in one
row, four on a side, at the back
of the throat and have striated
surfaces.
The short dorsal fin has a
moderately stiff, non-serrated
spine at its origin; it begins
behind the origin of the pelvic
fins. The anal fin is hooked
and has a slightly stiffened
spine at the origin. The caudal
fin is forked. Pectoral fins have
15–18 rays and a stiff, hard
spine with a finely serrated rear
margin. Small specimens lack
spines on their fins.
Scales are very small on the
Top: Silver carp are native to the large lowland rivers of East Asia from the body, and the head and oper-
Amur River south to the Pearl River. Bottom: Silver carp are invasive in cules are scaleless. Juveniles are
the Mississippi River system and have been reported elsewhere silvery all over. The backs and
(Adapted from Nico 2009.)
 SILVER CARP n 195

upper sides of adults turn

greenish, and gradually become
silvery below the lateral line.
Silver carp grow to more
than 3 ft. (1 m) long and may
weigh 60 lbs. (27 kg) or more.
Related or Similar Species.
Silver carp are most similar to In silver carp, a long keel on the belly is diagnostic, as is the long lateral
another Asian carp, the bighead line that bends toward the belly. (Michigan Sea Grant archives.)
carp (Hypophthalmichthys nobilis).
The bighead has a shorter ventral keel and irregular dark blotches over its body (see Fish,
Bighead Carp, for a more detailed description). The presence of a ventral keel separates both
silver and bighead carp from all native cyprinids except the golden shiner (Notemigonis cry-
soleucas), which has larger scales and five pharyngeal teeth on each side of its jaw rather than
the four found in Asian carps. Native shad could be mistaken for small juvenile silver carp,
but shad have no lateral line and fewer than 14 rays in the anal fin.
Introduction History. Silver carp were first imported into the United States in 1973 by a
private fish farmer in Arkansas for the purpose of controlling phytoplankton in his fish
ponds. It was soon being raised at six state, federal, and private hatcheries and, by the
1970s, stocked in several municipal sewage lagoons. It was first discovered in natural waters
in 1980, presumably the result of escapes from hatcheries and other facilities. Silver carp in
the Ouachita River in Louisiana likely stem from escapes from an aquaculture site upstream
in Arkansas. Florida received silver carp via a contaminated stocking of grass carp legally
introduced to control aquatic plants. In Arizona, silver carp also came in a contaminated
shipment of grass carp, but this was an illegal stocking. Ohio River fish may be products
of the stocking of nearby ponds or from populations originating in Arkansas.
Habitat. Silver carp is a freshwater fish that inhabits large rivers, lakes, and ponds. It
needs moving water for spawning and proper development of eggs. Flooded lowlands make
good nursery areas for larvae and juveniles.
Diet. Adult silver carp filters phytoplankters, bacteria, and detritus from the water with its
specialized gills. It reportedly also grazes aquatic plants. Juveniles feed on zooplankters. It feeds
by rapidly gulping water as it swims, then closing its mouth and pumping the water out through
its opercula. Those food particles taken in are ground by the pharyngeal teeth against a cartilagi-
nous plate.
Life History. Silver carp are sexually mature at three years of age. In their native range,
they migrate to communal spawning areas during the spring highwater period. They spawn
in small groups of 15–25 fish at dusk and at dawn, when the water temperature is between
65° and 68°F (18–20°C). They require water moving enough to aerate the eggs and let them
float downstream in the current.
Impacts. Silver carp is believed to have the potential to greatly alter native aquatic ecosys-
tems because of its consumption of plankton, a food required by larval fish and native mus-
sels. It could also compete with adults of some native fishes such as gizzard shad and
bigmouth buffalo (Ictiobus cyprinella) and introduce diseases.
A greater impact occurs on fishermen and recreational boaters. Silver carp swim near the
surface and are easily disturbed by boat motors. When startled, they leap high out of
the water, the reason they are sometimes called flying carp. Being struck by a 40–60 lb.
(18–27 kg) fish hurtling through the air can cause serious injury.
196 n VERTEBRATES (FISH)

Management. The U.S. Fish and Wildlife Service lists silver carp as an injurious species;
and under the Lacey Act, it is illegal to import it into the United States. The fear that this fish
and its close relative the bighead carp could devastate Great Lakes fisheries has led the U.S.
Army Corp of Engineers and the State of Illinois to construct electric barriers in the Chicago
Sanitary and Ship Canal to halt its dispersal into the lakes from the Mississippi River (see
also Fish, Bighead Carp).

Selected References
“Aquatic Invasive Species. Silver Carp.” Indiana Department of Natural Resources, 2005. http://
www.in.gov/dnr/files/SILVER_CARP.pdf.
Nico, Leo. “Hypophthalmichthys molitrix.” USGS Nonindigenous Aquatic Species Database, Gainesville,
FL, 2011. Revised January 11, 2011. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=549.
“Silver carp. Hypophthalmichthys molitrix.” Florida Integrated Science Center, USGS, 2005. http://
fl.biology.usgs.gov/Carp_ID/html/hypophthalmichthys_molitrix.html.

n Spotted Tilapia
Also known as: Black mangrove cichlid
Scientific name: Tilapia mariae
Synonyms: Tilapia dubia, T. meeki
Family: Cichlidae
Native Range. West Africa, in coastal rainforest streams and brackish lagoons from the
Tabou River in Côte d’Ivoire to the Pra River in Ghana and from southeast Benin to the
Kribi River in Cameroon.
Distribution in the United States. Established in at least eight counties of southern
Florida and in a few springs in Nevada. They are present in the Salton Sea, Colorado
River, and Los Angeles area of California, and possibly established in Arizona.
Description. Spotted tilapia have compressed oval bodies. The two dorsal fins appear as a
single fin; the forward one has 16 sharp spines and the rear one 12–13 soft rays. The tail is
fan-shaped. The anal fin has three spines at the front and 10–11 rays that taper to a point at
the rear. The mouth is terminal, and the large eye reddish.
Coloration is distinct. Juveniles bear no spots on the body but have a disruptive pattern of
black bars on a yellow-green or gold background. There is a prominent black spot on the
back of the dorsal fin, however. The bars fade with age, and 6–9 irregular square dark
blotches appear along the midline of the flanks of adults. Some show pink or red coloration
on chin or throat when spawning.
Some sexual dimorphism is present among spotted tilapia. Males have somewhat longer
dorsal and caudal fins, both of which display shimmering white spots not found in females.
Also, the foreheads of males have a steeper rise.
Spotted tilapia are smaller than most of the many other Africa cichlids established in
Florida and other Gulf states, and it is the most abundant. Florida specimens typically are
between 6 and 8 in. (15–20 cm), although they may grow to 12 in. (30 cm) or more and
weigh up to 3 lbs. (1.4 kg). Most fish larger than 10 in. (25 cm) are males.
Related or Similar Species. Spotted tilapia resemble some native sunfishes in body and
mouth shape. Occasionally, they are mistakenly called “oscars.” They are very similar to
the introduced redbelly tilapia (T. zilli), but that species has been eradicated in Florida.
 SPOTTED TILAPIA n 197

Introduction History. The

first known occurrence of spot-
ted tilapia in the United States
dates to April 1974, when it
was discovered in Snapper
Creek Canal in South Miami,
Miami-Dade County. Florida.
It quickly became established
in canals throughout eastern
Miami-Dade and southeastern
Broward County. It was estab-
lished in Everglades National
Park and Big Cypress National
Preserve by the late 1980s.
These fish originated as escapes
or intentional releases from
aquarium fish producers in
Dade County sometime bet-
ween 1972 and 1974.
Similarly in Nevada, intro-
duction was related to an aquar-
ium release. Spotted tilapia
have been abundant in Rogers
Spring, a thermal spring in the
Lake Mead National Recreation
Area, Clark County, since 1980.
Habitat. Spotted tilapia pre-
fer warm, slow-moving waters.
They thrive in canals in South
Florida and in warm springs in
Nevada. Adults and juveniles
stay close to sheltering struc- Top: Spotted tilapia are native to rainforest streams and brackish lagoons
tures such as vegetated shores in West Africa. (Adapted from Robins, n.d.) Bottom: Spotted tilapia are
or rock outcrops. They need invasive in Florida and also established in Arizona. They are present in
hard, flat surfaces when spawn- the Colorado River and Salton Sea. (Adapted from Nico 2009.)
ing. Optimal water tempera-
tures are 77–91°F (25–33°C); temperatures below 52°F (11°C) are not tolerated.
Diet. These fish feed primarily on “aufwuchs,” the attached algae, detritus, and associated
small invertebrates that grow on various plant and rock surfaces. They will also filter phyto-
plankton from eutrophic (plankton-rich) waters.
Life History. Spotted tilapia form pair bonds well before spawning begins and stay
together to care for eggs and fry. Typically, breeding colonies assemble on the spawning
grounds. In Florida, spawning generally occurs between November and March and may be
synchronized with the lunar cycle, more batches of eggs being laid just before the full moon.
These fish are substrate spawners and prefer gravel nests under flat rocks. Each female pro-
duces 200–2,000 turquoise eggs that stick to the gravel bottom. Two days after the eggs are
laid, she removes all fertile eggs to a nearby pit and eats all infertile eggs. Early-stage larvae
have “head glands” that produce a sticky strand by which the young attach themselves to
198 n VERTEBRATES (FISH)

A. Adult spotted tilapia. (© Noel Burkhead.) B. Juvenile spotted tilapia. (© Noel Burkhead.)

the substrate so that they will not drift away. Both parents guard and feed the hatchlings
until they are free-swimming some nine days later, when they are about 1 in. (2.5 cm) long
and have developed the cryptic barring typical of juveniles. Spotted tilapia are sexually
mature at 7 in. (18 cm).
Impacts. Spotted tilapia disperse rapidly and quickly become the dominant fish in both
numbers and biomass in habitats to which they are introduced, thereby reducing biodiver-
sity. By virtue of their numbers, they may compete with native fish for food. They may also
compete with native fish such as sunfishes for spawning areas, since they aggressively
defend their nests and broods.
Management. In the 1980s, Florida attempted to control spotted tilapia by introducing
yet another exotic fish, the predatory South America peacock cichlid (Cichla ocellaris) to feed
upon it. How successful this effort was is open to question; the two species seem to have
reached an equilibrium in their predator-prey relationship.
Prevention of further spread of tilapia is the main goal of management. It is illegal to
transport or possess live tilapia in Florida. This fish is edible, so fishermen wishing to eat
tilapia should immediately kill them and put them on ice.

Selected References
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Tilapia mariae (Fish).” ISSG Global Invasive Species Database, 2009. http://www.issg.org/
database/species/ecology.asp?si=1430&fr=1&sts=&lang=EN.
Nico, Leo. Tilapia mariae. USGS Nonindigenous Aquatic Species Database, Gainesville, FL, 2006.
http://nas.er.usgs.gov/queries/factsheet.asp?SpeciesID=482.
Robins, Robert H. “Spotted Tilapia.” Florida Museum of Natural History, n.d. http://www.flmnh
.ufl.edu/fish/gallery/Descript/SpottedTilapia/SpottedTilapia.html.

n Walking Catfish
Also known as: Magur
Scientific name: Clarias batrachus
Family: Clariidae
Native Range. Southeast Asia. Native to Bangladesh, eastern India, Indonesia, Malaysia,
Myanmar (Burma), Pakistan, Singapore, Sri Lanka, and Thailand. The fish introduced to
Florida were from Thailand.
 WALKING CATFISH n 199

Distribution in the United

States. Established throughout
most of Florida, including
Everglades National Park, Big
Cypress National Preserve,
Florida Panther National Wild-
life Refuge, and Pelican Island
National Wildlife Refuge. Re-
ported from California, Con-
necticut, Georgia, Massachusetts,
and Nevada.
Description. The walking
catfish has a flat, broad head
and an elongated body tapering
toward the tail. It has a typical
catfish appearance, with four
pairs of barbels (whiskers) and
wide, fleshy lips. The teeth are
small and bristle-like. The eyes
are small. The dorsal fin is long
and continuous and extends
two-thirds the length of the
back. There is no adipose fin as
in native catfishes. Anal fins are
also long, but end in a lobe sep-
arate from the caudal fin (tail),
which is rounded. Each pec-
toral fin has a strong, rigid spine
at the front that helps the fish
“walk.” Somewhat ungainly ter-
restrial locomotion is achieved
by using the pectoral spines to Top: Walking catfish are native to parts of South and Southeast Asia.
pull itself along while flexing (Adapted from Robins n.d.) Bottom: Walking catfish are invasive in
the body back and forth. Florida. They have been collected in other states. (Adapted from Nico
The scaleless body is usually 2009.)
a drab gray or gray-brown with
white flecks on the sides. Albino and calico colors are common in aquarium fishes, but wild
populations revert to the natural gray. (All of the original walking catfish introduced into
Florida were albinos.)
The gills of walking catfish are specially structured with tree-like organs to permit their
breathing on land and in stagnant water.
Related or Similar Species. Native catfish, such as the marine hardhead catfish (Ariopsis
felis) and gafftopsail catfish (Bagre marinus), as well as freshwater catfish such as the brown
bullhead (Ictalurus nebulosus) and channel catfish (I. punctatus) could be mistaken for walk-
ing catfish. All native catfishes, however, possess a forked tail, an adipose fin just forward of
the tail, and a dorsal spine.
Introduction History. Walking catfish were first imported to Florida in the early 1960s
for the aquarium trade. The introduction into local waters occurred in the mid-1960s when
200 n VERTEBRATES (FISH)

adult fish escaped either from a fish farm in northeastern Broward County or from a trans-
port truck carrying brood fish between Broward and Miami-Dade counties. Apparently pur-
poseful releases were made by fish farmers elsewhere in Florida in 1967–1968 after the state
prohibited the importation and possession of walking catfish.
By 1968, walking catfish were established in three counties; 10 years after the first
escapes, they were in almost 20 counties in the southern half of Florida. Their spread was
facilitated by the network of canals and drainage ditches in southeastern Florida and their
ability to cross land, especially on rainy nights.
Aquarium releases are the likely source of introductions in other states.
Habitat. Walking catfish may inhabit a variety of habitats, but they are most commonly
associated with freshwater habitats in which most native species do not thrive. They are
bottom-dwellers that become the dominants in warm, muddy ponds, swamps, canals, and
ditches and tolerate stagnant, low-oxygen conditions. Since they can “walk” from pond to
pond and also burrow into pond and river banks for a period of dormancy, they can also
inhabit intermittent streams and isolated temporary ponds. They are essentially tropical fish
and vulnerable to cold. Winter kills have been reported in Florida when water temperatures
drop below 50°F (10°C).
Diet. Walking catfish feed on large aquatic invertebrates, the eggs and larvae of amphib-
ians and fish, and small fish. They are known to have invaded aquaculture farms to feed on
fish.
Life History. Walking catfish become sexually mature at about one year of age. Mass
migrations and spawning seem to be linked to the rainy season. Nests are hollowed out in
detritus or submerged vegetation and guarded by the adults. The up to 1,000 eggs laid are
adhesive and stick to the substrate. Once the eggs have been fertilized, the male guards the
nest, and the female hovers nearby to drive off intruders. Embryos hatch in about 30 hours
and the fry stay under the protection of the parents for about five days, at which time the
yolk-sac is absorbed and the young begin to forage. Catfish grow rapidly and, in Florida,
usually reach a length of about 12 in. (30 cm). The largest reported walking catfish in the
United States was about 20 in. (50 cm) long and weighed about 3 lbs. (1.4 kg).
Impacts. When walking catfish were first released into Florida waters, they were forecast
to become the most harmful introduction known so far. They were prolific, and because
they could move across land, they dispersed rapidly. It was expected that they would out-
compete native catfish species and cause the decline of some native amphibians. However,
after an initial population explosion, numbers began to decline in the 1970s. Actual ecologi-
cal impacts are unknown. They have not eliminated any native fish and indeed do not seem
to have had any major negative effect on native fish or amphibian communities. It may
be that their preference for ponds that partially dry up each year and stagnant bodies of
water to which no natives are adapted has isolated them from native fish populations and
reduced their effects. They do, however, have an economic impact on fish farmers, who
must construct barriers to keep them out of their aquaculture ponds, and are still considered
undesirable. The walking catfish has been nominated as among 100 of the “World’s Worst”
invaders by the Invasive Species Specialist Group (ISSG).
Management. All members of the walking catfish family are on the U.S. Fish and Wildlife
Service’s list of Injurious Wildlife Species. Under the Lacey Act, it is a violation of federal law
to import them into the United States without a permit. Some states have laws that make it
illegal to possess a live walking catfish. Any walking catfish caught by anglers must be
quickly killed. Fish fences keep walking catfish out of aquaculture ponds.
 AFRICAN CLAWED FROG n 201

Selected References
Masterson, J. “Clarias batrachus (Walking catfish).” Smithsonian Marine Station at Fort Pierce, 2007.
http://www.sms.si.edu/IRLspec/Clarias_batrachus.htm.
Nico, Leo. “Clarias batrachus.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL, 2009.
Revised April 11, 2006. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=486.
Robins, Robert H. “Biological Profiles: Walking Catfish.” Florida Museum of Natural History, n.d.
http://www.flmnh.ufl.edu/fish/Gallery/Descript/WalkingCatfish/WalkingCatfish.html.
“Walking Catfish—Clarias batrachus.” MyFWS.com, Florida Fish and Wildlife Conservation
Commission, Tallahassee, n.d. http://myfwc.com/wildlifehabitats/profiles/fish/nonnative-fish/walking
-catfish/.

n Amphibians
n African Clawed Frog
Also known as: Upland frog, common platanna
Scientific name: Xenopus laevis
Family: Pipidae
Native Range. Highlands of sub-Saharan Africa in Angola, Botswana, Burundi,
Cameroon, Central African Republic, Democratic Republic of the Congo, southwestern
Kenya, Lestho, Malawai, Mozambique, Namibia, eastern Nigeria, Rwanda, South Africa,
Swaziland, Uganda, Zambia, and Zimbabwe. Absent from Congo Basin and warmer low-
lands of East Africa. Imports to the United States originated in South Africa.
Distribution in the United States. Disjunct populations are established in at least seven
counties in southern California, and reported from others. A single population is established
in artificial ponds at a golf course in Tucson, Arizona. African clawed frogs have been col-
lected in Colorado, Florida, Massachusetts, North Carolina, Nevada, New Mexico, Texas,
Utah, Virginia, Wisconsin, and Wyoming, but no established populations are known from
these states.
Description. The African clawed frog has a flattened body and relatively small head. It
lacks a visible external ear drum (tympanum) and has no tongue. Its eyes are small and lid-
less, located near the top of the head. The forefeet have four thin, unwebbed fingers that
usually point forward. The large hind feet have five fully webbed toes, the inner three of
which possess sharp, black claws. The skin is very smooth and slippery. The back is a mot-
tled olive-brown or gray, and the belly is cream colored. The lateral lines seem stitched along
the back. Snout to vent length is from 2.0 to 5.5 in. (5–14 cm.)
Tadpoles of this species are unique. They are transparent, so the internal organs are vis-
ible. Barbels occur at the mouth, making them resemble small catfish. The tail ends in a fil-
ament. They usually swim head down, vibrating the tail filament to stir up plankton.
Tadpoles grow to about 1.5 in. (3.8 cm) long.
Related or Similar Species. None. No native American frog or toad has clawed hind limb
toes. No native amphibians have transparent tadpoles or tadpoles with long barbells.
Introduction History. The African clawed frog was introduced to laboratories around the
globe in the 1940s and 1950s after it was discovered it could be used as a reliable test for
human pregnancy. An injection of urine from a pregnant woman stimulated a female frog
202 n VERTEBRATES (AMPHIBIANS)

to lay eggs. As the demand grew,

the frog was bred in captivity, so
that most free-living popula-
tions outside the natural range
of the species are actually feral
stock. The clawed frogs are easy
to care for and disease resistant,
and in the 1950s and 1960s,
they gained a foothold in the
pet trade. It is still an important
research animal, used primarily
for studies of developmental,
cell, and molecular biology.
Most introductions to the
wild were intentional releases
from laboratories once other
techniques to determine hu-
man pregnancy were devised.
Pets, no longer wanted by their
owners, were also rele-
ased. The first feral African
clawed frog was found in
Orange County, California, in
1968. Independent introduc-
tion events occurred in five
counties (Los Angeles, Oran-
ge, Riverside, San Diego, and
Santa Barbara); in 25–30 years,
the frogs spread throughout
most of the drainage systems
into which they were first re-
Top: The African clawed frog is native to the uplands of sub-Saharan leased. San Bernadino and
Africa. (Adapted from range map of Xenopus laevis at http:// Ventura counties were invaded
www.iucnredlist.org/apps/redlist/details/58174/0/rangemap.) Bottom: from neighboring counties. It is
Self-sustaining populations of African clawed frog occur in California expected that they will continue
and Arizona. It has been collected in several other states, where, to date, to spread along the many irri-
no populations have become established. (Adapted from Somma 2005.)
gation canals and drainage
ditches in southern California.
In Arizona, African clawed frogs were deliberately introduced to man-made ponds in
Arthur Park Golf Course, in Tucson, Pima County, in the late 1960s, and they persist there.
Spread to other bodies of water has been inhibited by the surrounding expanse of desert.
Other states where the frog was introduced but never became established, with the
recorded dates of introduction, are: Colorado, 1990; Florida, 1964; Massachusetts, 1993;
North Carolina, 1990s; Virginia, 1980s (eradicated 1987–1988); Wisconsin, 1972.
Habitat. The African clawed frog spends almost its entire life in aquatic habitats, leaving
water only if forced to migrate when a water body dries up or is poisoned. They may be
found in all types of aquatic situations except fast-moving rivers and water bodies with
predatory fish. They tend to prefer eutrophic ponds, both natural and artificial, slow
 AFRICAN CLAWED FROG n 203

streams, and natural waterways

that are frequently disturbed or
experience high environmental
variability. They are salt tolerant
and can live in estuaries.
Although they find optimal cli-
matic conditions in Medi-
terranean climates and water
temperatures between 60° and
80°F (15–27°C), adults can tol-
erate temperatures ranging
from 32° to 86°F (0–30°C)
and populations have persisted
under winter ice. Tadpoles sur-
vive in water temperatures
ranging from 50° to 86°F (10–
30° C). Adults can aestivate for African clawed frogs have flattened bodies and lack visible external ear
periods of up to eight months drums. (Todd Battey.)
and go without food for a year.
Eggs and tadpoles are viable in both acidic and alkaline water (pH 5–9).
Diet. Adults feed on slow-moving aquatic invertebrates that they suck into their mouths.
They will also scavenge dead and dying arthropods and take small fish and tadpoles, includ-
ing those of their own species. They rely on the lateral line system to detect scents in the
water and the movement of prey. Forelimbs gather food into the mouth, while the strong,
clawed hind legs shred larger prey items into ingestible sizes.
Tadpoles are filter-feeders. They extract small phytoplankters from open water. Food
includes one-celled algae such as diatoms, protozoans, and bacteria.
Life History. African clawed frogs in California breed from January through November.
A peak occurs in April and May. Males vocalize underwater during the evening to call
females. The mating call alternates long and short trills. The female either accepts him with
a rapping sound or rejects him with a slow ticking sound. (Very seldom does this answering
behavior happen with any other animal.) Mating usually takes place at night. Unlike most
other frogs, the mating embrace or amplexus is pelvic. Over the course of the next 3–4 hours,
the female releases, one at a time, hundreds of eggs that, fertilized by the male, adhere to
plants or other underwater structures. The eggs grow into tadpoles and hatch in 3–4 days.
The length of the larval stage is 5–12 weeks. Tadpoles are weak swimmers prone to preda-
tion by fish; they congregate in schools in deeper water to feed. As the tadpole metamorpho-
ses, the tail is absorbed, providing nutrition for the developing froglet. Metamorphosis takes
4–5 days. Sexual maturity is reached 6–10 months after metamorphosis. African clawed
frogs may live 9–15 years.
Impacts. The African clawed frog may be a threat to native amphibians and fish in
California and the Southwest since it preys upon tadpoles and fish fry. Its range overlaps
with such vulnerable species as western toad (Bufo boreas), red-spotted toads (Bufo puncta-
tus), California red-legged frogs (Rana draytoni), Pacific treefrogs (Pseudacris regilla), and
western spadefoot toads (Spea hammondi). It is known to eat larvae and recently metamor-
phosed western toads, but negative impacts on California red-legged frogs, though claimed,
have not been verified. Observations of African clawed frogs preying on fish are rare, but do
include consumption of the federally endangered tidewater goby (Eucyclogobius newberryi),
204 n VERTEBRATES (AMPHIBIANS)

African Clawed Frog as Pharmacopoiea

T he African clawed frog has long been used as a research animal because it is easy
to care for and breeds rapidly, and the development of its transparent embryos
is easy to observe. In 1987, Dr. Michael Zasloff discovered that the skin of the
African clawed frog has glands that secrete a broad-spectrum antimicrobial chemi-
cal effective against viruses, bacteria, fungi, and protozoa. When first discovered,
this defensive mechanism represented a totally new way that vertebrates were able
to protect themselves against infection. Previously, it was believed only the immune
system attacked invading pathogens. The frog skin chemicals work by creating a
hole in the target’s cell membrane, causing the cell contents to leak out.
Dr. Zasloff isolated two peptides from the skin of the African clawed frog and
named them magainins, from the Hebrew word magain, meaning shield.
Magainins were synthesized and manipulated in the laboratory to create thou-
sands of related antibiotics. There was much hoopla in the 1990s about their
potential use in new drugs to combat a variety of human infections. Experiments
showed that magainins killed bacteria such as E. coli, staphylococci, streptococci,
and enterobacteria. They gave promise in combating colon cancer. A topical cream
proved effective as a treatment for diabetic foot ulcers. Magainins in toothpaste
might fight dental plaque. They might control the tuberculosis bacillus or the plas-
modium that causes malaria, or they could work as a spermicide that not only pro-
tects against pregnancy but also against sexually transmitted diseases. Although
no magainin-containing drugs have completed the whole arduous course through
animal trials and human safety tests to FDA approval and the drug store shelf, their
discovery in the African clawed frog opened new paths of research in animal-
derived antibiotics and pointed to future discoveries of antibiotics and other help-
ful compounds in what remains of Earth’s biodiversity.
Since 1987 almost all frog species tested have been found to produce related
chemicals in their skin and in their digestive tracts; and other animals have found
to produce similar compounds in their saliva.
Source: Altman, Lawrence K. “Curiosity on Healing in Frogs Leads to a Gain in Antibiotics.”
New York Times, July 31, 1987. http://www.nytimes.com/1987/07/31/us/curiosity-on-healing-
in-frogs-leads-to-a-gain-in-antibiotics.html?ref=lawrencekaltman

arroyo chub (Gila orcutti), and another federally endangered fish, the unarmored threespine
stickleback (Gastaerosteus aculeatus williamsoni). Were African clawed frogs to get into the
isolated pools containing desert pupfish (Cyprinodon spp.), those rare fish would likely be
at severe risk.
Many scientists believe African clawed frogs carried the fungus Batrachochytrium dedroba-
tidis (see Fungi, Chytrid Frog Fungus) around the globe. This fungus is a major cause of the
worldwide decline and extinction of frogs and toads. African clawed frogs are immune to the
fungus, which is native to their African habitats. Museum specimens dating to 1938 show
evidence of chytridiomycosis, the skin disease associated with it.
Management. Most efforts to eliminate clawed frogs in California have not been success-
ful. Permanent eradication is known in only one instance, when an artificial pond on the
University of California, Davis, campus was poisoned by the California Department of Fish
 AMERICAN BULLFROG n 205

and Game. Elsewhere, poisoning has not worked. At Vasquez Rock in the upper Santa Clara
River drainage, the frogs simply left the water after rotenone was applied. Other methods of
control include draining ponds, removing frogs by seine or traps, electroshocking, and
introducing predatory fish; but none have proved effective in eliminating frogs or preventing
their reintroduction.

Selected References
Crayon, John J. “Xenopus laevis (Daudin, 1802) African Clawed Frog.” AmphibiaWeb: Information on
amphibian biology and conservation. Berkeley, California, 2009. http://amphibiaweb.org/.
Garvey, Nathan. “Xenopus laevis African Clawed Frog.” University of Michigan Museum of Zoology,
2000. http://animaldiversity.ummz.umich.edu/site/accounts/information/Xenopus_laevis.html.
Measey, John. “Xenopus laevis (Amphibian).” IUCN/SSC Invasive Species Specialist Group (ISSG)
Global Invasive Species Database, 2006. http://www.issg.org/database/species/ecology.asp?si=150.
Rorabaugh, Jim.“African Clawed Frog Xenopus laevis.” Online Field Guide to the Reptiles and
Amphibians of Arizona, 2008. http://www.reptilesofaz.org/Turtle-Amphibs-Subpages/
h-x-laevis.html.
Somma, Louis A. “Xenopus laevis.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL,
2009. Revised March 24, 2005. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=67.
Willigan, Erin. “African Clawed Frog (Xenopus laevis).” Introduced Species Summary Project, Columbia
University, 2001. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/
xenopus_laevis.htm.

n American Bullfrog
Scientific name: Lithobates catesbeianus
Synonym: Rana catesbeiana
Family: Ranidae
Native Range. Central and eastern United States and southern parts of Ontario and
Quebec provinces, Canada.
Distribution in the United States. The American bullfrog has become established outside
its native range in Arizona, California, Colorado, Hawai’i, Nebraska, Nevada, Oregon, Utah,
and Washington. It has also been introduced to locations where it was not historically found
within its natural range in Massachusetts (on the islands of Nantucket and Martha’s Vineyard,
and in the Wellfleet Bay sanctuary on Cape Cod), Iowa (in the DeSoto National Wildlife
Refuge on the Missouri River), and New Jersey (in the Cape May National Wildlife Refuge).
Description. Adult bullfrogs are green to brownish green, with dark blotches and bars on
the back and legs. The upper jaw is often a light green. The belly is cream to yellow. During
the breeding season, the throat of the male is yellow, while that of the female is white. The
absence of dorsolateral folds is diagnostic. However, there is a short fold in the skin running
from the eye, over the eardrum to the forelimb. The eardrum or tympanum is conspicuous
and exhibits sexual dimorphism. In males, the tympanum is larger than the eye; in females,
it is the same size as or smaller than the eye. The hindfoot is fully webbed with the exception
of the last joint of the fourth toe.
The tadpole is large (up to 6 in. or 15 cm long). Its back is yellow green and has black
spots; the belly is lighter. The dorsal fin is arched.
The call of the bullfrog is deep and very loud; it is usually described as “jug-o’rum” or
“br-wum.” They also sound a loud squeak of alarm before jumping into the water when
206 n VERTEBRATES (AMPHIBIANS)

surprised. Bullfrogs are active

both day and night and are
highly aquatic, rarely found
out of the water.
The bullfrog is the largest
frog native to North America.
From snout to vent, the length
of adults ranges from 3.5 to 8
in. (9–20 cm), the largest indi-
viduals being female. They may
weigh more than 1 lb. (0.5 kg).
They are celebrated jumpers,
and their long legs are consid-
ered a delicacy; they are often
hunted and farmed for meat.
Related or Similar Species.
The green frog (Lithobates
[=Rana] clamitans) is much
smaller (adults only 2–4 in. or
5–10 cm, snout to vent) and has
the prominent dorsolateral folds
characteristic of most North
American frogs. Its distribution
is similar to that of the natural
range of the American bullfrog.
Introduction History. Bull-
frogs were first introduced to
California in 1890s and Colo-
rado in the early 1900s. By the
end of the 1920s, they were
common on the lower Colorado
Top: The American bullfrog is native to the central and eastern United River near Yuma, Arizona. The
States and adjacent areas of Canada. Bottom: American bullfrogs have pathways of introduction are
been transplanted beyond their native range into all western states largely unknown. In Colorado,
(except Alaska), Hawai’i, and Puerto Rico. (Both maps adapted from bullfrog tadpoles at fish hatch-
McKercher 2009.) eries may have contaminated
fish stockings in trout streams
and lakes. Other possible means of introduction include the aquarium trade, frog-farming
operations, and the stocking of farm ponds to control agricultural pests. They may have been
deliberately released to provide a new game animal or introduced as a living feature of orna-
mental ponds. Once in the wild, bullfrogs are able to disperse through entire drainage systems.
In some western states, the competitive advantage that the American bullfrog has over
native frogs is enhanced by the introduction of nonnative fishes, such as bluegill sunfish
(Lepomis macrochirus). In the Pacific Northwest, bluegills and other sport fish were stocked
in lakes that had no native fishes. The nonnative fish eat the tadpoles of native frogs, but
leave bullfrog tadpoles alone. Bullfrogs coevolved with these alien fishes and are unpalatable
to them. Furthermore, the sunfish consume the nymphs of native dragonflies, species that
do prey upon bullfrog tadpoles.
 AMERICAN BULLFROG n 207

A. Adult American bullfrogs have no folds on the back or sides, but a short fold of skin does occur from the eye
over the conspicuous external eardrum to the forelimb. (Ilias Strachnis/Shutterstock.) B. The tadpole is very large
and has an arched dorsal fin. (Tommounsay/iStockPhoto.)

Bullfrogs have been introduced to Mexico, the Caribbean, South America, Europe, and
Asia.
Habitat. Shallow, still waters are preferred, especially where abundant aquatic vegetation
provides shelter. Bullfrogs are thus found in marshes and along the shores of lakes, ponds,
slow-moving rivers, and reservoirs or other impoundments. They need warm water for
breeding, and tadpoles prefer non-vegetated areas. Tadpoles are able to overwinter in water
beneath a cover of ice. Adults hibernate in mud during cold weather.
Diet. Adult bullfrogs are carnivorous and tend to eat whatever they can catch with their
large sticky tongues, including insects, crayfish, fish eggs, eggs of amphibians, tadpoles,
and frogs, including other bullfrogs. They also take snakes, birds, bats, and mice.
Tadpoles are mostly herbivorous and feed on cyanobacteria, algae, other plant material,
and small aquatic invertebrates.
Life History. American bullfrogs have a long breeding season, from May to July in
northern states and from February to October in warmer areas. A female deposits up to
20,000 eggs in a broad foamy sheet in warm, quiet water. Fertilization is external. The raft
of jelly-coated eggs may stretch 3 ft. (1 m) in diameter. It floats until just before hatching,
when it sinks onto underwater vegetation. Tadpoles hatch out 3–5 days after fertilization.
The gilled tadpoles develop slowly, and those in colder climes overwinter at least once.
The transformation from tadpole to froglet, when the tail is slowly absorbed, may take
1–3 years. The new froglets measure about 2 in. (5 cm) long. Bullfrogs become sexually
mature two years later, and live 7–9 years in the wild.
Impacts. The introduction of the American bullfrog to western states has likely exacer-
bated problems related to the demise of native frogs because bullfrogs feed on frogs and tad-
poles. Tadpoles may significantly alter aquatic community structure because they feed
heavily on algae. In Arizona, predation by bullfrogs has contributed to the extirpation of
populations of native leopard frogs and garter snakes. Most vulnerable are the Mexican
garter snake (Thamnophis eques megalops), and the federally threatened Chiricahua leopard
frog (Rana chiricahuensis). Introduced bullfrogs may also spread chytridomycosis, a fungal
disease of the skin implicated in the decline of amphibians worldwide (see Fungi, Chytrid
Frog Fungus). ISSG has nominated them as one of 100 of the “World’s Worst” invaders.
Management. Bullfrog populations are often managed as fisheries, and harvesting limits
population growth. Adults can be taken with gigs, nets, traps, bow and arrow, and guns;
208 n VERTEBRATES (AMPHIBIANS)

and they can be caught by hand. Spotlighting at night is a common practice since frogs are
immobilized by bright lights. Larvae can be killed by the same poisons used against fish.
Suctioning off the egg masses can be tricky but effective.
New introductions to areas where bullfrogs are not yet present should be prohibited.

Selected References
“Alien Species in Cahoots.” U.S. Geological Survey News Release, March 13, 2003. http://fresc.usgs.gov/
news/newsreleases.asp?NRID=3.
Bruening, S. “Rana catesbeiana.” Animal Diversity Web, University of Michigan Museum of Zoology,
2002. http://animaldiversity.ummz.umich.edu/site/accounts/information/Rana_catesbeiana.html.
Crayon, John J. “Rana catesbeiana (Amphibian).” Global Invasive Species Database, IUCN/SSC Invasive
Species Specialist Group, 2005. http://www.issg.org/database/species/ecology.asp?si=80&fr
=1&sts.
McKercher, Liz, and Denise R. Gregoire. Lithobates [=Rana] catesbeianus. USGS Nonindigenous Aquatic
Species Database, Gainesville, FL, 2009. Revised March 7, 2011. http://nas.er.usgs.gov/queries/
FactSheet.asp?speciesID=71.
Murphy, Martin. “North American Bullfrog (Rana castesbeiana).” Introduced Species Summary Project,
Columbia University, 2003. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/
inv_spp_summ/Rana_catesbeiana.htm.
Rorabaugh, Jim. “American Bullfrog Rana catesbeiana.” Online Field Guide to the Reptiles and
Amphibians of Arizona, 2008. http://www.reptilesofaz.org/Turtle-Amphibs-Subpages/
h-l-catesbeianus.html.

n Coqui
Also known as: common coqui, coquı́ común, Puerto Rican treefrog
Scientific name: Eleutherodactylus coqui
Family: Leptodactylidae
Native Range. Puerto Rico. Native to a variety of habitats and elevations.
Distribution in the United States. Coqui are established in Hawai’i on Hawai’i Island,
Kaua’i, Maui, and O’ahu. They are established but noninvasive in Florida, where they tend
to die off during winter freezes. Established populations also occur on St. Croix, St. John,
and St. Thomas, U.S. Virgin Islands.
Description. A small but very loud frog. Coqui are quite variable in color. The back may
range from light yellow to dark brown and may or may not be mottled with black spots.
There may be a cream line running along the midline of the back from the snout to the hind
legs or an indistinct dark “M” between the shoulders. The underside is light with brown stip-
ples. Eyes are gold or golden brown. The toes have suction cup-like pads like those of true
treefrogs (family Hylidae). Breeding males are slightly larger than 1 in. (34 mm) from snout
to vent; females average about 1.5 in. (41 mm). It appears that the average size of coqui frogs
is increasing in Hawai’i, since frogs continue to grow throughout their lives and may live up
to seven years. Coqui have been on the Big Island for 10 years. They also tend to be larger at
higher elevations. There is no tadpole stage.
The mating call of the male coqui is a rapid and loud “ko-KEE.” Males begin to call at
dusk and may continue all night, especially if it is raining. According to one report, the
sound measures as much as 80–90 decibels, comparable to a lawn mower.
Related or Similar Species. In Hawai’i, the coqui most resembles another exotic amphib-
ian, its close relative the greenhouse frog (Eleutherodactylus planirostris). Greenhouse frogs
 COQUI n 209

have narrower snouts, a nar-

rower body shape, claw-like
toes, red eyes, and a warty-
textured skin. Adults are usu-
ally a mottled copper color and
are less than 1 in. (2.5 cm) long.
Most importantly, they have a
soft bird- or cricket-like chirp.
They live on the ground and
are not considered invasive.
In Florida, coqui might be
confused with native treefrogs.
Introduction History. The
coqui was probably introduced
to the island of Hawai’i acci-
dentally around 1988 in a ship-
ment of horticultural plants
such as bromeliads from
Puerto Rico. From a few in-
fected nurseries, the coqui was
likely spread in landscaping
materials to other sites and
other islands. Populations grew
rapidly and expanded beyond
the sites of initial entry due of
lack of predators (e.g., owls,
snakes, tarantulas) on the
islands. By 2003, there were
more than 200 infestation sites
on the Big Island, 40 or more
on Maui, 5 on O’ahu, and 1 site
on Kaua’i. These sites include Top: The coqui is native to Puerto Rico. Bottom: The loud little arboreal
commercial plant nurseries, frog is invasive in Hawai’i. The expansion of populations in Florida
the grounds of resort hotels, seems to be prevented by cold winter weather. (Adapted from Somma
ornamental plantings in parks, 2009.)
and in native forest.
Habitat. Coqui frogs are found in Hawai’i in a variety of ecological zones at elevations
ranging from sea level to 4,000 ft. (1,220 m). Most populations are in or near horticultural
sites, although they spread from nurseries, resorts, and residential areas into natural areas
on public lands. By day, they hide in shady, thick moist brush, leaf litter, dead leaves on
banana plants, empty flower pots, or holes in porous rock. They nest in cavities, folded or
rolled leaves, PVC pipe, or other sheltering structures. Coqui seem to especially like broad-
leaf plants such as heliconias (Heliconia spp.), Koster’s curse (Clidemia hirta), Loulu palms
(Pritchardia spp.), silktree (Albezia julibrissin), split-leaf philodendrons (Monstera spp.), and
ti (Cordyline terminalis). Adults prefer to be 3–9 ft. (1–3 m) above the ground; juveniles are
usually found less than 4 ft. (1 m) above the ground. Since they have no tadpole stage, they
do not require standing water for breeding.
Diet. Insects and other terrestrial invertebrates.
210 n VERTEBRATES (AMPHIBIANS)

Life History. Breeding occurs

all year, but tends to concentrate
during the rainy season. Coqui
use internal fertilization. The
female lays a clutch of about
two dozen eggs in a protected,
elevated cavity. Males guard the
eggs and keep them from drying
out. Four to six clutches may be
produced in a year, usually one
every two months. The fertilized
eggs undergo direct develop-
ment to tiny (approx. 0.5 in. or
1.25 cm long) froglets in
2–3 weeks. Coqui froglets are
sexually mature in about eight
months.
This coqui is resting in a tank bromeliad. (Rogeliao Doratt/USDA.) Impacts. Hawai’i has no
native frogs, so any introduced
frog poses the potential to disrupt native ecosystems. Experiments have shown that con-
sumption of leaf-eating insects by coqui can increase foliage production and decomposition
rates, thereby increasing nutrient cycling rates. It is possible that coqui will consume
endemic insects and spiders, some of which are already on the verge of extinction and some
of which are important pollinators of Hawai’i’s endemic flora. It could compete with native
Hawaiian birds, most of which are insectivorous. There is also concern that coqui might pro-
vide a food source for the nonindigenous brown tree snake (Boiga irregularis), should it
invade the islands.
By far the greatest problem with the coqui is noise. Tourists and residents complain of
sleepless nights. Property values have declined in infested areas. The tourist and real estate
industries are threatened. Controls to prevent the further spread to the coqui could be del-
eterious to the floriculture and nursery industries, the main pathways of dispersal for this
tiny invader. The combined value of the tourism, floriculture, and real estate industries
which are at risk in Hawai’i is many millions of dollars.
For its potential ecological and economic impacts, the coqui has been nominated as
among 100 of the “World’s Worst” invaders by IUCN/Species Survival Commission’s
Invasive Species Specialist Group (ISSG).
Management. Small populations may be controlled by hand-capture and trapping.
Larger infestations may be managed by spraying landscape plants with hot water (113°F)
or dipping potted plants in hot water. A solution of citric acid will kill adults and eggs and
is approved for use by the EPA. Prevention is key. Hawaiians need to eliminate the habitats
that coqui prefer around their homes and vacation spots. They are encouraged to compost
yard wastes, prune and thin shrubbery, remove dead leaves especially from large-leaved
plants, and remove empty pots and other containers that collect rainwater during dry
seasons.
The inter-island transport of coqui frogs is now prohibited in Hawai’i. If strict controls
including certification of frog-free flowers and plants were to be placed on floriculture
exports—a not unreasonable prospect, this would create a financial hardship on one of
Hawai’i’s major industries.
 CUBAN TREEFROG n 211

Selected References
Chun, S., A. H. Hara, and R. Y. Niino-DuPonte. “Greenhouse Frog or Coqui Frog?” College of Tropical
Agriculture and Human Resources, University of Hawai’i at Manoa, 2003. http://www2.ctahr
.hawaii.edu/oc/freepubs/pdf/coqui_id.pdf.
“Control of Coqui Frogs in Hawai’i.” Department of Plant and Environmental Protection Sciences,
College of Tropical Agriculture and Human Resources, University of Hawai’i at Manoa, 2008.
http://www.ctahr.hawaii.edu/coqui/.
Kraus, Fred, Earl W. Campbell, Allen Allison, and Thane Pratt. “Eleutherodactylus Frog Introductions to
Hawaii.” Herpetological Review 30 (10): 21–25, 1999. Available online at http://www.hear.org/
articles/pdfs/herp_review_frogs_1999v30n1.pdf.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Eleutherodactylus coqui (Amphibian).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?fr=1&si=105.
Somma, Louis A. “Eleutherodactylus coqui.” USGS Nonindigenous Aquatic Species Database,
Gainesville, FL, 2009. Revised May 14, 2006. http://nas.er.usgs.gov/queries/FactSheet.asp
?speciesID=60.

n Cuban Treefrog
Also known as: Giant tree frog, rana platernera
Scientific name: Osteopilus septentrionalis
Synonyms: Trachycephalus insulsus, Hyla septentrionalis
Family: Hylidae
Native Range. Cuba, Isla de Pinos, the Bahamas, and the Cayman Islands.
Distribution in the United States. Established in Florida. Also established in Puerto Rico
and St. Croix and St. Thomas, U.S. Virgin Islands; collected from Georgia, but apparently
not established there. Single waifs have been reported from Colorado, Indiana, Maryland,
and Virginia. A report of a population on O’ahu, Hawai’i, has not been verified.
Description. The Cuban treefrog is the largest treefrog in the United States. It has very
large toe pads equal in size to the eardrum (tympanum). The very large eyes of the Cuban
treefrog give them a “bug-eyed” look. The back is warty, like a toad, and the belly is granular.
A skin fold extends from the eye rearward to the tympanum. Adults can be distinguished
from all native treefrogs because the skin on the top of the head is fused to the skull.
Color varies from gray to gray-green or tan-brown, or even cream, and individuals are able
to change color rapidly. The body is usually heavily mottled and there are often stripes on
the backside of the legs. A yellowish tinge occurs where the legs join the body. The rear toes
are slightly webbed, and there is a distinct tarsal fold along the entire ankle (tarsus). Most
adults in Florida are from 1 to 4 in. (2.5–10 cm) long, although adult females may grow to
6 in. (15 cm) or more.
Juveniles are difficult to identify because they lack warts and strong patterning.
Sometimes they lack the lateral stripe that is found on many native treefrogs. Tadpoles have
round bodies with dark or black backs. The intestinal coil is visible through the light-
colored belly. The tail is pigmented with lighter areas near the point of attachment to the
body. Scattered dark flecks appear on the wide fin.
The mating call of the male treefrog is a rasping snarl of varied pitch. It has been
described as sounding like a squeaky door. The male’s vocal sac looks like a double bubble.
Males also will call during the day when it rains.
212 n VERTEBRATES (AMPHIBIANS)

Cuban treefrogs secrete a

sticky substance that can irritate
the skin and mucus membranes
of people. The burning and itch-
ing sensation that results can
last for an hour or more. Wash
your hands after handling one.
Related or Similar Species.
In Florida, two native species
could be mistaken for Cuban
treefrogs. The green treefrog
(Hyla cinera) has a smooth,
bright-green skin. It has a point-
ed snout and a clear white line
along each side of the body.
The breeding call consists of a
bell-like ringing, repeated many
times. Maximum length is about
2.5 in. (6 cm). Squirrel treefrogs
(Hyla squirella) are much small-
er, adults being only about
1.5 in. (4 cm) long. They, too,
can change their color, but are
usually are muddy green on the
back; belly and feet are light
and spotted. Like all indigenous
treefrogs, they have a single,
rounded, balloon-like vocal sac.
The breeding call sounds like a
duck, while the rain call sounds
like a chattering squirrel.
Top: The native range of the Cuban treefrog consists of Cuba, Isla de Introduction History. Cuban
Pinos, the Bahamas, and the Cayman Islands. Bottom: Cuban treefrogs treefrogs were in Key West,
are established in Florida and Puerto Rico and possibly on O’ahu in Florida, sometime before 1928.
Hawai’i. Individual treefrogs have been collected in a number of other Most likely they arrived as acci-
states. (Adapted from Somma 2009.) dental hitchhikers in shipping
crates from the Caribbean. By
the mid-1950s, they had spread through the keys to Key Largo. The first specimen from main-
land Florida was collected in 1951. By the 1970s, they were being found throughout southern
Florida, and in 2007, breeding populations occurred as far north as Cedar Key on the Gulf
side of the peninsula and Jacksonville on the Atlantic side. Individuals have been reported
in Georgia and South Carolina. Cuban treefrogs disperse as stowaways on horticultural plants
(especially palm trees), building materials, cars, trucks, and boats. They respond to tropical
storms and hurricanes by immediately breeding and dispersing. It is expected that they will
continue to spread along the Gulf coast and then move southward into Mexico. Northward
expansion into the United States seems to be checked by cooler climatic conditions.
Habitat. Cuban treefrogs thrive in both natural and man-made habitats. In Florida, they
can be found in swamps and on hardwood hammocks as well as in pine forests. In
 CUBAN TREEFROG n 213

A. The skin on the head of the Cuban treefrog is fused to its skull. (Mike Pingleton, University of Illinois at
Urbana-Champaign, Bugwood.org.) B. The very large toe pads are characteristic of this treefrog, which often
invades homes. (Wayne Smaridge.)

residential areas, they are most frequently seen near ornamental ponds and near outside
lighting, where they wait on walls, windows, porches, and potted plants to catch insects.
When they get into homes, they are often encountered in the toilet. They also inhabit orange
groves and plant nurseries. Active at night, by day they hide in tight, moist areas such as cel-
lars, cisterns, drains, and plants.
Diet. These frogs are ambush predators that feed mainly on insects, spiders, and several
of Florida’s native treefrogs. They are also cannibalistic. Toads, lizards, and small snakes
are additional items reportedly consumed by Cuban treefrogs.
Tadpoles are omnivores and may consume the eggs and tadpoles of native frogs and even
those of their own kind.
Life History. The breeding season in Florida extends from May to October. Breeding is
stimulated by warm summer rains. Breeding events usually last only one night. Eggs are
deposited as a thin floating sheet in warm quiet waters of just about any size that lack preda-
tory fish. Their gelatinous films of eggs can be found in ponds, ditches, abandoned swim-
ming pools, and the stagnant waters of discarded containers. There is no parental care.
Tadpoles hatch about two days later. The new hatchlings have a body length of about 0.05
in. (1.2 mm) and a tail about 0.15 in. (3.8 mm) long. The tadpoles metamorphose into small
froglets in 3–8 weeks; the actual rate is determined by water temperature. At metamorpho-
sis, they are about 1 in. (26–28 mm) long. Males mature when they are about 1.5 in.
(40 mm) long; females mature more slowly and therefore are much larger than males at
maturity. Mature males participate in all breeding events and typically do not live as long
as females, which do not participate in every breeding event. Mature males may only live
two months, whereas mature females may survive more than two years.
214 n VERTEBRATES (REPTILES)

Impacts. Cuban treefrogs could negatively affect native treefrog species through preda-
tion and/or competition for food and space. Anecdotal evidence suggests they have replaced
green frogs and squirrel frogs in residential areas. But mostly these frogs are a nuisance in
and around homes and other buildings. Their calls, though not particularly loud, can be
annoying. As they feed and defecate on windows and walls, they create unsightly messes.
They enter homes on house plants, by jumping through an open window or door, or by
making their way into bathroom vent pipes. They end up in the toilet, a startling surprise
for whomever raises the lid. They are also known to have clogged sink drains. Perhaps the
most serious feature of these invaders is the irritating secretions from their skin, which can
cause strong reactions from people with asthma or allergies.
Management. Around the home, Cuban treefrogs can be caught by hand or attracted into
traps of PVC pipe and then, when it is determined the culprit really is this exotic invader,
humanely euthanized. Potential breeding sites can be eliminated by removing any containers
that could collect water. Small-mesh aquarium nets can be used to scoop out eggs and dump
them on the ground to dry out.
Other methods to prevent the spread or breeding of the Cuban treefrog include screening
cisterns and fumigating imported plants. Monitoring its dispersal and taking quick action to
eradicate new populations may slow its spread.

Selected References
Johnson, Steve A. “The Cuban Treefrog (Osteopilus septentrionalis) in Florida.” University of Florida,
IFAS Extension, 2007. http://edis.ifas.ufl.edu/uw259.
National Biological Information Infrastructure (NBII), Comité français de l’UICN (IUCN French
Committee), and IUCN SSC Invasive Species Specialist Group (ISSG). “Osteopilus septentrionalis
(Amphibian).” ISSG Global Invasive Species Database, 2008. http://www.issg.org/database/species/
ecology.asp?si=1261&fr=1&sts=&lang=EN.
Somma, Louis A. “Osteopilus septentrionalis.” USGS Nonindigenous Aquatic Species Database,
Gainesville, FL, 2009. Revised February 11, 2009. http://nas.er.usgs.gov/queries/FactSheet.asp
?speciesID=57.

n Reptiles
n Brown Anole
Also known as: Cuban brown anole, Bahamian brown anole
Scientific name: Norops sagrei
Synonym: Anolis sagrei
Family: Polychrotidae
Native Range. Cuba; the Bahamas; Cayman Brac; Little Cayman; Swan Island, Honduras.
Distribution in the United States. Established throughout peninsular Florida and in coastal
counties of southernmost Georgia. Populations are reported from Hawai’i, Louisiana, North
Carolina, South Carolina, and Houston, Texas; but establishment in those states is uncertain.
Description. This is a small anole lizard adapted for life on the trunks of trees and on the
ground. It has long toes with relatively small toe pads that enable it to run swiftly and jump.
The snout is short and rounded. It is distinguished as an anole by its dewlap, the red-orange
or yellow throat fan. This is much larger in males than females and, when not extended,
 BROWN ANOLE n 215

appears as a pale line on the

throat. A crest along the neck
and back can be erected. The
somewhat laterally compressed
tail sometimes bears a crest-
like ridge. The stripe down the
middle of the back is often
boldly patterned with waves,
zigzags, or diamonds in females,
but it is indistinct in males.
Body color varies from pale gray
to dark brown and even black;
mottling, spots, chevrons, or
light-colored lines may be pre-
sent. Brown anoles can change
their skin color in a matter of
minutes in response to environ-
mental stimuli, aggression, or
reproductive activity.
Adult males have a snout-
vent length (SVL) of about 2.3
in. (6 cm) and weigh about 0.25
oz. (6–8 g). Females are usually
less than 2 in. (5 cm) snout-
vent, and weigh 0.1 oz. (3–4 g).
The tail is normally longer than
the body, giving a total length of
5–8 in. (13–21 cm); but it can
be broken off and regrown, in
which case it will be smaller.
Brown anoles are active
during the day and commonly Top: The brown anole’s native range consists of Cuba, Isla de Pinos, the
seen head-down on a low perch Bahamas, Cayman Brac, Little Cayman, and Swan Island, Honduras.
waiting to ambush prey or bob- Bottom: Brown anoles have established populations throughout
bing their heads and flashing peninsular Florida and in some of the southernmost counties of
their dewlaps in territorial and Georgia. Although reported from a number of other states, they
currently do not seem to have established populations elsewhere.
mating displays. When startled,
(Adapted from Savannah River Ecology Laboratory, n.d.)
these alert and quick lizards run
away on the ground.
Related or Similar Species. The only anole native to the temperate southeastern United
States is the green anole (Anolis carolinensis). It tends to be smaller and more delicately built
than the brown anole. The head and snout are more slender. It is often bright green, but can
change color to a brown that might lead one to mistake it for a brown anole.
In the Miami area, eight other introduced anoles might be confused with the brown anole.
This is especially true of the bark anole (A. distichus), which is smaller and strictly arboreal.
Brown anoles are sometimes erroneously referred to as chameleons because of their abil-
ity to change color, but true chameleons are Old World reptiles with prehensile tails, mitten-
like feet, and eyes mounted on turrets that move independently of each other.
216 n VERTEBRATES (REPTILES)

A. The brown anole has a short, rounded snout. Its long toes with small toe pads enable it to run quickly and
jump, adapting it for life in the trees and on the ground. (John Anderson/iStockPhoto.) B. The brown anole is
typically seen in a head-down position on the lower trunks of trees. The large dewlap identifies it as an anole.
(David Sischo/iStockPhoto.)

Introduction History. The brown anole was first reported from the Florida Keys in 1887. In
the 1940s, it was introduced in at least six different ports in Florida. Most probably, it was a
stowaway on ships. Both major subspecies, the Cuban brown anole (N. sagrei sagrei) and the
Bahaman brown anole (N. sagrei ordinatus), arrived on the mainland, but through subsequent
interbreeding, they have lost their genetic distinctiveness. The brown anole has continued to
expand its range northward, hitchhiking on motor vehicles and in ornamental plants.
Habitat. The brown anole is a habitat generalist that prefers open vegetation in moist
areas such as forest edge, disturbed sites, and the edge habitats created in urban and subur-
ban settings. They spend much of their time on the ground or on trees just a few feet above
the ground. This is a tropical to subtropical species; its northern limits seem to be controlled
by cold winters.
Diet. Brown anoles are predators that feed on a wide range of animals, including annelid
worms, amphipods, isopods, moths, crickets, beetles, flies, spiders, snails, and small verte-
brates. They also eat the hatchlings of the green anole and probably of their own species.
Life History. Brown anoles begin to establish breeding territories in March and April and
defend them for another 5–6 months. Within each male’s territory may be two or more
female territories. Males fight intensely to ward off other males, locking jaws and knocking
one another off their perches. The dewlaps are displayed as part of the aggression ritual
and as courtship behavior to attract females. Females lay a single round egg at time, once
every week or two, throughout the breeding season. The egg is deposited under moist
decaying leaf litter on the ground. It takes 2–3 months for them to hatch; the first usually
 BURMESE PYTHON n 217

Anoles and Evolution

A noles in general are excellent dispersers, and their spread and subsequent
evolution around the Caribbean has made them the subject of much research.
In many respects they are comparable to Darwin’s finches on the Galapagos Islands
and the honeycreepers of the Hawaiian Islands. They diversified into more than
300 species by adapting morphologically and behaviorally to a variety of niches.
The brown anole itself has been used as a model for understanding ecology, ani-
mal behavior, and evolution. Recent work with brown anoles observed natural
selection in action.

emerge in June. Recent hatchlings are 0.6–0.7 in. (15–18 mm) long, SVL. Both males and
females mature the following summer, and most die during the subsequent winter when
about 18 months old.
Impacts. The spread of the nonnative brown anole often appears to coincide with the decline
of the native green anole, but much of the evidence is anecdotal. It has been demonstrated, how-
ever, that where the two species coexist, the green anole shifts its spatial niche higher into the
canopy of trees and seldom utilizes ground perches. When the brown anole is absent, the green
anole is a ground-trunk species. To what degree predation by brown anoles on green anole
hatchlings affects populations of the latter is unknown. In places where the brown anole thrives,
it can become the most numerous reptile in the area. Indeed, it is now one of the most abundant
lizards, and possibly the most abundant vertebrate, throughout Florida.
Management. No attempts to control or eradicate brown anoles have been made. Any
measures taken would likely prove fruitless since the lizard is so prolific and such a generalist.

Selected References
“Brown Anole.” Lizards of Georgia and South Carolina, Savannah River Ecology Laboratory,
Herpetology Program, University of Georgia, n.d. http://www.uga.edu/srelherp/lizards/
anosag.htm.
Campbell, T. “The Brown Anole (Anolis sagrei Dumeril and Bibron 1837).” Institute for Biological
Invasions: The Invader of the Month, February 2001. University of Tennessee, Knoxville, 2002.
http://invasions.bio.utk.edu/invaders/sagrei.html.
Casanova, L. “Norops sagrei” Animal Diversity Web, University of Michigan Museum of Zoology, 2004.
http://animaldiversity.ummz.umich.edu/site/accounts/information/Norops_sagrei.html.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Norops sagrei (reptile).” ISSG Global Invasive Species Database, 2006. http://www.issg.org/
database/species/ecology.asp?si=603&fr=1&sts=&lang=EN.

n Burmese Python
Scientific name: Python molurus bivittatus
Family: Boidae
Native Range. Southern Asia. Found from northeastern India and Bangladesh, through
Myanmar, Thailand, Laos, Cambodia, and Vietnam, into southern China. Also a number
218 n VERTEBRATES (REPTILES)

of isolated populations in
Nepal; Sichuan, China; and Java,
Bali, and Sumbawa, Indonesia,
that may represent relics of a for-
merly much larger distribution
area or may represent introduc-
tions by humans.
Distribution in the United
States. Established in southern
Florida, including Everglades
National Park and Big Cypress
Preserve, and also in Puerto
Rico. They can be found outside
Everglades National Park along
its eastern boundary and
increasingly in more distant
areas, such as Collier-Seminole
State Park and Manatee Coun-
ty. Several individuals have
been observed on Key Largo,
but at the moment, they do not
seem to be established in the
Florida Keys.
Description. One of the
world’s largest snakes, the
Burmese python may grow to
lengths greater than 25 ft.
(7.6 m) and weights close to
300 lbs. (137 kg). The largest
found to date in the Everglades
was 16 ft. (4.8 m) long and
Top: The native range of the Burmese python extends from northeastern weighed 152 lbs. (69 kg).
India through Southeast Asia into southern China. (Adapted from Barker Females become longer and
and Barker 2008.) Bottom: Burmese pythons are invasive in southern heavier than the males, which
Florida, including Everglades National Park and Big Cypress Preserve. are distinguished by their larger
They are also established on Puerto Rico. (Adapted from USGS 2007.) cloacal spurs, the two projec-
tions on either side of the vent
that may be vestigial hind limbs. The light-colored skin of this heavy-bodied snake is beau-
tifully marked with a mosaic of large reddish-brown blotches that are outlined in black. The
pattern begins on the large rectangular head as an arrow-shaped patch. The pet trade has
bred a number of variations, including albino, that may be found in wild Florida popula-
tions, since they all stem from released or escaped pets. Burmese pythons are semiaquatic
but also climb trees.
Related or Similar Species. Two large native pit vipers might be mistaken for young
pythons. The eastern diamondback rattlesnake (Crotalus adamanteus) is also heavy-bodied
and reaches lengths up to 7 ft. (2.1 m), but its head is triangular and the yellow-and-
brown diamond pattern down its back is distinctive. Some Florida cottonmouths
(Agkistrodon piscivorus) have a brown-and-black banded pattern and may reach lengths of
 BURMESE PYTHON n 219

A. The Burmese python, one of the world’s largest snakes, is beautifully marked with large reddish-brown
blotches outlined in black. (Vassil.) B. The arrow-shaped patch on its squarish head is charactersitic. (Vassil.)

6 ft. (1.8 m). However, this snake has a triangular head and a dark cheek stripe that runs
through the eye. Both of the pit vipers are venomous. Another brightly patterned snake that
could be confused with a small python is a colubrid and a constrictor, the corn or red rat
snake (Elaphe guttata guttata). It is common in residential areas and can reach lengths of 6
ft. (1.8 m). It has a spear- or arrow-shaped pattern on the head. Typically, the body is
orange-brown with red or brown blotches outlined in black. The belly is marked with a
black-and-white checkerboard, and two black stripes appear on the underside of the tail.
Introduction History. Burmese pythons were first collected from the wild in southern
Florida in the 1980s. They were first deemed established in Everglades National Park in
2000 based on collections made along Main Park Road in the 1990s. Since then, the number
of pythons in the park has increased dramatically and is now estimated to be 10,000 or
more. The first python in the Florida Keys was documented in 2007 at Key Largo
Hammock Botanical State Park. Pythons are well able to the swim the six miles from the
Everglades to Key Largo.
Burmese pythons in Florida derive from the pet trade. They were either intentionally or
accidentally released by pet owners. Burmese python hatchlings continue to be imported
into the United States from Southeast Asia by the thousands, and domestic breeding of dif-
ferent color morphs also continues.
Habitat. Burmese pythons can live in a variety of open habitats such as swamps, marshes,
grasslands, and woodlands, including the brackish glades and mangroves at the south end of
the Everglades. In Florida, they generally inhabit the same places that alligators do. These
excellent swimmers are usually close to water, which they must enter before shedding, and
can stay underwater without breathing for as long as 30 minutes.
Diet. The Burmese python is a carnivore that prefers live prey but will consume carrion.
Rodents and other small mammals are dietary staples, but it also takes amphibians, reptiles,
and birds. Researchers in Florida have found alligators, limpkins, ibises, coots, House
Wrens, rabbits, squirrels, mice and rats, muskrats, raccoons, Virginia opossums, bobcat,
and white-tailed deer, as well as domestic cats and geese, in the digestive tracts of pythons
captured in the wild.
220 n VERTEBRATES (REPTILES)

Other Big Snakes in Florida

T he Burmese python is perhaps the most publicized invasive snake established in

Florida; however, it is not the only giant snake on the loose. A population of
boa constrictors (Boa constrictor) inhabits a county park and possibly adjacent
areas in southeastern Miami. (It is difficult to know if individuals found beyond
the park are dispersers from the park population or recently released or escaped
pets.) In 2010, researchers reported establishment of another large constrictor,
the Northern African python (Python sebae), in a small area southeast the
Tamiami Trail at State Route 997. Other constrictors with free-living individuals in
Florida, but apparently not reproducing populations, include the green anaconda
(Eunectes murinus), the yellow anaconda (E. notaeus), the reticulated python
(Broghammerus reticulates) and the white-lipped python (Leiopython albertisii).
Sources: Reed, R. N., K. L. Krysko, R. W. Snow, and G. H. Rodda. “Is the Northern African
Python (Python sebae) established in southern Florida?” IRCF Reptiles & Amphibians 17(1):
52–54, 2010.
“Northern African Python, African Rock Python [Non-Native],” Florida Museum of Natural
History, http://www.flmnh.ufl.edu/herpetology/fl-guide/Pythonsebae.htm.

The python has poor eyesight and hunts by using its sense of smell. It also has heating
sensing pits along the upper lip with which it detects the body heat of its prey. It may either
stalk or ambush its victims. Prey items are killed by constriction and swallowed whole.
Life History. Burmese pythons are solitary except when they come together to mate
between December and April. Then groups of one female and several males often form.
Males locate the females chemically, detecting the pheromones secreted by the female. The
male wraps himself around the female’s body and internal fertilization takes place. Two to
three months later, in May and June, the female lays a clutch of 12–36 eggs (up to 100), each
weighing about 7 oz. (200 g). She incubates the eggs for two months by coiling on top of
them. The mother rarely if ever leaves her eggs and will raise their temperature above ambi-
ent temperature by shivering.
The eggs hatch 2–3 months later, in July and August, and parental care ends. The hatch-
lings are 18–24 inches long and grow rapidly if food is abundant. Both males and females
are sexually mature in 2–3 years and at lengths of about 8 ft. (2.6 m). Average lifespan is
15–25 years.
Impacts. Burmese pythons have been in Florida’s Everglades too brief a time for definitive
assessments of their impacts. The concern is that they will compete with native species and
further endanger rare species upon which they prey. Among their known prey are two wad-
ing birds of special concern, the Limpkin (Aramus guarauna) and White Ibis (Endocemus
albus). On Key Largo, prey included the endangered Key Largo woodrat (Neotoma floridana
smalli). They may also may be preying upon native mangrove fox squirrels (Sciurus niger avi-
cennia) and Wood Storks (Mycteria americana). Due to an overlap in diet, they could com-
pete with the state and federally threatened eastern indigo snake (Drymarchon couperi) for
food and space.
With the population explosion that pythons in the Everglades are currently experiencing,
they could become a major ecological problem and hamper efforts to restore the greater
 GREEN IGUANA n 221

Everglades ecosystem. Their future establishment is a major concern in the biologically sen-
sitive Florida Keys.
Pythons pose a threat to small children and pets in residential areas and to small livestock
and poultry in agricultural areas.
Management. Once pythons are established, it becomes impossible to eradicate them.
Efforts therefore focus on controlling numbers and preventing new infestations. The main
strategy involves tracking, capturing, and euthanizing. In southern Florida, volunteer mem-
bers of an Eyes & Ears Team spot pythons on roads and call in a person trained to capture
and dispose of them. Among the people trained to find snakes are mail carriers and package
delivery drivers who daily drive the region’s roads. In March and April 2010, a special
python hunting season was instituted in Florida. Permit holders could also take African
pythons and Nile monitors in this effort to kill as many large exotic reptiles as possible.
Research continues to learn the ways of the python and monitor its spread.
Recent legislation in Florida requires owners of pythons and some other large introduced
reptiles to buy a permit and implant a microchip in the animal to identify its owner. A
federal bill, the Non-native Wildlife Invasion Prevention Act (H.R. 669) that came before
the Congress in 2009–2010, would prohibit the importation of species determined by the
U.S. Fish and Wildlife Service to likely become invasive in the United States.

Selected References
Austin, Jill. “Stopping a Burmese Python Invasion.” The Nature Conservancy, 2009. http://
www.nature.org/wherewework/northamerica/states/florida/science/art24101.html.
Barker, David G., and Tracy M. Barker. “The Distribution of the Burmese Python, Python molurus bivit-
tatus.” Bulletin Chicago Herpetological Society 43(3): 33–38, 2008. Available online at http://
www.vpi.com/sites/vpi.com/files/Barkers.pdf.
Harvey, Rebecca G., Matthew L. Brien, Michael S. Cherkiss, Michael Dorcas, Mike Rochford, Ray W.
Snow, and Frank J. Mazzotti. “Burmese Pythons in South Florida: Scientific Support for Invasive
Species Management.” Publication #WEC242, IFAS Extension, University of Florida, 2008. ttp://
edis.ifas.ufl.edu/pdffiles/UW/UW28600.pdf.
National Biological Information Infrastructure (NBII), Puerto Rico Department of Natural and
Environmental Resources, and IUCN SSC Invasive Species Specialist Group (ISSG). “Python molurus
bivittatus (Reptile).” ISSG Global Invasive Species Database, 2010. http://www.issg.org/database/
species/ecology.asp?si=1207&fr=1&sts.
Padgett, J. “Python molurus.” Animal Diversity Web, University of Michigan, 2003. http://
animaldiversity.ummz.umich.edu/site/accounts/information/Python_molurus.html.
“Python molurus bivittatus.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL, 2009.
Revised October 24, 2007. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=2552.

n Green Iguana
Also known as: Common iguana, gallina de palo
Scientific name: Iguana iguana
Family: Iguanidae
Native Range. Mexico south through Central America to Ecuador on the Pacific side and
southeastern Brazil on the Atlantic slope of South America. Also in the Lesser Antilles
(Curacao, Grenada, St. Lucia, St. Vincent, Trinidad and Tobago, and Utila). Most iguanas
imported as pets into the United States come from captive farming operations in
Honduras, El Salvador, Colombia, and Panama.
222 n VERTEBRATES (REPTILES)

Distribution in the United

States. Established in southern
Florida. Feral populations
are also established on Maui,
Hawai’i; in the Rio Grande
Valley, Texas; on Puerto Rico;
and on the U.S. Virgin Islands.
Description. The green
iguana is one of the largest liz-
ards found in the United
States. Only young animals are
bright green; adults assume a
uniform, grayish-green color,
which they can alter somewhat
in response to social and envi-
ronmental cues. A month or so
before courtship begins, the
males acquire a bright orange
wash on the neck and forelimbs
that persists through the mating
period. Color may also change
diurnally, becoming darker
when the body is cold so as to
absorb more solar energy.
A crest of large spines along
the back and tail is particularly
well developed on males, as is
the very large dewlap under
the throat. The fleshy dewlap is
used in threat and courtship
displays, but also helps absorb
Top: Green iguanas are native to southern Mexico, Central America, and and dissipate heat. The tail is
tropical South America. They also are native to the Lesser Antilles. long and tapering, and ringed
(Adapted from “Green Iguana,” The Wild Ones Animal Index, http:// with broad dark stripes. The
www.thewildones.org/Animals/iguana.html.) Bottom: Released or tympanum is prominent and
escaped pets, green iguanas have established populations in southern covered by a membrane. Large
Florida, on Mau’i in Hawai’i, in the Rio Grande Valley of Texas, and on
scales appear on both sides of
Puerto Rico.
the head; beneath the tympa-
num is a large rounded scale
known as a subtympanic plate. The eyes are set on the sides of the head and covered by
an immobile upper lid and a movable lower lid. On the midline of the top of the head is a
white, light-sensing organ known as the parietal “eye.” It helps coordinate photoperiod with
reproductive status, but also detects the shadows of overhead predators. Sharp teeth are
positioned along the inner sides of both jaws. Nasal glands help expel excess salts.
Adult males may achieve a total length of 6 ft. (1.8 m) and weight of as much as 17.5 lbs.
(8 kg), although 9–12 lbs. (4–6 kg) is more common in Florida specimens.
Related or Similar Species. Some other large introduced lizards might be mistaken for
young green iguanas. The black spinytail iguana (Ctenosaura similis) also has a crest of spines
 GREEN IGUANA n 223

along the back, but has black banding on the body. Spiny scales circle around the tail. The
young are bright green like young green iguanas, but have broken bands of black around
the mid-body. The largest of these lizards may be 3–4 ft. (approximately 1 m) in total length.
The brown basilisk (Basiliscus vittatus) has a large crest behind the head and a brown body; it
only grows up to 2 ft. (60 cm) long. Knight anoles (Anolis equestris) and Jamaican giant
anoles (A. garmani), are both green but only about 1 ft. (30 cm) long.
Introduction History. Baby green iguanas are imported into Florida by the tens of thou-
sands each year in the pet trade. They were first reported outside captivity in the Miami area
in 1966, but populations probably did not become established until the 1980s, when they
became popular pets. Mature green iguanas become powerful animals that can bite, scratch,
and whiplash owners with their tails. When they become too much to handle, they are often
released into the wild. The earliest sightings of free-living iguanas were in Key Biscayne,
Hialeah, and Coral Gables, and in the vicinity of Miami International Airport. Undoubtedly,
these reptiles were intentional releases by pet owners or accidental escapes. Since that time,
they have expanded northward on the Atlantic coast, establishing populations in Broward
County by 2001 and in Palm Beach County by 2003. Along the Gulf coast, iguanas were
reported as established in Monroe County (Coral Reef State Park) in 1995 and in Lee County
(Fort Myers and Cape Coral) sometime in the 1990s. Increasing numbers are seen in Martin
and St. Lucie counties, but the iguana does not yet seem to be established in those areas.
Iguanas have also been reported in Pinellas County on the Gulf coast. Expansion much farther
north seems to be halted by irregular freezes. Green iguanas are relatively common in the
Everglades, and individuals have been reported as far south in the Keys as Stock Island.
Habitat. In Florida, the green iguana occurs in frost-free coastal areas. They thrive only
where temperatures range from 79° to 95°F (24–32°C) and where sunlight is sufficient to
let their bodies produce vitamin D. They are mostly arboreal and prefer large trees overhang-
ing water, into which they dive to escape predators. Therefore, they are likely to be near
canals, ditches, and ponds, and in mangroves lining the shores of shallow bays. Iguanas also
inhabit urban and suburban yards and like to bask on tree branches, sidewalks, docks, and
seawalls, or in open, mowed areas.
Diet. Green iguanas are herbivores and feed on leaves, flowers, and fruits. Juveniles con-
sume insects and other animal matter in addition to plant foods. Digestion occurs in the
hindgut, and juveniles eat the droppings of adults in order to acquire the microflora neces-
sary to break down plant material. Among preferred food plants are hibiscus, orchids, roses,
garden greens, and squashes and melons.
Life History. Green iguanas breed during the dry season so that the young hatch during
the wet season when food will be most plentiful. Iguanas of both sexes come together in
sandy areas to mate during a single nesting season each year. Dominant males scent-mark
territories and females with pheromones secreted from femoral pores on the undersides of
their thighs. Fertilization is internal. About 65 days after mating, each female excavates a
burrow 1.5–3 ft. (0.5–1.0 m) deep and, over a period of three days, deposits 20–70 eggs
in it. If nesting space is scarce, several females will use the same nest. No parental care is
given to the nest or eggs. Eggs measure roughly 0.6 in. (15 mm) in diameter and 1.5 in.
(35–40 mm) long. Hatchlings emerge 10–15 weeks later, usually during July and August,
and look like miniature females without spiny crests. They are 6.5–10 in. (17–25 cm) in
total length. The juveniles remain in family groups for a year; curiously, the young males
protect the females from predators. Green iguanas become sexually mature at 3–4 years of
age. The reproductive life of a female lasts several years. After mating, she is able to store
sperm for several years and fertilize her eggs even if a male is not present.
224 n VERTEBRATES (REPTILES)

A. The green iguana has a crest of long spines and a very large dewlap. (Susan Woodward.) B. This large lizard
has a long tail ringed with dark bands and a prominent external ear drum. (Susan Woodward.)

Impacts. Threats to native lizards are probably slight, since adult green iguanas are herbi-
vores and all native species are carnivores. Green iguanas sometimes use the burrows of the
Florida Burrowing Owl (Athene cunicularia floridana), a federally endangered subspecies, and
could prevent the birds from nesting or destroy their eggs and nestlings. Iguanas eat the
native butterfly sage (Cordia globosa), a plant of special concern in Florida, and consume yel-
low nickerbean (Caesalpinia bonduc), a primary host for the larvae of the endemic and endan-
gered Miami blue butterfly (Hemiargus [=Cyclargus] thomasi bethunebakeri). They are
potential dispersers of any invasive plants upon which they feed.
In residential areas, iguanas can destroy vegetable gardens and ornamental plants and
pose a health hazard by spreading salmonellosis when they defecate in pools or on docks
and sidewalks. Their droppings are also unsightly and smelly.
In Puerto Rico, basking green iguanas are a runway hazard at the Luis Muñoz Marı́n
International Airport and must be shooed away before planes can take off.
 NILE MONITOR n 225

Management. Iguanas can be removed from private property without special permits.
They can be caught legally in Florida by hand, noose pole, nets, or live traps. It is illegal,
however, to release captured animals; so they must be kept as pets or captive breeding stock
or be destroyed. Feral adults rarely tame down. The best control measures are to discourage
iguanas by removing protective cover from the yard, sheathing trees with metal to prevent
their climbing up to sun bathe, not planting their favorite food plants, and protecting valued
plants in screened enclosures.
In open country, it is very difficult to capture iguanas because they are such strong
swimmers and dive into water at the first sign of danger. Sometimes in cold weather, when
they are sluggish, they can be picked off branches or off the ground. Local iguana popula-
tions along canals and in mangroves have been reduced using boats to collect those knocked
off branches when temperatures are in the 40s.
In Florida, iguanas are protected only by anticruelty laws. In Hawai’i, strict laws regulate
the importation and possession of green iguana, and violations can lead to fines of $200,000
and up to three years in jail.

Selected References
Gibbons, Whit, Judy Greene, and Tony Mills. Lizards and Crocodilians of the Southeast. Athens:
University of Georgia Press, 2009.
Gingell, F., Biology of Amphibians and Reptiles, and J. Harding. “Iguana iguana” Animal Diversity Web,
University of Michigan Museum of Zoology, 2005. http://animaldiversity.ummz.umich.edu/site/
accounts/information/Iguana_iguana.html.
“Green Iguana.” Wikipedia, 2009. http://en.wikipedia.org/wiki/Green_Iguana.
“Green Iguana—Iguana iguana.” Florida’s Exotic Wildlife. Species detail. MyFWC.com, Florida Fish
and Wildlife Conservation Commission, n.d. http://myfwc.com/wildlifehabitats/nonnatives/
reptiles/green-iguana/.
Kern, W. H., Jr. “Dealing with Iguanas in the South Florida Landscape.” Publication #ENY-714, IFAS
Extension, Univeristy of Florida, 2009. http://edis.ifas.ufl.edu/in528.
Masterson, J. “Iguana iguana: Green iguana.” Smithsonian Marine Station at Fort Pierce, 2007. http://
www.sms.si.edu/irlspec/Iguana_iguana.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive species Specialist Group
(ISSG). “Iguana iguana (Reptile).” ISSG Global Invasive Species Database, 2006. http://
www.invasivespecies.net/database/species/ecology.asp?si=1022&fr=1&sts.

n Nile Monitor
Also known as: Money monitor
Scientific name: Varanus niloticus
Family: Varanidae
Native Range. Much of Africa between 15° N and 15° S. It occurs in forests and savannas
near permanent water. It is absent from desert regions, but has been found at elevations
greater than 6,500 ft. (2,000 m).
Distribution in the United States. Known to be established in at least two counties in
southern Florida: Lee County (Cape Coral) and adjacent Charlotte County. Sightings of
Nile monitors are reported from several other counties, including Broward, Collier,
DeSoto, Miami-Dade, Orange, and Palm Beach.
Description. This is the largest lizard now occurring outside of captivity in the United
States and the longest lizard in its native Africa. Nile monitors can reach SVLs averaging
226 n VERTEBRATES (REPTILES)

23.5–31.5 in. (60–80 cm) and

total lengths over 7.5 ft. (2.4 m).
They may weigh more than 22
lbs. (10 kg). Like all monitors,
they have long necks, long tails,
and an obviously forked tongue.
The head is narrow and wedge-
like and shorter than the neck.
The tongue is blue. The back-
ground color is gray-brown or
olive green. The jaws and head
bear cream-colored stripes that
graduate into chevrons on the
neck. Six to nine bands of yellow
dots encircle the body. Said to
look like coins, these dots are
the origin of one of the African
names for this species, the
“money monitor.” Juveniles are
more brightly patterned than
adults. Nile monitors have large,
strong claws for digging and a
muscular tail that is laterally com-
pressed for swimming and usu-
ally 1.5 times as long as the body.
Related or Similar Species.
The Asian water monitor
(Varanus salvator) looks a bit
like the Nile monitor and grows
even longer, reaching lengths of
10 ft. (3 m). It lacks the bands
Top: The Nile monitor is native to forests and savannas in much of Africa. of “coins” adorning the Nile
(Adapted from map at http://en.wikipedia.org/wiki/File:Nile_monitor_ monitor. As far as it is known,
range.PNG.) Bottom: The Nile monitor is established in at least two water monitors are not living
counties in southern Florida and has been reported from several others. free in Florida, but they are
(Adapted from Somma 2007.) easily obtained through the pet
trade, and it is entirely possible
that they will become established in the state. Some reported observations of individual Nile
monitors may actually have been escaped or released water monitors.
Introduction History. Free-living Nile monitors were first spotted at Cape Coral in 1990.
Their origins are unknown but presumed to be related to the pet trade. One rumor has it
that a pet store that went bankrupt in the 1980s released their Nile monitors into what
was then a wilderness of saw palmettos, slash pine, and mangrove. Another story suggests
that wholesalers of exotic animals purposefully released monitors to establish a breeding
population from which they could capture young animals to sell. It is perhaps most likely
that the animals came from pet owners who found that the cute little hatchlings they had
purchased had turned into large, temperamental reptiles they no longer wanted or could
manage, so they “humanely” released them. The animals are also quite able to escape from
 NILE MONITOR n 227

cages on their own. As the cape

was developed, monitors came
into more frequent contact with
residents and the number of
observations increased, espe-
cially along canals in the western
part of the peninsula. By 2002,
monitors had crossed Matlacha
Pass onto Pine Island. Sanibel
Island to the south lies within
reach. Since some populations
in Africa are adapted to temper-
ate climate, monitors from those
regions, if introduced to the
United States, theoretically could
spread throughout Florida and
the Gulf states and possibly as
far north as the Carolinas. The Nile monitor has a narrow wedge-shaped head and a laterally com-
Habitat. Nile monitors are pressed tail for swimming. (Dr. Gordon E. Robertson.)
well adapted to both terrestrial
and aquatic habitats. They prefer cover near permanent bodies of water. In Florida, favor-
able conditions exist in mangrove swamps, along the edges of both freshwater and saltwater
marshes, and on the banks of rivers and canals. In Lee County, they also live in pine flatwoods,
on golf courses with ponds, and in suburban neighborhoods and urbanized areas. They shelter
in burrows that they dig into canal banks or in those excavated by other animals. In Africa, they
live in a variety of climates at different elevations and will hibernate during cold weather in
more temperate regions.
Diet. Nile monitors are indiscriminate carnivores and scavengers. They will consume inver-
tebrates such as cockroaches, mangrove tree crabs, snails, and clams; amphibians, including the
introduced Cuban treefrog (see Amphibians, Cuban Treefrog); lizards, snakes, baby alligators,
and turtles, as well as the eggs of reptiles; birds and bird eggs; and small mammals such as
rodents and cats. They also eat carrion, garbage, and feces. Young monitors are more arboreal
than mature adults and feed almost exclusively on fast-moving invertebrates and lizards. With
increasing age, they consume less active prey that is more armored and must be crushed by
strong jaws and teeth before being swallowed. These intelligent animals will hunt cooperatively.
Life History. What information is available on life history comes from studies done in the
monitor’s native habitats in Africa and may or may not be directly applicable to populations
in Florida. In the Sahel region of Africa, females are sexually mature when they are two years
old and have a SVL of about 14 in. (36 cm). They breed every other year, so during any given
breeding season, only 50 percent of adult females lay eggs. In some areas, females feed and
accumulate fat in spring and summer. Eggs are laid during the winter dry season, buried
in the ground and in active termite mounds. Clutches average 35 eggs or more, with smaller
females producing many fewer than larger ones. Each egg is about 3.5 × 1.6 in. (6 × 4 cm).
Hatchlings emerge 6–10 months later, near the beginning of the wet season. They are 6–12
in. (15–32 cm) in total length and will more than double in size during the first year of life.
Nile monitors can live more than 10 years in the wild.
Impacts. Nile monitors are potentially the most destructive lizard introduced to Florida.
Nile monitors in Lee County, Florida, could have negative effects on native crocodilians,
228 n VERTEBRATES (BIRDS)

(the American alligator [Alligator mississippiensis] and, especially, the American crocodile
[Crocodylus acutus]), for in Africa, Nile monitors not only have diets similar to those of croco-
diles, but also are the major predators of crocodilian eggs and hatchlings. Their propensity to
occupy existing burrows raises concern for two protected species on the Cape Coral peninsula,
the Burrowing Owl (Athene cunicularia) and the gopher tortoise (Gopherus polyphemus), whose
nesting burrows could be taken over and eggs and young consumed. If Nile monitors were to
invade Sanibel Island, the large rookeries of pelicans and herons would be threatened. If they
gain access to Sanibel’s beaches, the nests of sea turtles would be vulnerable to their depreda-
tions. Should Nile monitors make their way into southeastern Florida and the Keys, a number
of listed species could become prey, including the southeastern beach mouse (Peromyscus polio-
notus niveiventris), Key Largo cotton mouse (Peromyscus gossypinus allapaticola), Key Largo
woodrat (Neotoma floridana smalli), and silver rice rat (Oryzomys palustris argentatus).
When cornered, Nile monitors become very defensive and attack with their teeth, sharp
claws, and strong tails. Humans and pets are at risk of serious wounds that can become sep-
tic due to the bacteria in monitors’ mouths. Small dogs and domestic cats are also at risk
since they can be caught easily and eaten by monitors. Reports from Cape Coral suggest a
decline in pet populations as the monitor population increased.
Management. Once established, Nile monitors are very difficult if not impossible to
eradicate. Population reduction and prevention of further infestation are the main manage-
ment strategies. Monitors may be captured with nooses or live traps or by digging them
out of the ground. Arboreal hatchlings can be caught at night by hand. Regulations can keep
monitors from being purchased in the first place by casual, untrained pet owners.

Selected References
Enge, K. M., K. L. Krysko, K. R. Hankins, T. S. Campbell, and F. W. King. “Status of the Nile Monitor
(Varanus niloticus) in Southwestern Florida.” Southeastern Naturalist 3(4): 571–82, 2004.
Gore, Jeff, Kevin Enge, and Paul Moler. “Nile Monitor (Varanus niloticus) Bioprofile.” MyFWC.com,
Florida Fish and Wildlife Conservation Commission, n.d. http://www.scribd.com/doc/26904049/
NILE-MONITOR-Varanus-Niloticus-Bioprofile-Compiled-By.
Kruse, Michael. “Nile Monitor Lizards Invaded Florida and They’re Winning the Battle.” St. Petersburg
Times, June 21, 2009. Available online at http://www.tampabay.com/news/environment/wildlife/
article1011745.ece.
“Nile Monitor.” Florida’s Exotic Wildlife, MyFWC.om, Florida Fish and Wildlife Conservation
Commission, Tallahassee, 2007. http://myfwc.com/wildlifehabitats/nonnatives/reptiles/nile-monitor/.
Somma, Louis A. “Varanus niloticus.” USGS Nonindigenous Aquatic Species Database, Gainesville, FL,
2009. Revised August 28, 2007. http://nas.er.usgs.gov/queries/FactSheet.asp?speciesID=1085.
Youth, Howard. “Florida’s Creeping Crawlers.” ZooGoer 34(3), Smithsonian National Zoological Park,
2005. http://nationalzoo.si.edu/Publications/ZooGoer/2005/3/reptilefeature.cfm.

n Birds
n Cattle Egret
Also known as: Buff-backed Heron
Bubulcus ibis
Family: Ardeidae
Native Range. Parts of Africa and Asia and southern Spain and Portugal.
 CATTLE EGRET n 229

Distribution in the United

States. This bird has been
reported in all states, but breed-
ing takes place primarily in the
southeastern United States as far
north as Virginia in the east and
Kansas in the center of the coun-
try. Resident populations can
also be found in southernmost
California, Hawai’i, and Puerto
Rico.
Description. The Cattle
Egret is a small white heron
about 20 in. (46–56 cm) long
with a wingspan of about 36
in. (88–96 cm). It is stocky and
has a thick neck, shorter than
its body length. When standing
still, it typically assumes a
hunched position. Most of
the year its plumage is white,
but for a short time during the
mating season, orange-buff
feathers occur on the crown,
throat, and back. The bill is rel-
atively short and yellow, and
the legs yellow to gray-green
during the non-breeding sea-
son, but both turn red during
the breeding season.
In flight, the Cattle Egret
holds its neck tight to the body. Top: The Cattle Egret’s vast native range spans much of non-desert Africa,
Cattle Egrets are gregarious and the southern Iberian peninsula of Europe, and South and Southeast Asia.
fly to and from feeding areas in (Adapted from map at http://wapedia.mobi/en/File:Ardea_ibis_map.svg.)
flocks. When it walks, it tends Bottom: Cattle Egrets can be found in all states, but breeding populations
to sway in an exaggerated strut occur primarily in the southeastern states, southern California, Hawai’i,
and then suddenly dart forward and Puerto Rico. (Adapted from map by Cornell Lab of Ornithology.
http://www.allaboutbirds.org/guide/Cattle_Egret/lifehistory.)
to stab its prey. Most often,
Cattle Egrets forage alongside
grazing animals, waiting for the livestock to flush insects. They may also pick parasites off
large herbivores and be seen standing on the animals’ backs.
Cattle Egrets are usually silent. Their voice consists of a low, nasal “rik-rak.”
Related or Similar Species. The Cattle Egret is smaller than any native herons or egrets
that have white plumage. It might be confused with the somewhat taller Snowy Egrets
(Egretta thula) and juvenile Little Blue Herons (Egretta caerulea). Snowy Egrets are distin-
guished by their black legs and yellow feet, as well as a black bill. They are adorned with
long, lacy white plumes on the neck and back during the breeding season. Juvenile Little
Blue Herons have dull, greenish legs and a pale gray or greenish bill. Both of these birds
230 n VERTEBRATES (BIRDS)

A. The Cattle Egret is a small, stocky heron. (Robbie Taylor/Shutterstock.) B. Cattle Egrets commonly hunt
insects flushed by cattle or other large mammals. (Donna Beeler/Shutterstock.)

are associated with freshwater, where they feed on fish and aquatic invertebrates; whereas
the Cattle Egret is generally found in upland pastures and feeds mostly on insects.
Introduction History. The introduction of the Cattle Egret to the Americas appears to
have been a natural event. It suddenly arrived in Suriname, South America, in the late
1870s and 1880s, presumably having simply flown across the Atlantic Ocean from the
African continent. It spread throughout South America, filling a vacant niche as a unique ter-
restrial member of the heron family. By 1917, it was in Colombia, and in the 1940s, it
showed up in Florida. In the United States, its long-distance wandering habits let it rapidly
expand its range. By the 1960s, it was in California as well as Canada.
In contrast to the natural range expansion that carried the Cattle Egret to all of the
continental United States, the bird was deliberately introduced to Hawai’i in 1959. Local
ranchers funded efforts by the Hawaiian State Board of Agriculture and Forestry to establish
Cattle Egrets as a biological control for flies and other insects that were cattle pests. Twenty-
five egrets were released at Kipu Ranch on Kaua’i, and 105 others were distributed among
ranches on Hawai’i, Maui, Moloka’i, and O’ahu. The Honolulu Zoo also received some birds.
The egrets began nesting soon after release and established large populations on Kaua’i and
O’ahu, where they came to attain pest status.
Habitat. Cattle Egrets commonly inhabit pastures, marshes, and ploughed fields. They
thrive in altered habitats and tolerate busy roadsides and urbanized areas well. Most often,
when feeding, they associate with cattle or other livestock. Cattle Egret roosts are usually
 CATTLE EGRET n 231

in trees or shrubs near water. They are colonial nesters and frequently nest in dense rook-
eries with other herons and egrets.
Diet. Insects are the mainstay of the Cattle Egret diet. They consume live insects flushed
by grazing mammals, but are also known to follow tractors, plows, lawn mowers, and even
airplanes to snatch insects disturbed by the machines. Most of their food consists of grass-
hoppers, crickets, flies, and beetles, but when insects are not abundant, they will take spi-
ders, certain moths, frogs, and crayfish as well as bird eggs and nestlings. They also
scavenge in refuse for edible leftovers.
Life History. Males establish breeding territories within large colonial nesting areas from
spring through early summer. Courtship displays attract females, and pair-bonds are estab-
lished that last the season. Bulky nests of sticks are built by the female from materials carried
to her by the male. Nest-building and mating commonly last only three days. The breeding
colors are lost as soon as mating is over. When egg-laying begins, one pale blue egg is pro-
duced every other day. The eggs are not brooded until the last egg is laid. Clutches usually
consist of 34 eggs. Both adults incubate the eggs for approximately 24 days. The eggs hatch
in the order in which they were laid; usually only the first two survive to fledge. The siblings
begin to compete with each other for food when less than a week old. Two to three weeks
after hatching, they leave the nest to climb around the rookery but continue to beg for food
from the parents. They begin to fly a week or so later and become independent about
2.5 months after hatching. Juveniles disperse hundreds of miles in apparently random direc-
tions. Even as adults, the egrets are highly migratory and wander widely. Cattle Egrets
become members of the breeding population at 2–3 years of age.
Impacts. In most instances, Cattle Egrets have little or no impact on native heron species.
They utilize different habitats for feeding, have a different diet, and breed after the nesting sea-
sons of native birds. Some concern does remain that they could potentially displace native her-
ons and egrets in rookeries, since they occur in such large numbers. Cattle Egrets are now more
numerous in North America than all other herons and egrets combined.
In Hawai’i, the birds are not quite as benign as on the mainland. They are known to feed
on the eggs and young of endangered wetland birds such as the Black-necked Stilt or A’eo
(Himantropus mexicanus) and could compete with such insect-eating species as frogs, toads,
and skinks. Furthermore, they have become a nuisance for aquaculture on Oahu because
they feed on prawns and are a hazard at airports in Honolulu, Lihue, and Hilo.
Large rookeries anywhere can become nuisances due to noise, odor, and potential public
health threats.
Management. Population reduction and control efforts in Hawai’i are likely to have only
temporary effects, since the birds are so mobile and will return to areas from which they
have been removed. Techniques that repel Cattle Egrets may have some success at airports
and nuisance rookeries. Elsewhere, the bird is usually not viewed as a problem and may
even be welcomed as a way of helping to control insect pests of cattle.

Selected References
Ivory, A. “Bubulcus ibis.” Animal Diversity Web, University of Michigan Museum of Zoology, 2000.
http://animaldiversity.ummz.umich.edu/site/accounts/information/Bubulcus_ibis.html.
Masterson, J. “Bubulcus ibis (Cattle Egret).” Species Report, Smithsonian Marine Station at Fort Pierce,
2007. http://www.sms.si.edu/irlspec/bubulcus_ibis.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Bubulcus ibis (bird).” ISSG Global Invasive Species Database, 2006. http://www.issg.org/
database/species/ecology.asp?fr=1&si=970&sts.
232 n VERTEBRATES (BIRDS)

“The Cattle Egret.” Hawaii Nature Focus, Booklet No. 10, Kilauea Point Natural History Association,
n.d. http://www.kilaueapoint.org/education/naturefocus/hnf10/index.html.

n Common Myna
Also known as: Indian Myna, House Myna
Scientific name: Acridotheres tristis
Family: Sturnidae
Native Range. Southern Asia from southeastern Iran and Afghanistan through Pakistan,
India, Nepal, and Sri Lanka to Southeast Asia and southern China.
Distribution in the United
States. Established on all the
major islands of Hawai’i and in
southern Florida.
Description. This member of
the starling family is a stocky
brown bird with a glossy black
head and neck, yellow bill, and
yellow legs. A patch of bare yel-
low skin occurs behind the eye.
Adults are 9.0–9.8 in. (23–
25 cm) long and have a wing-
span of 18 in. (46 cm). In flight,
the white tips of tail feathers
and white wing patches are
clearly visible. Immatures have
duller colors and browner
heads than adults. Common
Mynas have a large repertoire
of raucous calls, squeaks,
clicks, and whistles. Both males
and females sing and often fluff
their feathers and bob their
heads when vocalizing. At
dawn and dusk they engage in
loud choruses and at their com-
munal roosts keep up a noisy
chatter well after dark.
Common Mynas strut across
the ground, rather than hop-
ping as many birds do. They
tend to travel in pairs.
Top: The native range of the Common Myna stretches across southern Related or Similar Species.
Asia from southwest Iran to Vietnam. (Adapted from “Common Mynas Common Mynas are related to
aka Indian Mynas aka Talking Mynas.” AvianWeb, 2006.) Bottom: the European Starling (see
Common Mynas are established in southern Florida and on all the
Birds, European Starling) and
major islands of Hawai’i. (Adapted from “Common Myna Acridotheres
tristis” 2003.) share some of their annoying
 COMMON MYNA n 233

behaviors. The Hill Myna

(Gracula religiosa) is the more
common pet talking myna. It is
larger than the Common Myna
and has a black body with
fleshy yellow wattles on the
head that extend back from
below the eyes to the nape.
The bill is red or orange. This
mostly arboreal bird has esca-
ped captivity and may be en-
countered in urban gardens
and parks in Miami, Florida,
and in several large cities in so-
uthern California and Hawai’i.
Introduction History. The
Common Myna was brought to
Hawai’i from India in 1865 by
Dr. William Hildebrand in an A bare patch of yellow skin behind the eye distinguishes the Common
Myna. The white wing patch is more conspicuous in flight. (K. W.
effort to control army worms
Bridges, “Common Campus Birds,” University of Hawai’i at Manoa. http://
(Spodoptera mauritia) and army www.botany.hawaii.edu/biology101/birds/common_myna.htm)
cutworms (Euxoa auxiliaris) that
were devastating the islands’ sugar cane and pastures. They did help control cutworms, but
also adapted well to urban life and were abundant in Honolulu by 1879. Eventually, they
inhabited all the main islands.
Common Mynas’ ability to “talk,” i.e., mimic human voices, and their general intelligence
made them part of the pet trade. Mynas in Florida most likely derive from escapes or inten-
tional releases of pet birds. The first sighting and confirmed breeding of the Common Myna
in the state was in Miami in 1983. It has since been reported from 19 other counties within a
300-mi. (480 km) radius of that city and may have spread as far north as Sapelo Island,
Georgia. Although the bird has dispersed rapidly, its numbers have not exploded, and small
scattered populations seem to be the rule.
Habitat. Common Mynas prefer open country such as farmland and suburban and urban
parks and yards. The small populations in Florida apparently prefer shopping mall parking
lots. In their native range, they inhabit open lowland woodlands and the edges of settle-
ments and roost in isolated stands of tall trees. They are cavity nesters, but are not restricted
to holes in trees, building their bulky nests in any nook or cranny available.
Diet. Common Mynas are omnivorous. They take insects—especially grasshoppers, small
vertebrates, and carrion, feeding mostly on the ground. Its generic name Acridotheres means
“grasshopper hunter.” Mynas also feed on grains and fruits and sometimes on the eggs and
nestlings of other bird species. In Florida, they are known to beg for French fries at fast food
establishments.
Life History. Common Mynas begin to build their nests of grass, leaves, and twigs in late
February or early March. Both parents construct the nest, which may be placed in tree
cavities, crevices in buildings, martin houses, and the tops of coconut and date palms, and
aggressively defend their nesting territories. Between March and July, the female lays 2–5
blue eggs, and she and the male both incubate them. Chicks hatch in 13–18 days. They
fledge when 3–4 weeks old. Parental care continues for another month while the young
234 n VERTEBRATES (BIRDS)

learn to forage for themselves. A female may produce 1–3 clutches a year. Both sexes reach
maturity at one year of age.
Juveniles form small flocks once they become independent of their parents. Adults forage
in loose groups of 5–6 birds. During the nonbreeding season, Common Mynas will roost
together in huge flocks numbering more than 1,000 birds.
Impacts. The noise and droppings from large flocks of Common Myna can make them a
nuisance in urban and suburban settings. However, due to the large numbers of insects they
consume and their bold antics, many people like this bird. In Hawai’i, they serve as a reser-
voir for avian malaria (see Microorganisms, Avian Malaria) and are thus implicated in the
decline of endemic island birds. They disperse the seeds of lantana (Lantana camara; see
Volume 2, Shrubs, Lantana)—which is an invasive weed in both Hawai’i and Florida—and prey
upon the eggs and nestlings of native songbirds and seabirds such as the Wedge-tailed
Shearwater (Puffinus pacificus). In Florida, they compete with Purple Martins for nest sites, but
do not congregate in the large flocks typical elsewhere, so are not as problematic as in Hawai’i.
The Common Myna has been introduced to many parts of the world, including many
islands in the Pacific. It is such an agricultural pest in fruit- and wheat-growing areas (espe-
cially in Australia) that the IUCN has included the Common Myna on its nomination list for
“100 of the World’s Worst invasive alien species.”
Management. Although the Common Myna has not become the pest it has in other parts
of the world, its importation into the United States is now prohibited.

Selected References
“Common Myna Acridotheres tristis.” Florida’s Breeding Bird Atlas: A Collaborative Study of Florida’s
Birdlife, 2003. http://legacy.myfwc.com/bba/docs/bba_COMY.pdf.
“Common Myna—Acridotheres tristis.” Florida’s Nonnative Wildlife. Species detail. MyFWS.com,
Florida Fish and Wildlife Conservation Commission, 2009. http://myfwc.com/wildlifehabitats/
nonnatives/birds/common-myrna/.
“Common Mynas aka Indian Mynas aka Talking Mynas.” AvianWeb, 2006. http://www.avianweb.com/
commonmynas.html.
IUCN/SSC Invasive Species Specialist Group (ISSG). “Acridotheres tristis (Bird).” ISSG Global Invasive
Species Database, 2006. http://www.issg.org/database/species/ecology.asp?si=108&fr=1&sts.
Lin, T., and T. Root. “Acridotheres tristis.” Animal Diversity Web, University of Michigan Museum of
Zoology, 2007. http://animaldiversity.ummz.umich.edu/site/accounts/information/Acridotheres
_tristis.html.
Pranty, Bill. “Common Myna (Acridotheres tristis).” In Pranty, B., J. L. Dunn, S. C. Heinl, A. W. Kratter,
P. E. Lehman, M. W. Lockwood, B. Mactavish, and K. J. Zimmer, “Annual Report of the ABA
Checklist Committee: 2007–2008,” 37. Birding 40: 32–38, 2008. Available online at http://
www.aba.org/birding/v40n6p32.pdf.
“The Common Myna.” Nature Focus Booklet #5. Kilauea Point Natural History Association, n.d. http://
www.kilaueapoint.org/education/naturefocus/hnf5/index.html.

n Eurasian Collared-Dove
Scientific name: Streptopelia decaocto
Family: Columbidae
Native Range. South Asia. Prior to the 1600s, the native range was apparently restricted
to India, Sri Lanka, and Myanmar. Its range later expanded by natural means and through
deliberate introductions to Turkey and southeastern Europe, and by the end of the twentieth
century, it was found throughout Europe.
 EURASIAN COLLARED-DOVE n 235

Distribution in the United

States. Southeastern and cen-
tral United States; Los Angeles
region of California. The range
is rapidly expanding, however,
and it may soon be in all 49
continental states.
Description. Like several
native doves, the Eurasian
Collared-Dove is a medium-
size bird with short legs and a
small head. The body is pale
gray with a slightly pinkish
wash over the head and breast.
The slender bill is black, the
legs and feet are mauve, and
the eyes are red. A black line
forms a distinct “collar” on the
back of the neck; above it is a
white line. The square tail is
white underneath. The sexes
look alike. Adults are 12–13
in. (30–33 cm) long and have a
wingspan of 18–22 in. (45–
55 cm). Average weight is 7 oz.
(200 g).
Juveniles are similar to
adults except that their feathers
have pale reddish edges and
their eyes are brown. The legs
are a darker brownish red.
Until they are three months Top: The Eurasian Collared-Dove is native to South Asia. Prior to the
old, they display no clear collar. 1600s, it seems to have been restricted to the area shown on the map,
Eurasian Collared-Doves but since that time its range has expanded through Turkey to all of
have a rhythmic “coo COOO Europe. Bottom: The range of the Eurasian Collared-Dove is rapidly
cup” resembling that of the expanding in the United States. Currently found in southeastern and cen-
tral states, it may eventually invade all of the continental United States.
native Mourning Dove (Zenaida
(Adapted from Sibley 2000.)
macroura), but lower in pitch.
During display flights, they pro-
duce a harsh, nasal “krreew” call. The wings do not whistle when the bird takes flight.
Related or Similar Species. The native Mourning Dove is smaller and lighter and has a
long pointed tail. When it takes flight, the wings produce a whistling sound. The native
White-winged Dove (Zenaida asiatica) is about the same size as the Eurasian Collared-
Dove and has a square tail, but the tail is dark underneath with a white edge. Its white wing
patch is distinct at rest and in flight. Neither native species has a black collar.
Closely related and very similar in appearance to the Eurasian Collared-Dove is the somewhat
smaller, introduced Ringed Turtle-Dove (Streptopelia risoria) for which it was initially mistaken.
The turtle-dove is a feral bird of extremely limited distribution in the United States. It is
236 n VERTEBRATES (BIRDS)

distinguished from the Eurasian

Collared-Dove by its very light
color and white tail and undertail
coverts as well as its call.
Introduction History. The
Eurasian Collared-Dove came
to the New World as a cage
bird. It first became established
in the wild in the Bahamas in
1974 when birds escaped from
a breeder. They are excellent
colonizers, as their history in
Europe demonstrates; and by
the 1980s, they had expanded
their range into southern
The Eurasian Collared-Dove is pale gray with a distinct black “collar” on Florida. By 2009, they were
the back of the neck. (Gregg Williams/Shutterstock.) well established in the south-
eastern United States, especially
along the Atlantic and Gulf coasts; isolated populations were established in the southern
Plains states and southern Rocky Mountain states as well as in southern California. It has
been reported in Alaska and in the Great Lakes region, but these may represent local escapes
of captive birds. In some or the disjunct areas, the dove has been deliberately released for
sport hunting or represents an accidental introduction. At least some of these populations
are expected to serve as nuclei from which further range expansion will occur.
Habitat. Eurasian Collared-Doves are most abundant in coastal, agricultural, and suburban
habitats. They forage in open areas and tend to avoid forested areas and areas that are intensively
cultivated. Nests are constructed in trees and shrubs and on man-made structures.
Diet. These birds are primarily seed and fruit eaters, but also consume small inverte-
brates. They generally forage on the ground but will eat at bird feeders.
Life History. Eurasian Collared-Doves are monogamous and have long breeding seasons;
they may breed year-round in warm climates. In Florida, a pair may raise three or more
broods each year. Each clutch consists of two eggs, the first laid being significantly larger
than the second. Both parents incubate the eggs until they hatch about 15 days after laying.
Hatching is asynchronous, with the second egg hatching 12–40 hours after the first. The
young are fed regurgitated crop milk; as they get older, seeds become part of the nestlings’
diet. Juveniles fledge at about 18 days of age and are independent of the parents when
30–40 days old. The young are sexually mature by their first spring.
In the nonbreeding season, the birds congregate at communal roosts. Eurasian Collared-
Doves live a relatively long time; the oldest bird on record was more than 13 years old.
Impacts. When they occur in large numbers, Eurasian Collared-Doves can deter other
birds from using bird feeders and may even aggressively defend the food source. Large flocks
also become a noise problem and produce large amounts of unsightly droppings. They can
transmit the parasite Trichomonas gallinae to native doves using the same feeders or birdbaths
and to hawks that prey upon them. Potentially, they could damage crops, but such impacts
have not been reported.
Management. Eradication is no longer a possibility. Numbers may be controlled by hunt-
ing. As an introduced species, it is not protected by law; but state and local hunting regula-
tions still apply.
 EUROPEAN STARLING n 237

Selected References
Johnson, Steve A., and Gay Donaldson-Fortier. “Florida’s Introduced Birds: Eurasian Collared-Dove
(Streptopelia decaocto).” Document WEC256, Department of Wildlife Ecology and Conservation,
University of Florida/ Institute of Food and Agricultural Services, 2009. http://edis.ifas.ufl.edu/
uw301.
National Biological Information Infrastrucutre, IUCN French Committee, and IUCN SSC Invasive
Species Specialist Group. “Streptopelia decaocto (bird).” ISSG Global Invasive Species Database,
2008. http://www.issg.org/database/species/ecology.asp?si=1269&fr=1&sts=&lang=EN.
Sibley, David Allen. “Eurasian Collared-Dove.” In National Audubon Society, The Sibley Guide to Birds,
256. New York: Alfred A. Knopf, Inc., 2000.
Sibley, David Allen. “Ringed Turtle-Dove.” In National Audubon Society, The Sibley Guide to Birds, 256.
New York: Alfred A. Knopf, Inc., 2000.

n European Starling
Also known as: Common Starling
Scientific name: Sturnus vulgaris
Family: Sturnidae
Native Range. Europe. In lowlands from the United Kingdom westward across Ukraine
and Central Asia to western China. The Starling is a summer visitor in the northern part of
its range where the climate is continental. Migratory birds winter in the Mediterranean
region in Spain, Portugal, North Africa, and across the Middle East into Pakistan.
Distribution in the United States. The European Starling is common in all of the contigu-
ous 48 states and is also found in Hawai’i. In Alaska, resident populations occur around
Anchorage and the Kenai Peninsula; breeding birds summer in the Yukon River valley.
Description. European Starlings are robin-size birds that at a distance appear to be black.
They have short, squared tails, pointed wings, and long, thin, pointed bills. The feathers of
nonbreeding males and females are glossy black with white tips, giving the bird a spotted
appearance. Bills are grayish-black, and the eye is dark-colored. Legs are a dull red.
During the breeding season, the feathers become iridescent purple and green, and most of
the spots vanish. The male sports long feathers on the breast. The bill of both sexes becomes
yellow when they are in breeding condition, but males have a blue spot at the base of the
beak whereas females have a reddish pink spot. Juvenile birds have uniformly gray-brown
plumage and brownish-black bills.
Full grown Starlings are 8–9 in. (20–23 cm) long, have a wingspan of 12–16 in. (31–
40 cm), and weigh 2–3 oz. (60–90 g). Their calls tend to be harsh squeaks, whistles, and
gurgles, but they are good mimics of the calls of other birds and can imitate other sounds
in the environment as well.
Starlings walk; they do not hop. In flight, they resemble tiny fighter planes, with their tri-
angular wings and short tails. Nonbreeding birds gather in huge flocks and perform spec-
tacular mass aerial displays, especially at dusk as they ready themselves to settle at a
roosting site.
Related or Similar Species. European Starlings often occur in mixed flocks with other
“blackbirds” with which they could be confused. The Common Grackle (Quiscalus quiscala)
is larger, has a more elongated body and a proportionately much longer tail. Its iridescent
feathers are not speckled, and its bill is never yellow. The eye, however, is bright yellow.
The male Brown-headed Cowbird (Molothrus ater) is roughly the same size as a Starling,
238 n VERTEBRATES (BIRDS)

but its head is distinctly brown,

the feathers are not spotted, and
the bill is conical in shape and
always dark brown in color.
Male Brewer’s Blackbirds (Eu-
phagus cyanocephalus) also have
glossy black plumage with a
green or dark-blue sheen in the
breeding season. They are simi-
lar in size to Starlings and have
straight bills, but have bright
yellow eyes.
Introduction History. The
arrival of the European Starling
in North America traces to late
nineteenth-century attempts
by acclimatization societies to
create a familiar landscape
for European immigrants.
The American Acclimatization
Society, under the leadership
of Eugene Scheiffelin, released
80–100 Starlings into New
York’s Central Park in 1890–
1891. Schieffelin’s goal was to
have all the birds mentioned in
the works of William Shakes-
peare brought to the United
States. (He also brought over
the House Sparrow from
England. See Birds, House
Top: the European Starling is native to Europe and western Asia. (Adapted Sparrow.) In Henry IV, Hotspur
from map at http://en.wikipedia.org/wiki/File:Sturnus_vulgaris_map.png.) says, “I’ll have a Starling . . . to
Bottom: The European Starling is found in all 50 states and Puerto Rico. speak nothing but ‘Mortimer.’ ”
(Adapted from “European Starling” [map]. Cornell Lab of Ornithology. Several attempts to intro-
http://allaboutbirds.org/guide/european_starling/id.) duce Starlings were made from
1850 to 1900, but the first suc-
cess came with the release of 60 birds in Central Park in 1890. Fifteen pairs survived. The
following year, another 40 were set free. During the first 10 years, the population was con-
tained in the greater New York City area. Thereafter, European Starlings rapidly increased
their numbers and expanded their range north, south, and west. They reached Alaska by
1970. Within 75 years of introduction, Starlings had dispersed across the entire continent.
Habitat. Starlings are commonly seen in urban areas and other disturbed sites. They feed
on the ground in open areas with short grass, including suburban lawns and city parks.
They also infest feed lots and agricultural areas, where they feed on grains and other crops.
They may fly great distances from roosting sites in tree tops, under bridges, or in other open
structures to their feeding grounds. They are secondary cavity-nesters: they do not excavate
their own nest sites, but occupy those made by other birds, or mammals or humans. They
 EUROPEAN STARLING n 239

A. European Starlings are about the size of a robin but walk, rather than hop. (Steve Byland/Shutterstock.)
B. Nonbreeding birds assemble in huge flocks and perform what appear to be well-coordinated mass aerial dis-
plays. (Vasily A. Ilyinsky/Shutterstock.)

are not fussy about location and will build nests in holes in trees, crevices in buildings, rain
gutters, under the eaves of roofs, in roof vents and attics, and in birdhouses.
Diet. European Starlings are omnivores. They consume a variety of invertebrates, includ-
ing earthworms, beetles, grasshoppers, spiders, and snails, and also feed on seeds, berries,
and fruits such as apples, pears, and cherries. By inserting their bills into the ground or into
food items and then opening their beaks, they can pry fruits open to extract any seeds or
insects inside. Favorite foods reportedly include the berries of poison ivy and Virginia
creeper, blackberries, mulberries, and elderberries.
Life History. Breeding pairs form and begin to select nesting sites in late winter or early
spring. The nests are constructed of dried grasses and other materials and typically fill the
nest cavity. The female lays a clutch of 4–6 blue eggs. She may produce two or three clutches
during the breeding season, which extends into July. The eggs incubate for 15 days, and
hatchlings remain in the nest another 21–23 days. Both parents incubate eggs and feed the
young birds. Fledglings follow their parents and beg for food for several days after leaving
the nest. When they become independent, the juveniles congregate in flocks with other
young Starlings. At the end of the breeding season, the parents again become gregarious
and spend the nonbreeding season feeding and roosting as members of large flocks.
Impacts. Much of the trouble associated with Starlings comes from their being nuisances
due to the fact that they congregate in such large, noisy flocks. In cities, their acidic guano
can coat buildings and sidewalks and corrode statues and the paint on cars. Diseases such
as salmonella and histoplasmosis can proliferate at established roosts. Droppings have the
potential to contaminate animal feed and water sources. Large flocks are able to inflict seri-
ous damage to crops such as grain, grapes, olives, and cherries. Starlings massing near air-
port runways pose a real danger to arriving and departing airplanes because they can clog
engines, damage planes, and, when an entire flock collides with a plane, can cause planes
to crash.
The ecological impacts of the European Starling in the United States are related to their
aggressive takeovers of nest cavities sought or occupied by other bird species. Declines of
woodpeckers, martins, tree swallows, and bluebirds have been blamed on the presence of
Starlings. Starlings are also implicated in the spread of the seeds of exotic weeds.
240 n VERTEBRATES (BIRDS)

The European Starling has also been introduced to South Africa, Australia, and New
Zealand. It has been nominated by the Global Invasive Species Programme as one of the
100 “World’s Worst” invaders.
Management. Direct population reductions may be accomplished through poisoning
with Starlicide Complete, trapping, or shooting, but most efforts try to repel or exclude
Starlings. At airports, livestock facilities, and some urban roosting sites, frightening the birds
with noises, including recordings of their own distress calls, has had at least temporary suc-
cess. Barricading entrances to cavities with screens or other covers will eliminate nesting
sites. Habitat modification involving the removal of water sources and foraging sites has
had some success at airports.

Selected References
Adeney, Jennifer Marion. “European Starling (Sturnus vulgaris).” Introduced Species Summary Project,
Columbia University, 2001. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/
inv_spp_summ/Sturnus_vulgaris.html.
Johnson, Steve A., and Walter Givens. “Florida’s Introduced Birds: European Starling (Sturnus vulga-
ris).” Document WEC255, Department of Wildlife Ecology and Conservation, University of
Florida/IFAS, 2009. http://edis.ifas.ufl.edu/UW300.
Withers, David Ian. “Origins of the European Starling in the United States.” Tennessee Department of
Environment and Conservation, 2000. http://www.state.tn.us/environment/tn_consv/archive/
starlings.htm.

n House Finch
Also known as: Linnet
Scientific name: Carpodacus mexicanus
Family: Fringillidae
Native Range. Western North America. Prior to the late nineteenth century, House
Finches were found in the southwestern states. They expanded their range northward and
were first reported in the Columbian Basin of Oregon in 1885. Dam construction and irriga-
tion projects apparently facilitated their spread into eastern Washington State in the early
twentieth century; they were found in the western part of the state by the 1950s. Today, nat-
urally occurring populations inhabit the drier habitats west of the Rocky Mountains from
southern British Columbia to southern Mexico. Originally occupying undisturbed desert
scrub, desert grasslands, chaparral, oak savannas, and low elevation open coniferous forests,
they adapted to human-dominated environments and moved into suburban and agricultural
areas. They also crossed the Rockies onto the High Plains.
Distribution in the United States. House Finches are now common birds throughout the
eastern United States and have also been introduced into Hawai’i.
Description. House Finches are small, sparrow-like songbirds. Adults are 5–6 in. (13–
14 cm) long and have wingspans of 8–10 in. (20–25 cm). The male usually has a rosy-red
forehead, stripe over the eye, breast, and rump. The brown back has dark brown streaks,
and light-brown streaks appear on the flanks and belly. Some males are orange or even yel-
low; plumage color depends on diet. Females are brown with a plain brown head, finely
streaked underparts, and two pale wingbars. They have no eye stripe. Both sexes have squar-
ish, slightly notched brown tails and blunt, rounded beaks. Juveniles look like females;
males acquire their color in their second spring.
 HOUSE FINCH n 241

House Finches have a

melodic warbling song with a
few harsh notes and a down-
ward trend.
Related or Similar Species.
The most similar bird in the
eastern United States is a conge-
ner, the Purple Finch (Carpo-
dacus purpureus). Purple Finches
are somewhat more robust.
Males have more red on their
heads and breast than House
Finches and lack brown streaks
on their flanks and bellies.
Females have broad white stripes
over and under the eye, a larger
beak, and bolder striping on
breast and belly than female
House Finches. Female House
Sparrows (see Birds, House
Sparrow) could also be mistaken
for female House Finches. The
former have light brown stripes
on the back and unstreaked
undersides. Pine Siskins (Cardu-
elis pinus) are considerably small-
er and more heavily streaked
than female House Finches; they
have yellow patches on the wings
(not always obvious) and slen-
der, pointed bills.
Introduction History. House Top: The House Finch is native to the western United States, where its
Finches were introduced into range has been expanding since the 1960s. Bottom: The House Finch is
Hawai’i in the 1880s. Finches a native transplant and now a common songbird in the eastern half of
in the eastern United States the United States. It has also been introduced into Hawai’i. (Both maps
trace their origins to the 1930s, adapted from All About Birds, 2009.)
when House Finches were sold
in pet stores in New York City as “Hollywood Finches.” The selling of wild birds became ille-
gal in 1940, and some dealers apparently released their captive finches on Long Island to
avoid legal action. For the first few years, the population’s survival was tenuous, but by
the 1950s, House Finches were established in New York City. The population then irrupted,
and finches spread rapidly up and down the East Coast, probably helped by the increasing
presence of bird feeders in suburban and urban areas of the United States. By the 1980s
House Finches had reached the Mississippi River, and today the eastern and western distri-
bution areas seem to have merged; House Finches are now found across the continent.
The population explosion ceased in the early 1990s with appearance of mycoplasmal
conjunctivitis. This infection causes the eyelids to become red, swollen, and encrusted and
can lead to blindness and starvation. The disease first appeared in the Washington, D.C.,
242 n VERTEBRATES (BIRDS)

A. The male House Finch has a rosy-colored forehead and light brown streaks on the flanks and belly.
(Stubblefield Photography/Shutterstock.) B. The female House Finch has a plain brown head and finely streaked
underparts. There are two pale wing bars. (Chris Hill/Shutterstock.)

area during the winter of 1993–1994. Two years later, it had spread across the
Appalachians, and by the following winter, it was in the Midwest and beginning to show
up on the Great Plains. The disease is most prevalent in winter when the birds flock at feed-
ers. By 1999, scientists were reporting that House Finch numbers were roughly 40 percent
less than expected before the epidemic began. The rapid decline in House Finch populations
coincided with an increase in the number of House Sparrows in some affected areas. The vir-
ulence of the disease may be abating. Current estimates are that only 5–10 percent of the
population is infected, whereas at the peak of the disease, 50 percent of the eastern finch
population may have been died from it. (In 2004, the disease appeared in the Pacific
Northwest and has reached epidemic proportions there.)
Habitat. Where they have been introduced, House Finches inhabit backyards, urban parks,
farmland, and forest edge. They congregate at bird feeders and like to perch in high trees close
by. Nesting sites include ledges on buildings, shrubs, debris piles, hollows in trees, and birdhouses.
Diet. House Finches consume almost any type of seed and also eat ripening fruits, flow-
ers, and buds. In addition to bird seed at feeders, where sunflower seeds are preferred, the
finches also extract seeds from thistle, dandelion, and mistletoe and forage on the ground.
The males acquire their rosy color after the postnuptial molt by eating carotenoid-rich red
berries and fruits. Cherries and mulberries are among those preferred.
Life History. Pair formation begins in late winter. Females tend to choose the males with the
brightest coloration. The breeding season starts in early spring when the female constructs a
small cuplike nest of fine grasses, hair, or other fiber. She lays 3–6 small, pale blue eggs that
are flecked with black on the larger end. The female incubates the eggs for 12–14 days, while
the male feeds and guards her. The nestlings fledge when 11–19 days old. The male will feed
the fledglings for several days until the female has built a new nest. Most House Finches produce
at least two clutches during the breeding season. The young of the year disperse widely and con-
gregate at food sources. Nonbreeding adults and juveniles are especially gregarious in fall and
winter and may form large, mobile foraging flocks. Some in the northeastern United States
migrate in winter. House Finches are mature at one year of age and may live more than 10 years.
Impacts. Most people welcome and encourage these colorful little songsters at their back-
yard feeders. However, as the House Finch spread through the eastern United States,
 HOUSE SPARROW n 243

dramatic declines in Purple Finches and House Sparrows took place in newly invaded areas.
The range of the Purple Finch has shifted northward, confining it more and more to the
conifer forests avoided by the House Finch. Part of this range shift may be due to climatic
change and not solely the influence of increasing House Finch populations.
House Sparrow numbers declined wherever House Finches invaded, suggesting the latter
were outcompeting the former. (Later, when finch numbers decreased, the sparrows
rebounded, reinforcing this interpretation.) Recent research has also found evidence of an
evolutionary response among House Sparrow populations, a phenomenon known as charac-
ter displacement. At bird feeders, the sparrows prefer smaller seeds such as millets, whereas
the finches prefer sunflower seeds. Where the two species both occur, sample measurements
indicate that House Sparrow beak depths are smaller than before the finch invasion and they
are becoming better adapted to specialize on the small seeds neglected by finches.
Since 1993, a bacterial eye infection (mycoplasmal conjunctivitis) has swept through
eastern House Finch populations. There is a possibility that this disease could be spread at
feeders to other wild birds, such as Blue Jays (Cyanocitta cristata) and American
Goldfinches (Carduelis tristis), as well as to domestic poultry.
Management. Since most people like having these birds around, management is largely
directed at reducing the infection rate of conjunctivitis among House Finches. This can be
accomplished by homeowners cleaning bird feeders and disposing of old seed and accumu-
lated bird droppings. For those who find House Finches to be a nuisance, fruits can be pro-
tected from finch depredation with plastic netting, and potential nesting sites can be eliminated.

Selected References
Cooper, Caren, Wesley Hochachka, and André Dhondt. “Why Did House Sparrow Numbers Rise,
Then Fall?” Birdscope 21(2): 2007. http://www.birds.cornell.edu/Publications/Birdscope/
Spring2007/sparrow_numbers.html.
Dewey, T., K. Kirschbaum, and J. Pappas. “Carpodacus mexicanus,” Animal Diversity Web, University
of Michigan Museum of Zoology, 2002. http://animaldiversity.ummz.umich.edu/site/accounts/
information/Carpodacus_mexicanus.html.
“House Finch.” All About Birds, Cornell Lab of Ornithology, 2009. http://www.allaboutbirds.org/
guide/house_finch/id.
Johnson, Steve A., and Jill Sox. 2009. “Florida’s Introduced Birds: House Finch (Carpodacus mexica-
nus).” Document WEC 253, Department of Wildlife Ecology and Conservation, University of
Florida/IFAS. http://edis.ifas.ufl.edu/uw298.
Kammermeier, L. “Population Dynamics of the House Finch.” Birdscope, 13(2): 15, 1999.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Carpodacus mexicanus (bird).” ISSG Global Invasive Species Database, 2005. http://
www.invasivespecies.net/database/species/ecology.asp?si=485&fr=1&sts.
Wootton, J. Timothy. “Ecology and Evolution of Invading and Restored Species.” Department of
Ecology and Evolution, University of Chicago, n.d. http://woottonlab.uchicago.edu/index/
invasive-species/ecology-and-evolution-of-invasive-and-restored-species.

n House Sparrow
Also known as: English Sparrow
Scientific name: Passer domesticus
Family: Passeridae
Native Range. Eurasia and North Africa. Occurs from the United Kingdom eastward
through Siberia and southeastward to the Arabian Peninsula and Indian subcontinent, but
244 n VERTEBRATES (BIRDS)

is not native to Italy. In North

Africa, it is found in coastal
areas and at isolated oases in
the Sahara. They have been
associated with human settle-
ments for centuries.
Distribution in the United
States. Established in all 48 of
the contiguous states and in
Hawai’i and Puerto Rico.
Description. This Old World
sparrow is a small brown bird
common in towns and cities. It
has a relatively large head and
short wings. The legs are pink.
The male has a black bib and
mask, gray cap and rump, and
chestnut-colored nape. The side
of the head and underparts are
pale gray, and the brown back
is streaked with black. The
conical bill is black in summer
and yellowish in winter. Non-
breeding and immature males
have less black on the throat
and breast. The female is brown
all over, with tawny streaks on
her back and a strong buff-
colored eye line edged below
by a dark brown stripe. Her
throat and breast are light and
Top: The native range of the House Sparrow includes all of Europe, much unstreaked. The bill is yellow.
of Asia, and parts of North Africa. (Adapted from map at http:// Young birds resemble females.
en.wikipedia.org/wiki/File:PasserDomesticusDistribution.png.) Bottom: House Sparrows are 5.25–
The House Sparrow is a common bird of the built environment in all 48 6.25 in. (133–159 mm) long.
contiguous states, Hawai’i, and Puerto Rico. In the relatively short time they
have been naturalized in the
United States, they have evolved in accord with regional climatic differences across North
America. Northern birds are larger than southern ones; desert populations have much
lighter coloration than populations in the humid east or Pacific Northwest.
The song of the House Sparrow is a series of chirps.
Related or Similar Species. Only one other Old World sparrow has become established
in the United States: the European Tree Sparrow (Passer montanus), restricted to St. Louis,
Missouri, and nearby Illinois. It can be distinguished by its rufous crown, white collar,
and prominent black patch in the white cheek.
Introduction History. Some early introductions of the House Sparrow (then called the
English Sparrow) were attempts to control insect pests, especially the elm spanworm
(Ennomos subsignaria), a defoliator of shade trees. Many introductions were the results of
 HOUSE SPARROW n 245

A. The male House Sparrow has a black bib and mask. (Francis Boose/Shutterstock.) B. The female House
Sparrow is brown with tawny streaks on the back and a strong buff-colored eye line. (Stubblefield
Photography/Shutterstock.)

activities of acclimatization societies; and at first, citizens tenderly cared for the birds. The
first introduction into the United States consisted of eight pairs from England released in
Brooklyn, New York, in 1851. More birds were imported the following year, and 50 of them
were released. Neither group thrived. In 1853, birds held and nurtured in a tower in the
Greenwood Cemetery were released and protected. These sparrows did survive; they multi-
plied, and people carried them to other towns and cities in the region. Other deliberate
introductions, commonly by acclimatization societies, include those in Portland, Maine
(1854), Peace Dale, Rhode Island (1858), Boston, Massachusetts (1858), and New Haven,
Connecticut (1867). House Sparrows were brought to San Francisco, California, in 1875.
The birds prospered in part by eating seeds in the dung of urban horses. They rapidly dis-
persed along carriage roads into agricultural areas, where they became pests. House
Sparrows are now found in almost all populated parts of the country. Numbers are believed
to have peaked in the early twentieth century before motorized vehicles replaced horses.
Another period of decline followed the conversion to modern industrial agriculture in the
1960s.
Habitat. These birds are creatures of human-modified environments. They are found in
cities, suburbs, and agricultural settings, but not in undisturbed forests, grasslands, or
deserts. By preference cavity nesters, they build their nests in crevices in and on buildings,
in the hollow posts of traffic signs, in the rafters of porches, in birdhouses, in holes in trees,
or in any other sheltering location.
Diet. House Sparrows primarily forage on the ground for seeds, but also feed on fruits,
insects, and garbage. It is not unusual for them to extract dead insects from the grills of auto-
mobiles and to hop about the parking lots of fast food restaurants looking for discarded
food.
Life History. House Sparrows begin to nest as early as February, when monogamous
pairs form. The nests, usually in small colonies, consist of dried plant material, feathers,
string, and shreds of paper. The female usually lays 3–6 speckled pale blue eggs per clutch
and may produce up to four clutches a year. Incubation starts after the last egg in a clutch
has been laid and continues for 10–12 days. The young fledge about 14 days after hatching.
They may live more than 12 years.
Impacts. Today, the House Sparrow is considered mainly a nuisance species in cities and
towns when large groups create noise and messy, corrosive droppings. In the early years of
246 n VERTEBRATES (BIRDS)

their invasion of the United States, they were an agricultural pest, eating ripening grains,
destroying fruits, and consuming or fouling seeds and other feed. To some extent, the neg-
ative impacts were balanced by the birds’ consumption of insect pests, including cabbage
worms and cotton caterpillars. House Sparrows will evict native birds from their nest cavities
and were implicated in declines of the Eastern Bluebird (Sialis sialis), Purple Martin (Progne
subis), and some woodpeckers during the twentieth century. Sparrow populations have
declined since the advent of large monoculture farms in the 1960s. Declining numbers of
sparrows in the central and eastern United States and recovery programs targeting bluebirds
and martins have mitigated their effects on native birds.
Management. Habitat modifications that eliminate roosting and nesting sites can reduce
local House Sparrow populations. So, too, can reduction of food sources by cleaning up gar-
bage dumps and protecting berries and other small crops with bird netting. The birds can be
repelled with noise and scarecrows. Since these nonnatives are not protected by the
Migratory Bird Act, they may be trapped or shot.

Selected References
“House Sparrow.” All about Birds, Cornell Lab of Ornithology, 2009. http://www.allaboutbirds.org/
guide/House_sparrow/id.
Laycock, George. The Alien Animals. The Story of Imported Wildlife. New York: Ballantine Books,
1966.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Passer domesticus (bird).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?fr=1&si=420.
Roof, J. “Passer domesticus.” Animal Diversity Web, University of Michigan Museum of Zoology, 2001.
http://animaldiversity.ummz.umich.edu/site/accounts/information/Passer_domesticus.html.

n Japanese White-Eye
Also known as: Me-jiro, Dark-green White-eye
Scientific name: Zosterops japonicus
Family: Zosteropidae
Native Range. Eastern Asia. Native to Japan, eastern and southern China, Vietnam to
Burma in Southeast Asia, Taiwan, Hainan Island, Ryukyu Islands, and the northern
Philippines.
Distribution in the United States. Established on all the major islands in Hawai’i, where
it is now the state’s most abundant bird.
Description. This extremely active and acrobatic little bird has an olive-green head, neck,
and back. The white eye-ring appears almost to be embroidery. The throat and underside of
the tail are yellow, the breast gray, and the belly dull white. Flanks are brownish. The wings
and tail are dark brown; upper tail feathers and wing feathers are outlined in green. Legs and
feet are black. They measure 4–4.5 in. (10–12 cm) long. Juveniles lack the white eye-ring.
White-eyes forage in trees in small flocks and often hang upside to down to reach food.
Introduction History. The Japanese White-eye was first introduced to O’ahu in 1929 by
the (Hawaiian) Territorial Board of Agriculture and Forestry to control insects. It was taken
to the island of Hawai’i in 1937 and now can be found on all the major islands.
 JAPANESE WHITE-EYE n 247

Habitat. In Hawai’i, the

Japanese White-eye lives in
trees and shrubs in rainforest,
open deciduous forests, agricul-
tural areas, towns, and city
parks. It occurs from sea level
to treeline in both arid and
humid parts of the islands.
Diet. Japanese White-eyes
probe foliage at all levels for
insects, including beetles, fly
larvae, and spiders. They also
eat seeds, nectar, and fruit.
They switch from one food
source to another depending
on availability.
Life History. The breeding
season extends from February to
December, but there is a distinct
peak in activity in July and
August. Neat cuplike nests are
built at various levels in trees.
They are constructed of grass,
string, cobwebs, leaves, and
mosses and attached to the fork
of a branch with spider webs. If
human habitation is nearby, the
nest is often lined with human
hair. The female lays 3–4 white
eggs that hatch after an 11-day
incubation period. Newborn
chicks are altricial; their eyes Top: The Japanese White-eye is native to East Asia. (Adapted from
open about five days after hatch- MacKinnon, J., and K. Phillips, A Field Guide to the Birds of China.
ing. The young fledge 10–12 days Oxford: Oxford University Press, 2000.) Bottom: The Japanese White-
after hatching. Often the young eye is invasive only in Hawai’i, where it has become the state’s most abun-
cannot fly when they first leave dant bird.
the nest, but they acquire this
ability in 1–6 days. Fledglings remain with their parents for 15–20 days, at which point the
adults begin to build a new nest and force the juveniles out of the breeding territory. The white
eye-ring is apparent at 23 days of age; and by 30 days, the young look like adults. Young birds
gather in flocks until the next year, when they will begin to form pairs and breed.
Impacts. The Japanese White-eye, unlike many exotic birds in Hawai’i, has invaded
native montane forest and is currently the most abundant land bird on the main islands. It
appears to compete directly with some endemic Hawaiian Honeycreepers for food. As a gen-
eralist, the White-eye can switch its feeding specialty to those items most prevalent in a vari-
ety of forest types or to less desirable insects, fruits, and flowers when preferred food sources
become depleted. In wet forests, it competes with the Elepaio (Chasiempis sandwichensis) for
foliage insects on low branches and the Apanane (Himatione sanguinea) for nectar. In high-
248 n VERTEBRATES (BIRDS)

altitude forests, it competes

with the Iiwi (Vestiaria coccinea)
for nectar. Evidence suggests
that the presence of White-eyes
leads to undernourishment of
the young of some native birds
such as the endangered Akepa
(Loxops coccineus), resulting in
stunted adults more vulnerable
to diseases such as avian malaria,
higher infestations of chewing
lice, higher mortality rates, and
declining populations. It may
have lesser impact on the
Common Amakihi (Hemignathus
virens), which has a similarly
broad niche.
The Japanese White-eye is an active and acrobatic little bird that sports a
The Japanese White-eye has
distinct white eye-ring. (K. W. Bridges, “Common Campus Birds,”
University of Hawai’i at Manoa. http://www.botany.hawaii.edu/biol- the potential to spread invasive
ogy101/birds/japanese_white_eye.htm.) plants into undisturbed native
Hawaiian forests. It is known to
disperse, for example, the velvet tree (Miconia calvescens; see Volume 2, Trees, Velvet Tree),
lantana shrubs (Lantana camara; see Volume 2, Shrubs, Lantana), and the fire tree (Myrica faya;
see Volume 2, Trees, Fire Tree). On a positive note, the Japanese White-eye may be replacing
extinct species of honeycreeper that dispersed native shrubs.
Management. There appears to be no effort to control the spread of these birds, and in-
deed, it is unlikely any measure would be effective.

Selected References
Foster, Jeffrey T., and Scott K. Robinson. “Introduced Birds and the Fate of Hawaiian Rainforests.”
Conservation Biology 21(5): 1248–57, 2007.
“Introduced Japanese White-Eyes Pose Major Threat to Hawaii’s Native and Endangered Birds.”
ScienceDaily, 2009. http://www.sciencedaily.com/releases/2009/09/090917131540.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Zosterops japonicus (Bird).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?si=954&fr=1&sts.
“Zosterops japonicus.” Senior Seminar: Introduced Species in Hawaii. Department of Biology, Earlham
College, 2002.

n Monk Parakeet
Also known as: Quaker parrot, Quaker conure
Scientific name: Myiopsitta monachus
Family: Psittidae
Native Range. South America. Four subspecies are recognized, three in the lowlands east
of the Andes: M. m. monachus from southeastern Brazil (Rio Grande do Sul) into Uruguay
and northeastern Argentina; M. m. cotorra from eastern Bolivia south through Paraguay;
 MONK PARAKEET n 249

and M. m. calita in the Pata-

gonian region of Argentina. The
disjunct upland population in
Bolivia, currently recognized as
the subspecies M.m. luchsi, is
probably a separate species as
evidenced by its different habitat,
appearance, and nesting behav-
ior. Genetic evidence traces the
origins of the Monk Parakeets in
the United States to the region
from Entre Rios, Argentina, to
Rio Grande do Sul, Brazil, near
the Uruguayan border. Other
records confirm that most ani-
mals trapped for the pet trade
were exported from eastern
Argentina and Uruguay.
Distribution in the United
States. Populations come and
go, but the bird appears to be
established in Alabama, Conn-
ecticut, Delaware, Illinois,
Florida, Louisiana, New Jersey,
New York, Oregon, Rhode
Island, Texas, and Virginia;
and perhaps also in Colorado,
Missouri, Ohio, and South
Carolina. The largest popula-
tions are in Florida and Con-
necticut, and in both states, they
are increasing in numbers and Top: The Monk Parakeet is native to temperate lowlands east of the Andes
expanding their ranges. Monk Mountains in South America. (Adapted from Russello et al. 2008.)
Parakeets were eradicated from Bottom: Monk Parakeets are established in several urbanized locations
California in the 1970s. Monk across the United States. (Adapted from Johnson and Logue 2009.)
Parakeets have been observed
in Hawai’i, but have not become established. They are well established in Puerto Rico.
Description. The Monk Parakeet is a small, stocky, mostly green parrot about 11.5 in.
(30 cm) from head to tail. The wingspan is about 19 in. (53 cm), and the weight is 2–3 oz.
(90–120 g). The forehead, throat, and breast are gray with white barring. The lower abdo-
men and vent areas are yellow, and the flight feathers are dark blue. The eyes are brown,
the bill pale yellow or orange, and the legs are gray. Immatures have green foreheads.
Monks are highly social and noisy; they chatter continually at the nest. They possess a
variety of calls and squawks and are especially loud when flying.
Monk Parakeets weave large communal nests of sticks, usually 30 ft. (10 m) or more
above the ground. They utilize trees, power poles and towers, and the nests of other birds
as supports for these constructions, which may become 3 ft. (1 m) in diameter and weigh
up to 440 lbs. (200 kg). Each pair has its own nest cavity within the structure, the entrance
250 n VERTEBRATES (BIRDS)

pointing downward. The nest is

used for breeding and for shel-
ter year round, and is probably
a major reason that the Monk
Parakeet has been able to sur-
vive in northern states. It is the
only parrot that builds a stick
nest; others species are cavity-
nesters.
Related or Similar Species.
Three other parakeets intro-
duced to the United States are
about the same size and might
be confused for Monk Para-
keets. The White-winged Para-
keet (Brotogeris versicolurus)
and the Yellow-chevroned
The Monk Parakeet is a stocky, mostly green parrot about a foot long. Parakeet (B. chiriri) are both
They are highly social. (Steve Baldwin, BrooklynParrots.com.) smaller (8.75 in. or 222.5 mm
total length) than monks and
have green heads and bodies with yellow patches showing on their folded wings. In flight,
both display yellow bands on the underwing (i.e., the greater coverts are yellow), which
are lacking in the Monk Parakeet. The Nanday Conure or Black-hooded Parakeet
(Nandayus nenday) has a black head and bluish breast. Budgerigars (Melopsittacus undulatus),
a fourth exotic parrot, are much smaller (7 in. or 178 mm total length) than Monk Parakeets
and lack the gray foreheads and throats. Their long pointed tails and yellow wing stripe dis-
tinguish them in flight.
Introduction History. From the late 1960s into the early 1970s, some 64,000 Monk
Parakeets were imported into the United States. This intelligent species ranks as one of the
10 best talkers among parrots, and its cost is reasonable. Birds were exported from
Argentina, perhaps because they were considered agricultural pests in that country. Their
importation into the United States was not banned until the Wild Bird Conservation Act of
1992, but these popular cage birds continue to be bred by aviculturalists here. Releases,
intentional or not, continue to replenish feral populations and start new ones.
Feral populations had become established in at least 21 urban locations in seven states by
the early 1970s. All stemmed from accidental or intentional releases by zoos, the pet indus-
try, or pet owners. The concern over the species becoming an agricultural pest led the U.S.
Fish and Wildlife Service between 1970 and 1975 to capture or shoot free-roaming parrots
and try to eradicate feral populations. They eliminated monks from California and reduced
the overall number of populations to seven in five states. Nonetheless, the number of and
size of populations grew after 1975. By 1995, the bird could be found at 76 sites in 15 states.
Monk Parakeets were first reported from Florida in 1972. They were widespread by 1992
and today may number 50,000 to 150,000 birds in 16 counties. The urbanization of eastern
Florida with the planting of thousands of ornamental plants that together produced fruit and
nectar year round provided an optimal habitat for the bird. Birdseed at feeders supple-
mented the diet, particularly in winter.
Monk Parakeets in New Jersey are believed to be descendents of birds released in New
York in the 1960s. In 1968, parakeets escaped from a damaged crate at John F. Kennedy
 MONK PARAKEET n 251

Carolina Parakeet

T he eastern United States was once home to North America’s only native para-
keet, the Carolina Parakeet (Conuropsis carolinensis), and some have suggested
that the Monk Parakeet might come to fill its empty niche—and, like it, become an
agricultural pest. Well into the nineteenth century, the colorful bird was an abun-
dant resident of sycamore and bald cypress bottomland forests from the Gulf coast
northward almost to the Great Lakes. Like most parrots, it was highly social, noisy,
and fed on a variety of plant foods, including cockleburs, sandspurs, pine seeds,
and bald cypress seeds. It also foraged on grain and orchard crops.
The Carolina Parakeet was up to 13 in. (33 cm) long from the head to the tip of
the tail. Its body was bright green, and its head yellow. The forehead and lores
were bright orange. Unlike Monks, Carolina Parakeets nested in tree cavities. It
became popular in the early pet trade because of its colorful plumage, but was a
poor talker. Its bright feathers were also in demand for a time for women’s hats.
The birds had the unfortunate habit of flocking to trapped or injured parakeets
in response to their distress calls, and this made it relatively easy for farmers or
hunters to shoot to trap large numbers at a time.
John James Audubon noted a rapid decline in the number of parakeets in 1831.
The last confirmed sightings in the wild were made in 1904 at Lake Okeechobee in
Florida by the respected ornithologist Frank Chapman. The last known captive bird
died in 1918 at the Cincinnati Zoo, and the Carolina Parakeet was officially
declared extinct in 1939.
The demise of the Carolina Parakeet was likely due to a combination of factors.
Slaughter by farmers trying to protect crops usually is identified as the cause of
extinction, but the birds were also trapped for food, feathers, and pets.
Destruction of their bottomland habitat was undoubtedly a contributing factor.
Other possibilities include introduced diseases—especially those originating in
poultry—and competition for nesting and roosting cavities by feral honey bees, a
species introduced to North America during the colonial period.

Airport. The first parakeet was recorded from Oregon in 1977. In 1980–1981 a nest was
built near the Portland International Airport, and a small population has persisted since that
time.
Habitat. In their native lands, Monk Parakeets prefer open country with tall, isolated
trees for nesting. They are found in savannas and woodlands, and in farmland, open euca-
lyptus forest, and palm groves. The widespread planting of eucalyptus trees as windbreaks
in the pampas allowed them to expand their range onto the grasslands. In the United
States, all populations are urban.
Diet. Monk Parakeets are largely granivorous and consume primarily the seeds of wild
grasses, sedges, and sunflowers. They also feed on berries, fruits, and occasionally insects.
In agricultural settings, they forage on domesticated grasses such as maize, wheat, rice,
and sorghum and are considered major agricultural pests in South America. In the cities of
North America, they feed on the buds, seeds, and fruits of ornamental plants and, in winter,
almost exclusively on birdseed put out for them on feeders.
Life History. Breeding begins in North America in spring as the photoperiod increases.
Clutches contain 4–7 eggs. Eggs hatch asynchronously, beginning about 20 days after the
252 n VERTEBRATES (BIRDS)

first one was laid. The nestlings are initially covered with yellow down and remain in the
nest for about 40 days, fed by the parents. Fledglings rarely move far from their birthplace,
and often add their own chambers to the communal nest. Immature birds become sexually
mature at age 2, but may not breed. Instead, they stay with their parents and help maintain
the nest and care for the next generation of their siblings.
Impacts. In the United States, the Monk Parakeet is not the agricultural pest it was pre-
dicted to become in the 1970s. All populations are, to date, urban. There is little or no evidence
suggesting they affect native wildlife. Instead they seem to have brightened the lives of many
urban dwellers who delight at seeing these colorful birds at their winter bird feeders. The large
nests do pose a problem on transmission line towers and distribution poles, which can be
damaged. When they are wet, the nests can cause short circuits and disruption of electric
power service. It is costly and can be dangerous to remove nests and repair poles and towers.
Monk Parakeets can also be a nuisance because of the noise they make.
Management. Control of monk parakeets is difficult. The U.S. Fish and Wildlife Service’s
efforts, which came early in the history of introductions of the birds to the United States,
only resulted in eliminating the monks in California. Destruction of nests just causes the
birds to build new ones, often nearby. Controlling population sizes may be achieved by lim-
iting accessible food resources, for example, by removing bird feeders or not growing fruit-
bearing ornamental plants. Trapping and selling the parakeets might be an option, since
there remains a strong market for the birds. The issue of management is complicated by
the fondness many people have for these birds and the political strength of animal rights
organizations. Since any eradication program would be labor intensive, costly, and contro-
versial, and since the birds are not currently an ecological problem, many in wildlife man-
agement think the effort to reduce populations simply is not worth the trouble.

Selected References
Campbell, T. S. “The Monk Parakeet, Myiopsitta monachus.” Institute for Biological Invasions,
University of Tennessee, Knoxville. Invader of the Month, 2000.
Gluzberg, Yekaterina. “Monk Parakeet (Myiopsitta monachus).” Introduced Species Summary Project,
Columbia University, 2001. http://www.columbiauniversity.org/itc/cerc/danoff-burg/invasion
_bio/inv_spp_summ/Myiopsitta_monachus2.html.
Johnson, Steve A., and Sam Logue. “Florida’s Introduced Birds: Monk Parakeet (Myiopsitta monachus).”
Document #WEC257, Department of Wildlife Ecology and Conservation, University of Florida/
IFAS, 2009. http://edis.ifas.ufl.edu/UW302.
Russello, Michael A., Michael L. Avery, and Timothy F. Wright. “Genetic Evidence Links Invasive
Monk Parakeet Populations in the United States to the International Pet Trade.” BMC Evolutionary
Biology 8: 217, 2008.
Stafford, Terri. “Pest Risk Assessment for the Monk Parakeet in Oregon.” Oregon Department of
Agriculture, 2003. http://www.oregon.gov/OISC/docs/pdf/monkpara.pdf.

n Mute Swan
Scientific name: Cygnus olor
Family: Anatidae
Native Range. Eurasia. The Mute Swan breeds in temperate areas from Northwest Europe
to Russia, Ukraine, and Kazakhstan. It occurs south of the taiga in southern Siberia and
northern China. Wintering grounds are primarily in subtropical regions that lie south of
 MUTE SWAN n 253

nesting areas. These areas

include the eastern Mediterra-
nean, the Black and Caspian
seas region, the Persian Gulf
region, Central Asia, and the
Yellow Sea coast.
Wild populations in Western
Europe were virtually eliminated
by overhunting from the thir-
teenth through the nineteenth
centuries. Large landowners pre-
served groups of semidomesti-
cated birds on their estates; so
most populations today descend
from feral swans. Conservation
efforts in the late nineteenth
century allowed Mute Swans
to regain much of the former
territory of their wild ancestors.
Birds in the United Kingdom
and Western Europe are semi-
migratory, in winter moving only
short distances, often to the coast,
where they congregate in flocks.
During severe winters, when
lakes and rivers freeze over, they
will fly much longer distances to
open water.
Distribution in the United
States. Mute Swans occur
in the eastern United States
from southern Maine to South Top: The Mute Swan was once widely distributed in the temperate
Carolina and from the Atlantic regions of Eurasia, but populations were largely extirpated by
coast inland to the Mississippi overhunting from the thirteenth through the nineteenth centuries. Many
River drainage. Small numbers populations existing today derive from semidomesticated stock.
are found in western states. (Adapted from Delany 2006.) Bottom: Mute Swans have established
Mute swans are considered populations in southern New England, Long Island Sound, Chesapeake
Bay, and Great Lakes areas. Small numbers occur in some western
established in southern New
states. (Adapted from “Winter Christmas Bird Count” [map]. http://
England, Long Island Sound, www.discoverlife.org/mp/20q?search=Cygnus+olor.)
the Chesapeake Bay, and the
Great Lakes region. The largest
populations are in Rhode Island, Connecticut, New York, New Jersey, and Maryland. They
are not truly migratory in any part of their adopted range.
Description. Mute Swans are very large white waterfowl with long, gracefully curved
necks. When swimming, the bill points downward. The Mute Swan has an orange bill with
a diagnostic black knob at the base and a black nail at the tip. The lores are also black. Adults
may have a total length greater than 5 ft. (1.3–1.6 m) and weigh 20–25 lbs. (8–11 kg). The
wingspan can be 7 ft. (1.8–2.5 m). Males (“cobs”) are larger than females (“pens”).
254 n VERTEBRATES (BIRDS)

Immature swans (cygnets)

have two morphs, greyish
brown or white. The gray cyg-
nets have slate-gray bills, legs,
and feet. The white morph has
a tan bill and pinkish tan feet.
Both lack the basal knob on
the bill that characterizes
adults.
Related or Similar Species.
Native swans have long necks
that are held straight when
swimming, and black bills. The
Mute swans have long, gracefully curved necks. When swimming, they Trumpeter Swan (Cygnus bucci-
point their bills downward. (Ozerov Alexander/Shutterstock.) nator) is equal in size to the
Mute Swan. Its range is gener-
ally north of that of the Mute
Swan. The Tundra or Whistling Swan (Cygnus columbianus) is considerably smaller and
has yellow lores. Its distribution overlaps with that of the Mute Swan on its wintering
grounds and during migration. The white Snow Goose (Chen caerulescens) might also be
mistaken for a swan on the wintering grounds, but it is a much stockier bird with a shorter
neck and is about half the size of a swan. Its black wing tips are diagnostic in flight.
Introduction History. By most accounts, Mute Swans were introduced into the United
States from Europe in the late 1800s as ornamental birds on private estates, in urban parks,
and in zoos. Aviculturalists bred the birds, esteemed for their beauty and grace. Originally,
swans had their flight feathers cut to keep them from flying off. However, some owners
neglected to pinion their birds, and escapes established feral populations. The earliest
known free-ranging populations lived along the Hudson River (1910) and on Long Island,
New York (1912). These populations expanded their numbers and distribution within the
states of the Atlantic Flyway. New Jersey had feral Mute Swans by 1919, Rhode Island by
1923, and Maryland by 1954. Mute Swans began to colonize the lower Great Lakes in the
mid-1960s and were first sighted in South Carolina in 1993. Many populations in the
eastern United States grew especially rapidly in the 1990s, but have shown signs of decline
since that time. Mute Swans in the California and the Pacific Northwest are the products
of local releases and escapes.
There is no credible evidence to the support the hypothesis that Mute Swans were native
to North America and suffered continent-wide extinction during the Pleistocene Epoch. Nor
is there scientific support for the belief that Mute Swans regularly migrated from Siberia to
North America. Furthermore, none of the great early American naturalists and wildlife
artists, such as Mark Catesby, John James Audubon, Spencer Baird, or Elliot Coues, encoun-
tered these large birds.
Habitat. Lakes, ponds, rivers, bays, estuaries, fresh and saltwater marshes. They prefer
waters up to 4 ft. (1.2 m) deep where they can reach submerged vegetation, but will move
to deeper water when the shallow water freezes in winter.
Diet. Mute Swans consume submerged aquatic vegetation (SAV). Studies in the
Chesapeake Bay revealed a preference for widgeon grass (Ruppia maritima). They also con-
sume eel grass (Zostera marina), wild celery (Vallisneria americana), and several pondweeds
(Potamogeton spp., and others). They will pull up whole plants to eat or to feed to cygnets.
 MUTE SWAN n 255

Life History. Defense of large (6–13 ac. or 2.4–5.3 ha) nesting territories begins in
February, and nesting starts in March or early April. The mating ritual involves what is
known as “busking,” when males raise their wings and fluff their feathers while twirling in
place. Pair formation is not for life as legend has it, but swans are monogamous for the sea-
son and pairs may be stable for years at a time.
The female performs most of the nest building, using rushes, cattails, reeds, and stems of
other marsh plants to construct a large nest 4–6 ft. (1–2 m) in diameter, often on a mound
safely above water level. Both males and females are highly aggressive toward intruders in
their territories. Clutch size varies from 4 to 10 eggs. The young cygnets hatch about 35 days
after the last egg is laid. In the Mid-Atlantic region, hatching usually takes place in early
June. Mute swans generally nest only once a year, but if the eggs are destroyed early in the
season, they will nest a second time.
Cygnets are precocial and begin swimming within a day or two of hatching. Families typ-
ically remain on the breeding territory away from other family groups or nonbreeding adults
during the first two months or so. Cygnets grow rapidly and can fly at 4–5 months of age;
they are fully grown at 6 months. Juveniles are forced out of the nesting territory before
the next breeding season. Juveniles and subadults often gather in large flocks near open salt
water and molt, becoming flightless for a short time. Mute Swans become sexually mature at
age 2, but most will not nest until they are 3–5 years old. Swans can live in the wild up to
19 years, but the average lifespan is about 7 years.
Impacts. In the Chesapeake Bay, the Mute Swan’s consumption of SAV is of major con-
cern. SAV provides essential cover and nursery areas for a variety of fish and shellfish and
feeding grounds for a large number of native waterfowl, and it has already been seriously
depleted in estuaries of the bay due to development and pollution. Whereas native ducks,
geese, and swans use the area as wintering grounds and consume SAV for only a season,
Mute Swans are year-round residents and consumers. Plants such as wild celery reproduce
and recover from wintering grazing pressures when the native waterfowl are absent. Mute
Swans could impede the seasonal and long-term recovery of SAV and reduce winter forage
for native birds. Loss of SAV is implicated in population declines of Canvasbacks (Aythya
valisineria) and Redheads (A. americana).
The larger Mute Swan may also compete with native Tundra Swans, recently brought
back from near extinction, for food and shelter in winter. Aggression on nesting territories
has been directed against breeding Canada Geese (Branta canadensis), Mallards (Anas platy-
rhynchos), and American Black Ducks (Anas ubripes).
In Maryland, molting mute swans occupied a beach that was the last nesting area in the
state for two state-threatened birds, the Black Skimmer (Rynchops niger) and the Least Tern
(Sterna antilarium). The loafing swans trampled eggs and nestlings on the beach, leading to
the abandonment of the area by both skimmers and terns for three seasons.
Swans are very aggressive toward people, pets, and other waterfowl. They can inflict seri-
ous damage with their wings and can prevent recreational use of some shorelines.
Disturbance of SAV by swans can limit crabbing and fishing activities. Together with other
waterfowl that defecate in water, flocks of Mute Swans contribute to degraded water quality
and an increased coliform bacteria count. Cranberry farmers in New Jersey and
Massachusetts complain that swans get into their bogs and uproot cranberry plants as they
browse other submerged aquatic plants.
Management. In 2001, Mute Swans became a protected species under the Migratory Bird
Treaty Act. The U.S. Fish and Wildlife Service (USFWS) received the authority for managing
the birds. In 2002, the USFWS issued depredation permits to states for Mute Swan
256 n VERTEBRATES (BIRDS)

population control; and in 2003, it established the Atlantic Flyway Mute Swan Management
Plan, which encourages states to develop and implement location-specific management
plans. The chief means of controlling Mute Swans is by the labor-intensive practice of
addling eggs. Eggs are coated with oil to suffocate the embryos with the adults unaware of
any disturbance to their clutch. Trapping and relocating or killing adults are other popula-
tion control measures. A hunting season could be a viable option; swans were originally
domesticated in Western Europe for food. Several states prohibit the establishment of
new populations by outlawing the sale or importation of birds and requiring that
those already owned be pinioned. Every three years, extant populations in the Atlantic
Flyway are monitored by air in midsummer, when native swans and Snow Geese are in
the tundra.

Selected References
Delany, Simon. “The Mute Swan in Europe—a Preliminary Assessment of Numbers, Distribution and
Potential Risks in Dissemination of HPAI–H5N1.” Wetlands International, Feb. 14, 2006. http://
global.wetlands.org/LinkClick.aspx?fileticket=0hONijRYbSs%3D&tabid=56.
Ivory, A. “Cygnus olor.” Animal Diversity Web, University of Michigan Museum of Zoology, 2002.
http://animaldiversity.ummz.umich.edu/site/accounts/information/Cygnus_olor.html.
“Mute Swan.” New York State Department of Environmental Conservation, Fish, Wildlife and Marine
Resources, n.d. http://www.dec.ny.gov/animals/7076.html.
“Mute Swan Cygnus olor.” Invasive Species in the Chesapeake Watershed. Summary, Chesapeake Bay
Program, 2002. http://www.mdsg.umd.edu/issues/restoration/non-natives/workshop/mute
_swan.html.
“Mute Swan Management Plan.” State of Rhode Island and Providence Plantations, Department
of Environmental Management, Division of Fish and Wildlife, 2006. http://www.dem.ri.gov/
programs/bnatres/fishwild/pdf/muswan07.pdf.

n Rock Pigeon
Also known as: Common Pigeon, Feral Pigeon, Rock Dove
Scientific name: Columba livia
Family: Columbidae
Native Range. Europe, North Africa, southwestern Asia. At their maximum pre-
domestication range, wild Rock Pigeons could be found from the Faeroe Islands and south-
west Norway south through the United Kingdom and coastal France, to Spain and Portugal
and eastward along the shores of the Mediterranean in southern Europe and North Africa.
Its range also extended eastward across inland Europe into Russia, the Middle East, and
India and Nepal. Its natural habitat was cliffs, usually along coasts, where it nested in caves
and on ledges. Rock Pigeons were domesticated more than 5,000 years ago in the eastern
Mediterranean, and since then, feral pigeons have lived close to people in urban and agricul-
tural settlements. Most northern European populations of wild pigeons were driven to
extinction by overhunting or by genetic dilution from feral birds.
Distribution in the United States. Rock Pigeons are common birds, especially in cities,
throughout the United States and in Puerto Rico.
Description. Rock Pigeons are medium-sized birds with small round heads and
large chests. They are 12–13 in. (30–35 cm) long and have a wingspan of about 25 in.
(62–68 cm). When they walk, their heads bob forward and backward. The plumage is extremely
 ROCK PIGEON n 257

varied, a product of their domes-

tication. Among the two dozen
well-recognized colors and pat-
terns are those named blue-bar
or “wild type,” blue checker,
blue-T, dark (melanistic) check-
er, spread, white, pied, and Ash
red variants. The wild type is a
bluish-gray body with a dark
head. Iridescent feathers of blue,
green, and purple circle the
neck. The rump is white. Two
black wing bars mark the folded
wing, and a black band occurs
on the end of the tail. The eyes
are reddish, and the feet pink. A
white cere or fleshy swelling
appears at the top of the short
beak. Females tend to select
mates with a pattern different
from their own and so perpetu-
ate the variety of phenotypes in
a population. There also tend to
be geographic trends in color:
city populations tend to have
more checker and spread birds
than rural populations (except
in Honolulu, where 80% of the
pigeons are white); and pigeons
with red plumage are more fre-
quently encountered in the
American Southwest than in Top: The wild Rock Pigeon was a native species from Iberia to India and
northern and eastern states. in mountainous areas of North Africa, where it inhabited cliffs.
Pigeons are strong, agile Domesticated for more than 5,000 years and probably mixed with feral
flyers with pointed wings simi- stock for the same amount of time, the true native range is uncertain.
lar to falcons. When they glide, (Adapted from Kravtchenko, Viktor. “Columba livia distribution map.”
the wings are held in a V. They http://en.wikipedia.org/wiki/File:Columba_livia_distribution_map.png.)
Bottom: Rock Pigeons are common birds in cities and other human-
appear tame and often forage
modified habitats in all 50 states and Puerto Rico.
and roost in flocks. Rock
Pigeons produce a cooing
sound, but this call is mainly heard when they are on their nests or during courtship.
Related or Similar Species. The native Band-tailed Pigeon (Patagioenas fasciata) inhabits oak
and pine woodlands in California and parts of Arizona and New Mexico. It will frequent water-
holes, but is not a city bird. Another native pigeon, the White-crowned Pigeon (Patagioenas leuco-
cephala), breeds in mangroves and forages in inland hardwood forests. In the United States, it is
restricted to the southern tip of Florida and the Florida Keys and is also found in Puerto Rico.
Introduction History. Domestic pigeons were once important sources of meat and eggs,
and people carried them around the world. The first captive birds came to North America
258 n VERTEBRATES (BIRDS)

in 1606 with the settlement of

Port Royal in Nova Scotia. The
first pigeons to arrive in what
is now the United States came
with the settlers of Jamestown,
Virginia, in 1607. All Rock
Pigeons living free in the
United States derive from
escapes of domestic birds.
Habitat. Rock Pigeons are
feral birds found in proximity
to humans in farms, towns,
and cities. They require struc-
tures with ledges for nesting,
loafing, and roosting; and most
buildings serve well as surro-
Rock pigeons are familiar denizens of urban landscapes. The plumage of gate cliffs and caves. Nests will
these feral birds is extemely varied, a product of domestication. (Jaimaa/ be built wherever there is a dark
Shutterstock.)
opening, on a ledge on a sky-
scraper, in the rafters of an abandoned building, or under a bridge.
Diet. Rock Pigeons forage on the ground, mainly for medium-sized seeds. Preferences
include dried peas, wheat, oats, barley, millet, and maize. In urban settings, they also con-
sume grass seed, berries, bread crumbs, popcorn, and other particles of discarded food.
Life History. Pigeons in cities may breed all year. Most nesting, however, occurs between
March and October. Males will fluff their feathers and bow-coo in courtship of females at
any time. The pair will mate for life. The male selects the nesting site, and the pair constructs
a crude nest. The male brings nesting materials to the female, and she tucks the straw, twigs,
and feathers he brings around her body. The female typically lays two white eggs 40 hours
apart. The eggs hatch asynchronously 17–20 days later. The first egg hatches about a day
before the second. Male offspring are larger than females and often the first to hatch. The
larger nestling has a distinct advantage over its smaller nest mate, which will die in years
of food shortages. The younger sibling can be viewed as insurance for the parents if some-
thing should happen to the first one, and in good years, both will survive and fledge. For
the first 4–5 days, nestlings are fed pigeon “milk,” a cheesy secretion of the crops of both
parents that is rich in protein and fats. The crop milk is regurgitated to the young birds
and ceases to be produced after 10 days. As the pigeon milk supply declines, more and more
seeds are given to the squabs (birds 1–30 days old).
Squabs leave the nest in 4–5 weeks and are fully independent when about 7 weeks old.
A pair of pigeons may breed several times a year, sometimes constructing new nests on
top of old ones encrusted with droppings from previous broods. A typical pair in Kansas
produces 10 squabs a year from an average of 6.5 nests. Young pigeons are able to breed
at six months of age. The large reproductive potential of feral pigeons is a legacy of their
history as a domesticated species.
Impacts. Feral pigeons are no longer considered an agricultural pest in the United States.
In cities, they can be a nuisance, littering building lofts, façades, and sidewalks with their
droppings and nesting materials. Their dung can harbor the fungus Histoplasma capsulatum;
cleanup programs aimed at getting rid of pigeon and starling droppings have caused histo-
plasmosis infections in the lungs of humans in Arkansas, Ohio, Missouri, and Wisconsin.
 BLACK RAT n 259

Nonetheless, many people derive pleasure from feeding pigeons in urban parks and at feed-
ers, and eradication programs are few.
Management. The most effective way to control feral pigeon populations is to reduce or
eliminate food sources and barricade nesting and roosting sites. Cleaning up vacant lots
and reducing litter on urban streets and in parks will help with the former, as will using
feeders that minimize the amount of seed that scatters onto the ground. Netting, metal
spikes on ledges or other perches, and electric exclusion fences can help with the latter.

Selected References
Hetmanski, Tomasz, and Anna Jarosiewicz. “Plumage Polymorphism and Breeding Parameters of
Various Feral Pigeon (Columba livia Gm.) Morphs in Urban Area.” Gandsk, north Poland. Polish
Journal of Ecology 56(4): 683–91, 2008. Available online at http://www.pol.j.ecol.cbe-pan.pl/
article/ab56_4_12.pdf.
Johnston, Richard F., and Marian Janiga. Feral Pigeons. New York: Oxford University Press, 1995.
“Rock Pigeons.” AvianWeb and Wikipedia, 2006. http://www.avianweb.com/rockpigeons.html.
Roof, J. “Columba livia.” Animal Diversity Web, University of Michigan Museum of Zoology, 2001.
http://animaldiversity.ummz.umich.edu/site/accounts/information/Columba_livia.html.
Youth, Howard. “Pigeons: Masters of Pomp and Circumstance.” Zoogoer, 276. Friends of the National
Zoo, 1998. http://nationalzoo.si.edu/Publications/ZooGoer/1998/6/pigeons.cfm.

n Mammals
n Black Rat
Also known as: House rat, roof rat, ship rat
Scientific name: Rattus rattus
Family: Muridae
Native Range. Most authorities agree that black rats originated on the Indian subconti-
nent and possibly in neighboring areas of Southeast Asia. They have been so long associated
with humans, migrating with people around the world, that it is difficult to know their ori-
gins with any certainty.
Distribution in the United States. Currently, the black rat is most common in the
southern United States, although before the introduction of the Norway rat in the eighteenth
century, it was the common rat in towns and on farms in the Northeast. In the lower 48
states, it is largely found in seaports, but small, isolated populations do occur inland.
Larger populations are known from California’s Central Valley and along the Pacific coast
north into the Puget Sound region of Washington State. In the eastern United States, they
are found from Norfolk, Virginia, south along the Atlantic coast and throughout the Gulf
states. Black rats also occur in Hawai’i, where they live in moist natural forests.
Description. The black rat is slender and has a tail longer than its head-body length. The
tail is used for balance, making this rat a most agile climber on overhead wires and tree
limbs. Its ears are relatively large and hairless. The body may be black all over, or the back
may be brown to gray and the underside a lighter tone. The tail is hairless, scaly, and uni-
formly colored. Head-body length ranges from 6 to 8 in. (160–220 mm) and tail length from
8.5 to 10 in. (190–240 mm). Black rats weigh 5–10 oz. (140–280 g). Males are larger than
females.
260 n VERTEBRATES (MAMMALS)

Related or Similar Species.

The closely related Norway rat
(see Mammals, Norway Rat) is
larger and its tail is shorter than
its head-body length. The ears
are relatively small. Woodrats
(Neotoma spp.) have white
undersides. The rice rat
(Oryzomys palustris) has small
hairy ears; its gray back is
streaked with black, and its
belly and feet are whitish and
its long tail is pale below.
Introduction History. The
black rat spread along the trade
routes of Asia and Europe,
entering the eastern Mediterr-
anean in Roman times and
Europe by early medieval times.
It is the infamous carrier of the
Black Death (bubonic plague)
that ravaged Europe several
times during the Middle Ages.
In the 1500s, it arrived with
Europeans in Central and
South America. Black rats were
aboard the earliest ships arriv-
ing at Jamestown, Virginia;
Captain John Smith complained
of thousands of rats devastating
grain stores in 1612. They sub-
Top: The Indian subcontinent is the most probable home of the black rat, sequently moved from port to
but it may also have occurred originally in Southeast Asia. The original port and then dispersed inland
source area is obscured by the animal’s long dispersal history in the wherever humans lived and
company of humans. Bottom: Since the introduction of the Norway rat farmed. The invasion of the
to the continental United States, the black rat has become almost Norway rat about the time of
exclusively a resident of southern states, where it is found primarily in
the American Revolution led to
seaports. In Hawai’i, however, it is found in moist forests. (Adapted
from Marsh 2005.) a range reduction among black
rats in North America, as the
larger newcomer replaced black rats in regions with more prolonged winters. Coastal and
insular populations may be constantly replenished today as black rats continue to arrive
on incoming vessels.
Habitat. Black rats prefer warm climates and areas inhabited by people. However, their
rarity in the more temperate regions of the United States may be due more to their displace-
ment by the later-arriving Norway rat than to environmental conditions, as they were for-
merly common in towns and farms in New England and elsewhere in the northern United
States. These rodents are agile climbers and often live in treetops in natural settings or the
upper floors and attics and rafters of buildings everywhere from the inner city to small
 BLACK RAT n 261

towns. The latter is particularly

true where their range overlaps
with Norway rats, which
occupy cellars and lower floors.
Black rats also commonly
occupy ships, gaining access by
climbing up ropes or stowing
away in the cargo. While they
may live in riparian vegetation,
black rats rarely swim.
Diet. Omnivores, these
rodents prefer grains, nuts, and
fruits, but also prey on inverte-
brates and the eggs and chicks
of birds. They also like poultry
and livestock feed and cat and
dog food. Black rats will cache
solid food to consume later
and must drink water daily.
Life History. These are social
animals living in mixed groups
of adult males and females.
Nonetheless, one male is domi-
nant, and usually two or three
females are dominant to all
others in the group except the
dominant male. Males breed
with more than one female and
defend their territories and
mates. In warm climates, breed-
ing occurs all year; in more sea-
sonal climates, breeding may
peak in summer and fall. Nests
are constructed in tree cavities, The agile black or roof rat has a tail longer than its head-body length.
among palm leaves, in hedge- (Falcon Scallagrim/iStockPhoto.)
rows, in protected niches in
buildings, or in other dense cover. After a gestation period of 20–22 days, females give birth
to 5–8 altricial pups. Their eyes open and pellage begins to show in about two weeks, at which
time they begin to move around. They will be weaned in at 3–4 weeks, but stay at the nest until
they reach adult size. They will be reproductively mature at 3–4 months of age. The young of
the season leave the area when mature; it is unknown how far they disperse. Under prime con-
ditions, a female will produce five litters a year. Average lifespan for a wild rat is about one year.
Impacts. Black rats have been implicated, directly or indirectly, in the extinction of native
species, particularly on islands. In Hawai’i they contributed to the extinction of some of the
islands’ endemic honeycreepers (Drepaniidae) in the 1800s and now threaten the survival of
seabirds such as Bonin Petrels (Pterodroma hypleuca) through predation on the birds’ eggs.
Similar depredations on islands around the world led to their nomination as one of the
100 “World’s Worst” invaders by the IUCN/SSC Invasive Species Specialist Group.
262 n VERTEBRATES (MAMMALS)

Apparent proof of the rat’s impact can be seen on Anacapa Island in the Channel Islands off
California, where the rare Xantus’s Murrelet (Synthliboramphus hypoleucus) is increasing in
numbers since the eradication of the black rat in 2000–2001.
As a vector for dangerous human diseases, the black rat is infamous. In addition to
bubonic plague, caused by the bacterium Yersinia pestis that is transmitted by rat fleas, black
rats also carry leptospirosis, toxoplasmosis, trichinosis, and typhus.
Damage to tree crops such as citrus, avocado, and some nuts is a problem in some places.
Sugarcane is damaged where rats feed because they open stalks to insects and pathogens.
Like their cousins the Norway rats, black rats are major pests in grain and other feed stor-
age facilities, contaminating food with their urine and feces. In buildings, they will gnaw
electrical wiring and damage insulation. In backyards, they will consume ornamental plants,
fruits, and vegetables.
Management. The best practice is to prevent the establishment of a rat colony in the first
place. Rat-proofing roofs; clearing dense, overgrown shrubs and vines; eliminating water
sources and removing pet food left outside; and securing food and feed in rat-proof contain-
ers are first lines of defense. Trapping and poisoning black rats can be problematic as they
distrust new objects in their environment and will avoid them. Successful eradication pro-
grams on islands have relied on anticoagulant baits.
Black rats are not a protected species and may be killed or captured at any time using
mechanical or chemical means. However, the chemicals used must be registered for rat con-
trol by federal and state agencies and used in accordance with label directions.

Selected References
“Black Rat, Rattus rattus.” eNature.com, 2007. http://www.enature.com/fieldguides/detail.asp
?recnum=MA0096.
Gillespie, H., and P. Myers. “Rattus rattus.” Animal Diversity Web, University of Michigan Museum of
Zoology, 2004. http://animaldiversity.ummz.umich.edu/site/accounts/information/Rattus
_rattus.html.
IUCN/SSC Invasive Species Specialist Group (ISSG). “Rattus rattus (Mammal).” ISSG Global Invasive
Species Database, 2006. http://www.issg.org/database/species/ecology.asp?si=19.
Marsh, Rex E. “Roof Rats.” Internet Center for Wildlife Damage Management, 2005. http://icwdm.org/
handbook/rodents/RoofRats.asp.

n Feral Burro
Also known as: Wild donkey
Scientific name: Equus asinus
Family: Equidae
Native Range. The ancestor of the burro, the African wild ass (Equus africanus), evolved
in the deserts of eastern Egypt, Sudan, and the Horn of Africa. The domestic breeds, which
were the founding stock for feral burros, were Spanish in origin and most probably came
into the United States from Mexico.
Distribution in the United States. Most feral burros are on public lands managed by the
Bureau of Land Management (BLM) in Arizona, California, and Nevada. Almost half are
located in the Lower Colorado River valley. A few small populations can be found in
Oregon, Utah, and in Custer State Park, South Dakota.
 FERAL BURRO n 263

Description. The typical

burro is gray with a white muz-
zle, white eye-rings, black sho-
ulder cross and black leg bar-
ring. Many, however, are black
or spotted (paint), and white
individuals are not uncommon.
Average shoulder height is 47
in. (120 cm). Their long ears
are distinctive.
Feral burros congregate in
small groups with transient
members. Groups generally
consist of bachelor males or
females and their immature off-
spring. Only mature jacks
(males) are solitary.
Introduction History. Most
feral burros stem from domestic
animals used by prospectors
and miners during the western
gold rushes of the nineteenth
century. By the end of that cen-
tury, the development of roads
and railroads marked the obso-
lescence of pack animals, and
the boom time of small-scale
gold, silver, copper, and lead
mining was largely over. Many
burros were abandoned to fend
for themselves at that time.
Habitat. Feral burros thrive Top: Two subspecies of wild African asses occurred historically in the
in warm desert habitats as long desert east of the Nile and in the Horn of Africa. They were likely first
as permanent water sources are domesticated in Upper Egypt some time prior to 5,000 years ago. In the
available. In spring and sum- colonial period, Spain was known for its fine breeds of domestic ass,
mer, burros tend to congregate which were transported to its colonies in the New World. (Adapted
from Woodward, S. L. “Feral burros of the Chemehuevi Mountains,
in riparian habitats where shade
California: The biogeography of a feral exotic.” PhD diss., University of
and food are available; in winter, California, Los Angeles, 1976.) Bottom: The current range of free-
they frequent interfluves where roaming feral burros in the United States consists mostly of desert areas
annual forbs may be abundant. managed by the U.S. Bureau of Land Management. (Based on U.S.
Diet. Burros are primarily Bureau of Land Management’s herd management areas maps and data.)
browsers. In a study area along
the Lower Colorado River in California, more than 60 percent of the diet of feral burros was
comprised of desert shrubs such as palo verde (Cercidium floridanum), mesquite (Prosopis
spp.), and arrowweed (Pulchea sericea). Thirty percent was composed of forbs, particularly win-
ter annuals such as woolly plantain (Plantago insularis). Grasses were also consumed.
Life History. Feral burros breed year round in warm deserts. The jenny (female) gives
birth to a single foal, who accompanies her for a year or longer. Juvenile males typically leave
264 n VERTEBRATES (MAMMALS)

A. The typical feral burro is gray with a white muzzle, white eye-rings, and black shoulder cross. However, black,
white, and spotted individuals are not uncommon. (Chris Curtis/Shutterstock.) B. Most burros occur in the
deserts bordering the lower Colorado River. (Susan Woodward.)

the mother when about one year of age and assemble in bachelor groups of varying size and
membership. Occasionally, a juvenile female will tag along with a small bachelor group or
single jack for a time. Mature jacks may or may not establish temporary territories near sour-
ces of water and attempt to breed with jennies passing through it. However, many breeding
attempts also occur beyond the limits of the territory by non-territorial males.
Females are sexually mature by 10 months, males by 12 months; but they usually do not
breed successfully until older. Full physical growth is not achieved until about two years of
age. Females come into estrous immediately after giving birth, but usually do not conceive at
that time. Gestation lasts almost 12 months, and jennies may nurse their foal for up to a year.
It is common for females to foal only once every two years even in optimal habitat. Lifespan
may be up to 15 years.
Impacts. Burros establish well-developed trails along the contours of desert slopes,
which, along with their dust wallows, could accelerate erosion. Trampling can lead to com-
paction of desert soils and destruction of archeological sites. Feral burros could compete
with desert bighorn sheep (Ovis canadensis nelsoni) for forage and water and could also com-
pete with the federally threatened desert tortoise (Gopherus agassizii) for forage and crush its
burrows. They are accused of fouling precious desert water holes with their feces; but
domestic cattle are more likely perpetrators, since burros are loathe to get their feet wet.
Management. The 1971 federal Wild Horse and Burro Act mandated that the BLM pro-
tect and manage feral burros on the public lands under their jurisdiction as living symbols
 FERAL CAT n 265

of our national heritage. At that time, an estimate 14,400 burros roamed the deserts of the
Southwest. Herd Management Areas were established, and appropriate population levels
were determined for each managed population. Excess animals were removed and put up
for adoption. Today an estimated 4,700 feral burros remain in a free-roaming state. Nearly
half of these are in Arizona. The other half is nearly evenly divided between California and
Nevada.
To mitigate the potential impact of burros on bighorn sheep, water tanks have been estab-
lished in some areas and fenced so as to permit sheep to enter but not burros.
Feral burros are also managed on special sanctuaries or preserves. Many more are held in
short-term corrals and long-term pastures awaiting adoption. Most feral burros have been
removed from those national parks in which they once occurred under the National Park
Service’s mandate to eliminate nonnative species.
In 2010, the secretary of the interior announced plans to more aggressively use fertility
control measures and manage sex ratios of free-roaming populations in the management of
feral horse—and presumably feral burro—herds.

Selected References
MacDonald, C. R. “Wild Burros of the American West: 2006 National Burro Status, A Critical Analysis
of the Current Status of Wild Burros on Public Lands.” 2007. http://www.wildhorsepreservation
.com/pdf/BurroAnalysis-2006-Public.pdf.
“National Wild Horse and Burro Program.” Bureau of Land Management, 2010. http://www.blm.gov/
wo/st/en/prog/wild_horse_and_burro.html.
Woodward, S. L. “The Social System of Feral Asses (Equus asinus).” Zeitshrift fur Tierpsychologie 49:
304–16, 1979.
Woodward, S. L., and R. D. Ohmart. “Habitat Use and Fecal Analysis of Feral Burros (Equus asinus),
Chemehuevi Mountains, California,” Journal of Range Management 29: 482–85, 1976.

n Feral Cat
Also known as: Alley cat, house cat
Scientific name: Felis silvestris catus
Family: Felidae
Native Range. The domestic cats from which feral cats derive were primarily European
animals transported with colonists to North America and Hawai’i. Recent genetic studies
point to the Near East as the site of the earliest domestication of the cat and its ancestor as
the Near Eastern wildcat, F. s. lybica. The propensity of free-roaming cats to interbreed with
wild cats has introduced genes from other subspecies of Felis silvestris, including the
European wildcat (F. s. silvestris) and the Central Asian wildcat (F. s. ornata).
Distribution in the United States. Throughout.
Description. Feral cats are indistinguishable from domestic house pets and display a full
range of coat patterns. After many generations in a free-roaming state, they tend to revert to
the “wild type” tabby color pattern, with varying degrees of white on belly and chest. Adults
have a shoulder height of 8–12 in. (20–30.5 cm) and weigh 3–8 lbs. (1.4–3.6 kg). Body
length is 14–24 in. (35.5–60 cm); the long, flexible tail adds an additional 8–12 in.
(20–30.5 cm). These agile predators have retractable claws, sharp teeth, long whiskers,
and keen hearing and eyesight, including acute night vision.
266 n VERTEBRATES (MAMMALS)

Related or Similar Species.

Free-roaming but owned
domestic cats are identical in
appearance, although they tend
to be tame. Bobcats (Lynx rufus)
are twice the size and have
black-tipped, stump tails.
Introduction History.
Domestic cats were first intro-
duced to the mainland of the
United States by European trad-
ers and colonists in the seven-
teenth century. They arrived in
Hawai’i in the 1800s. Cats were
commonly kept on sailing ships
to kill vermin such as rats and
mice. Before they became pam-
pered companion animals, they
were used as work animals to
control rodent pests in barns
and fields and were only loosely
managed. Many undoubtedly
escaped to a feral state. Feral
cat populations continue to be
supplemented by unneutered
pets that wander off and pro-
duce litters in the wild. How-
ever, large numbers are the
result of the deliberate aban-
donment of unwanted pets and
litters of kittens. Often, well-
Top: Cats were probably first domesticated in Mesopotamia. Their intentioned people release
ancestors were the Near East subspecies of wildcat, Felis silvestris lybica. domesticated animals into the
(Adapted from http://en.wikipedia.org/wiki/File:Wiki-Felis_sylvestris wild rather than subject them
.png) Bottom: Feral cats are found in all 50 states and Puerto Rico. to the likelihood of euthanasia
at animal shelters. The United
States, a country where cats are
the most popular household pet, has an estimated 30–60 million feral cats.
Habitat. Feral cats are highly adaptable and occupy a wide variety of habitats and climate
regions. They tend to congregate where food is readily available and shelter is nearby.
Colonies may live in urban alleyways, near waste collection sites behind restaurants and fast
food operations where food waste, as well as foraging rodents, may be abundant; in towns,
where people often feed them; and on farms. They are also common on college campuses,
at military bases, and in other places where the human population is transient and pet aban-
donment is frequent.
Diet. Midsize carnivores, feral cats prey upon small mammals, birds, reptiles, amphib-
ians, and insects.
 FERAL CAT n 267

A. Feral cats are indistinguishable from house pets and display a full range of coat patterns. (Matt Valentine/
Shutterstock.) B. After many generations in a free-roaming state, feral cats tend to revert to the “wild type” tabby
pattern. (Daz/Shutterstock.)

Life History. Feral cats are very prolific. Sexual maturity is achieved by females between 7
and 10 months of age and sometimes even younger. Gestation is 63–65 days. Reproduction
can occur year round. A female typically produces three litters of 4–6 kittens a year. The
young are weaned in 35–40 days. Life expectancy for a feral cat is only 2–3 years, compared
with 10–20 years for a cat that is a household pet.
Impacts. Cats, feral and free-roaming pets alike, are a major threat to wildlife because of
the great numbers of small mammals, reptiles, and birds that they kill. The destruction of
native birds often receives the most attention, but the number of small mammals such as
shrews, chipmunks, and rabbits killed each year by cats is even greater and may reach more
than a billion. A single cat may kill 100 or more small mammals and birds each year. While
island populations of endemic species are most vulnerable to losses, the depredations occur-
ring in suburban and urban “habitat islands” are significant.
In Florida, cats are considered a major threat to several federally endangered small mam-
mals, including the Key Largo cotton mouse (Peromyscus gossypinus allapaticola), rice rat
(Oryzomys palustris natator), Key Largo woodrat (Neotoma floridana smalli), Lower Keys
marsh rabbit (Sylvilagus palustris hefneri), Choctwhatchee beach mouse (Peromyscus poliono-
tus allophrys), Perdido Key beach mouse (Peromyscus polionotus trissyllepsis), Anastasia
Island beach mouse (Peromyscus polionotus phasma), and Southeastern beach mouse
(Peromyscus polionotus niveiventris). They are also known to prey upon federally listed birds
such as Roseate Tern (Sterna dougallii), Least Tern (Sternula antillarum), and Florida Scrub-
jay (Aphelocoma coerulescens) and on green sea turtles.
In Hawai’i, cat predation is one more factor pressing endangered birds—such as the
Hawaiian Crow or ‘Alalā (Corvus hawaiiensis); Hawaiian Goose or Nēnē (Branta sandvicensis);
Palila (Loxioides bailleui), a Hawaiian honeycreeper; and Hawaiian Petrel or ‘Ua’u (Pterodroma
sandwichensis)—toward extinction.
268 n VERTEBRATES (MAMMALS)

Feral cats can be reservoirs for feline diseases such as feline leukemia and FIV, threatening
native bobcats and mountain lions as well as domestic house cats. They also can transmit rabies
to pets and humans and may carry parasites such as Toxoplasmosis gondii, roundworm, and
hookworm. Fleas in cat colonies can spread to nearby human dwellings and workplaces.
Nominated by the IUCN as among “100 of the World’s Worst” invaders, feral cats are
notorious as introduced predators on islands all over the world.
Management. Feral cat control is an emotionally charged issue. Animal welfare advocates
and conservationists tend to have different goals and therefore different approaches to the prob-
lem. Humane organizations are interested in the well-being of the cats; conservationists focus on
halting depredation of native fauna. Trap-neuter-release (TNR) programs tend to be least con-
troversial in most communities, but do not prevent the ecological damage a dense colony of car-
nivores can impose on native small animal populations. While reducing the number of breeding
adults in a cat colony should over time reduce the feral population, recruitment of new members
will continue as long as people abandon or fail to sterilize their pets. Cats are thought of as soli-
tary, but they are only slightly territorial and, where food is abundant, will congregate in large
numbers. Permanent removal of animals from a feral cat colony tends to open space for new
arrivals. A sterilized cat released back to the colony, on the other hand, creates competition for
food and shelter and may actually increase the mortality of kittens born into a feral state.
Many feral cat colonies are managed and fed by volunteers, who capture, spay, and vac-
cinate new strays entering the population. Kittens may be removed, tamed, and put up for
adoption. Nonetheless, small mammals, birds, and reptiles will continue to be killed. As
every cat owner knows, even a well-fed cat kills.
On some islands, eradication of feral colonies by trapping and euthanasia may be pos-
sible, but on the mainland, the ecological impacts of cats will continue as long as millions
of pet cats are neither sterilized nor confined, and as long as people feed and care for strays.

Selected References
Driscoll, Carlos A., et al. “The Near Eastern Origin of Cat Domestication.” Science 37: 519–23, 2007.
LaBruna, Danielle. “Domestic Cat (Felis catus).” Introduced Species Summary Project, Columbia
University, 2001. http://www.columbia.edu/itc/cerc/danoff-burg/invasion_bio/inv_spp_summ/
Felis_catus.html.
Longcore, Travis, Catherine Rich, and Lauren M. Sullivan. “Critical Assessment of Claims Regarding
Management of Feral Cats by Trap-Neuter-Return.” Conservation Biology 23: 887–94, 2009.
Masterson, J. “Felis catus, Feral House Cat.” Smithsonian Marine Station at Fort Pierce, 2007. http://
www.sms.si.edu/IRLspec/Felis_catus.htm.
Verdon, Daniel R. “Feral Cats: Problems Extend to Wildlife Species, Ecologists Say.”
DVM Newsmagazine, 2002. http://veterinarynews.dvm360.com/dvm/article/articleDetail.jsp
?id=31506&sk=&date=&&pageID=1.

n Feral Goat
Scientific name: Capra hircus
Synonym: Capra aegagrus hircus
Family: Bovidae
Native Range. Goats were first domesticated in the Zagros Mountains region of western
Iran. Most island populations of feral goats in the United States likely derived from Iberian
domestic breeds.
 FERAL GOAT n 269

Distribution in the United

States. Invasive populations of
feral goats are or were found in
the Channel Islands of
California and all the major
islands of Hawai’i. Feral goats
also exist on the island of
Mona off Puerto Rico. Small
populations of feral goats
descended from escaped or
abandoned livestock are found
in isolated areas of the main-
land United States (e.g., in
California foothill habitats), but
generally are not considered a
problem because their numbers
are held in check by predation
and hunting.
Description. San Clemente
Island (California) goats are the
best-described population.
These animals adapted to an
island environment over several
hundred years in part by
becoming relatively small and
fine-boned. They are frequently
described as deer-like and have
a shoulder height of 21.5–29.5
in. (55–75 cm)—only slightly
larger than dwarf breeds of
domestic goats. Both males and
females possess horns that spi- Top: The wild ancestor of domestic goats, Capra aegagrus, is native to the
ral upward and outward. The mountains of western Iran, where domestication seems to have first taken
males have heavier horns than place. (Adapted from Altaileopard, http://commons.wikipedia.org/wiki/
females and also have beards. File:Capra_aegragus_map.png.) Bottom: Feral goats are notorious
Before extensive culling and invaders on islands. Today they remain on the Hawaiian Islands and on
removal of goats to the main- Mona Island, Puerto Rico.
land (see Management below),
San Clemente Island goats displayed a variety of coat colors; today, they are mostly red or
brown with black markings, especially on the head and forelegs.
Feral populations usually assemble in herds of 5–20 animals. Herds may consist of mixed
age and sex groups, all-male groups, or females and their young.
Related or Similar Species. Goats are often confused with sheep. An easy way to tell them
apart is to look at the tails: goats’ tails are held up, while sheep tails droop downward.
Introduction History. Early Spanish and Portuguese explorers frequently dropped goats
off on islands around the world to run wild and breed in order to serve as a meat supply
for sailors arriving in the future. It has long been believed that the goats on the Channel
Islands of California were introduced in this manner, but later supplemented by stock from
270 n VERTEBRATES (MAMMALS)

A. Feral goats on Mauna Loa, Hawai’i, in the 1970s. (National Park Service.) B. Slopes on Santa Catalina Island in
the Channel Islands of California were deeply gullied and overgrown with prickly pear cactus as a consequence
of overgrazing by feral goats when this photo was taken in the late 1970s. (Susan Woodward.)

California’s Spanish missions and later still by farmers living on the islands. Recent genetic
analysis of San Clemente goats, transplanted to that island from Santa Catalina in 1875, indi-
cated that the island goats were not Spanish in origin, but a genetically distinct breed.
Captain James Cook brought the first goats to Hawai’i in 1778.
Habitat. Goats are hardy animals and adapt to a variety of habitat types. In California,
feral goats subsist in Mediterranean scrub habitats, valley foothill hardwood, and valley foot-
hill hardwood–conifer woodlands. They seek shelter on steep slopes and rock outcrops. In
Hawai’i, they live from sea level to high alpine habitats with distinct dry seasons and prefer
rocky slopes and open lava fields. A population on an island off South Carolina adapted to a
hot, humid, and swampy environment.
Diet. Goats are browsing animals and thrive on brush and other coarse vegetation unpal-
atable to sheep and cattle. They also consume grasses.
Life History. Dominant males breed with estrous females serially. Gestation lasts 147–
155 days, after which time 1–3 precocial young are born. Twinning is common. Sexual
maturity in both sexes may be reached by six months and definitely by one year of age.
They may breed twice a year under favorable conditions.
Impacts. Severe overgrazing by feral goats on islands can leave areas essentially devoid of
vegetation. Not only are endemic plant species threatened in this manner, but the removal of
food and cover endangers native island birds and other animals as well. In less severe situa-
tions, vegetation structure and species composition can be altered. Soil erosion and down-
slope sedimentation are other consequences of unmanaged browsing by goats. Ecosystem
degradation and loss of biodiversity are potential outcomes.
Management. On both the Channel Islands and in Hawai’i, hunting was tried as a way to
reduce feral goat populations without success. Reports from Hawai’i Volcanoes National Park,
for example, indicate that 70,000 goats were removed between 1920 and 1970, but the goat
population remained high. On San Clemente Island, owned by the U.S. Navy since 1934, sys-
tematic removal of goats began in 1972. Large numbers were driven into traps with net-wing
fences and killed. Some 16,000 animals were removed between 1975 and 1979 and another
8,000 between 1979 and 1982, when an estimated 4,000 goats still roamed the island.
Goats have since been completely eradicated from Lana’i Island, Hawai’i, and San
Clemente Island using a combination of techniques including aerial hunting from helicop-
ters; specially trained goat-hunting dogs to flush out remaining survivors; and “Judas goats,”
 FERAL HORSE n 271

sterilized goats outfitted with radio-transmitters that seek out herds to join and thus detect
any goats missed by other methods.
When the navy proposed shooting goats from helicopters to eliminate them from San
Clemente, the Fund for Animals, an animal welfare group, stepped in and live-trapped the
remaining animals with nets and helicopters and removed some 3,000 goats to the main-
land, where most were sterilized and put up for adoption. A few animals were not neutered
and formed the base of a new breed of domestic goat, the San Clemente Goat. These hardy,
disease- and parasite-resistant animals are used as meat animals and also as pets.
On Santa Catalina Island (California) and parts of Hawai’i, exclusion fencing has kept
feral goats out of localized areas and allowed the regeneration of native plants.

Selected References
“Capra hircus.” Introduced Species in Hawaii. Senior Seminar Biology Department, Earlham College,
2002.
Coblentz, Bruce. “Capra hircus (Mammal).” USA and IUCN/SSC Invasive Species Specialist Group
(ISSG). ISSG Global Invasive Species Database, 2008. http://www.issg.org/database/species/
ecology.asp?si=40.
Edmundson, Leslie. “San Clemente.” Breeds of Livestock. Oklahoma State University, 1997. http://
www.ansi.okstate.edu/breeds/goats/sanclemente/index.htm.
“San Clemente Goat.” American Livestock Breeds Conservancy, n.d. http://www.albc-usa.org/cpl/
sanclementegoat.html.

n Feral Horse
Also known as: Wild horse, mustang; Banker horse
Scientific name: Equus caballus
Family: Equidae
Native Range. Europe. Feral horses descend from domestic horses bred in Spain and
parts of northwestern Europe. The earliest domesticated horses have been traced to the
steppes of Kazakhstan and Ukraine, but Iberian wild horses may have contributed to the
development of European breeds.
Distribution in the United States. Populations of free-roaming horses occur in 10
western states, primarily on the public lands administered by the Bureau of Land
Management. In addition, a population is maintained in Theodore Roosevelt National
Park, North Dakota. Other herds are found at several sites along the Atlantic coast, including
North Carolina’s Outer Banks and Assateague National Seashore and Chincoteague National
Wildlife Refuge in Virginia. About half all feral horses reside in Nevada.
Description. Feral horses are indistinguishable from domestic horses, although a few
populations have been isolated long enough to preserve or evolve unique genetic informa-
tion. Western horses generally stand 14–15 hands (4.75–5 ft. or 1.4–1.5 m); males weigh
795–860 lbs. (360–390 kg) and females 595–750 lbs. (270–340 kg). Assateague ponies
and other eastern insular horses tend to be smaller, with shoulder heights seldom more than
13 hands (4.3 ft. or 1.3 m). Feral horses come in all colors and patterns, including solid col-
ors ranging from black to cream, paints of every type, and leopards.
Of genetic significance are the few herds of pure Spanish mustangs descending from the
earliest domestic horses brought to the Americas. These horses have short, straight backs
and deep, narrow chests that are V-shaped when viewed from the front. The croup (rump)
272 n VERTEBRATES (MAMMALS)

is low, and the tail set low. They

usually have straight to concave
foreheads and convex noses.
Examples include the Kiger
mustangs in southeastern
Oregon and the Cerbat mustangs
found near Kingman, Arizona.
The feral horses of Theodore
Roosevelt resemble horse types
common in the nineteenth cen-
tury, but rare today. They have
large heads and short backs
and are frequently blue or red
roans with “bald” or “apron”
(white) faces and white patches
on their sides.
Related or Similar Species.
Feral horses are generally indis-
tinguishable from domestic
horses. In western states, ranch-
ers and Native Americans often
allow their horses to run free
when they are not using them,
and these animals could easily
be mistaken for feral horses.
Introduction History. Chris-
topher Columbus, on his sec-
ond voyage to the New World
in 1493, brought the first horses
to the Americas when he landed
them on Hispaniola. From that
Top: The wild ancestor of the domestic horse was native to the steppes of Caribbean island, Spanish con-
Eurasia, but it is now extinct in the wild. Bottom: Free-roaming feral quistadors introduced them to
horses still live in the deserts and grasslands of 10 western states and a mainland North America in the
few islands off the East Coast of the United States. (Based on U.S. early 1500s. Spanish horses
Bureau of Land Management’s herd management areas maps and data spread northward through
and other reported occurrences.)
Mexico along the chain of mis-
sions set up by the Spanish.
Native Americans raided the missions and settlements of the frontier, stole the horses, and
became adept horsemen, warriors, and bison hunters. They exchanged and stole horses from
other tribes, effectively spreading the animal throughout the continent. Some groups bred
horses and developed distinctive breeds, such as the original Appaloosa. Other Spanish
horses escaped from military outposts, where stallions were used as cavalry mounts.
Large herds of semidomesticated horses developed on the Great Plains, their numbers
and genes added to when European settlers with breeds from Western Europe pushed west-
ward from the Atlantic coast of the United States. Herds of wild horses from east of the
Mississippi were displaced westward with settlement. French breeds moved south and west
from the Detroit region and north from the New Orleans colony. The U.S. Cavalry in the late
 FERAL HORSE n 273

1880s and early 1900s intro-

duced old-style East Friesian
blood to the mix. These draft
horses were preferred for pull-
ing artillery and heavy wagons;
any that escaped would have
joined the herds of feral horses
roaming the Plains.
Settlers lost horses when
wild stallions broke through
fences and liberated mares.
Furthermore, it was common
practice to let horses roam
freely on the range until needed
for work on ranches. Many
likely escaped captivity and
added to the feral herds.
Wild horses existed in the Feral horses are generally indistinguishable from the domestic horses
badlands of North Dakota when from which they descend. However, a few populations have been
isolated long enough to preserve or develop unique genetic information.
Theodore Roosevelt visited in
(Robert Broadhead/Shutterstock.)
the 1880s. He recognized them
as escapes from nearby ranches
or Indian reservations, and many carried brands. Most horses were removed from the area
in 1954, a few years after the national park was established. A few small bands eluded cap-
ture and formed the core of the current herd managed by the National Park Service.
The twentieth century saw the value of free-roaming horses decline as ranching and other
interests grew, and many were shot or taken to slaughterhouses. Populations declined to
such an extent that some feared the feral horse was headed for extinction. The mustang
became celebrated by some as a romantic symbol of the Old West and its preservation
demanded. A successful campaign initiated by “Wild Horse Annie” (Velma B. Johnston),
who was aided by the letter-writing of schoolchildren across the country, resulted in a
federal law to save the wild horse, the Wild Free-Roaming Horse and Burro Act, passed in
1971. Responsibility for managing the herds remaining on public lands fell to the Bureau
of Land Management (BLM); the U.S. Forest Service gained jurisdiction over horses in
national forests and grasslands. Under the protection of these agencies, feral horse popula-
tions began to grow again. In 2010, BLM estimated some 33,700 horses were on the range-
lands they manage.
In their evolutionary history, horses evolved in North America and crossed the Bering Land
Bridge into Eurasia during the Pleistocene Epoch. They subsequently became extinct in North
America 11,000–13,000 years ago, but were domesticated in Ukraine about 5,000 years ago.
Horses were reintroduced—as domestic animals—by the Spanish early in the sixteenth century.
The origin of the species on this continent has led some to propose that feral horses are native,
not exotic species and that they are part of the natural heritage of the West.
Habitat. In the West, feral horses generally inhabit open shrublands and woodlands in
arid and semiarid regions. Optimal rangelands have grassy areas and riparian zones. On
the offshore islands of the Atlantic coast, feral horses are associated with salt marshes.
Diet. On western ranges, feral horses prefer grasses, forbs, and sedges but also browse
shrubs such as shadscale saltbush (Atriplex confertifolia), sagebrush (Artemesia tridentata),
274 n VERTEBRATES (MAMMALS)

and rabbitbrush (Chrysothamnus nauseosus). Assateague ponies graze preferentially on salt-

marsh cordgrass (Spartina alterniflora), but will eat American beach grass (Ammophila brevi-
ligulata) and three-square rush (Scirpus americanus); they also browse woody plants.
Life History. Most feral horses live in small, stable social groups of 5–15 animals. A band
consists of a mature stallion and his harem, a group of 4–6 mares and their younger offspring.
Sometimes a subdominant male also is part of the band. The group maintains a strict social
hierarchy and often roams in an established territory. The mares will stay together even when
the stallion is lost. Young stallions that have yet to acquire harems form bachelor herds. Old
stallions that have lost their harems to younger males are usually the only solitary animals.
The harem forms the breeding unit. Mating may occur year round but peaks during the
foaling season, which is typically early spring. Usually a mare produces a foal every other year
after a gestation period of about 340 days. When a mare is ready to foal, she temporarily leaves
the harem and gives birth to a single offspring. The newborn can run and swim shortly after
birth; and its mother brings it into the band a few hours later. The foal will continue to nurse
until its mother is about to give birth again. Both fillies and colts are driven from the band when
they reach sexual maturity around 2–3 years of age. The young mares will be collected into an
existing or new harem by a stallion that is not their father. Young males will form into bachelor
groups until such time as they can acquire mares of their own either by successfully challenging
an aging stallion or by assembling a group of mares through stealth or upon the death of
another male. Mature mares rarely move to another band.
In the absence of natural predators and on good range, feral horse populations may grow
rapidly, doubling in numbers in four years. Recruitment rates are especially high after major
population reductions due to severe winter weather or roundups.
Impacts. It is difficult to assess the real impacts of feral horses on native ecosystems
because of the high degree of emotion surrounding the issue of their role on western range-
lands. Apparently from 1600 to 1850, vast herds of horses occurred throughout the Great
Plains, where they had transformed life for the Plains Indians. They shared the range with
bison, pronghorn, elk, and mule deer. Extermination of horses accompanied the campaign
to remove Native Americans from the West and also the development of modern cattle
ranching. Although free-roaming horses were captured to become cow ponies and saddle
horses, cattlemen more and more regarded them as nuisances that may lure away their
domestic horses and compete with cattle for forage and water.
Scientific studies reveal a dietary overlap of 80 percent or higher between feral horses and
cows, both species of which are predominantly grazers. Since horses also browse, they are
actually better suited to much western rangeland than are cattle; but Americans do not con-
sume horsemeat, so the argument that horses should replace cattle is moot. With respect to
impacts on other animals, comparatively little research has been conducted. Their diets do
not seem to overlap significantly with deer, elk, or pronghorn, all of which are primarily
browsers. Potentially, horses trample the nests of ground-dwelling birds. Reptile diversity
and abundance may be reduced in areas occupied by feral horses. On well-managed ranges,
plant diversity has been shown to increase with a variety of large grazing and browsing
mammals, including feral horses, present. Greater abundance and diversity of large verte-
brates occurred where horse droppings accumulated and soil was intermediately disturbed
in California’s Anza-Borrego State Park. Trampling of vegetation and compacting of soils
on trails made by horses could be localized problems and could accelerate soil erosion.
Management. Feral horses are federally protected under the 1971 Wild Free-Roaming
Horse and Burro Act. The law is administered by the BLM (Department of the Interior) and the
U.S. Forest Service (Department of Agriculture). Shooting or poisoning feral horses in the wild
 FERAL PIG n 275

became a federal crime. When BLM assumed authority for feral horses, an estimated
17,300 mustangs roamed western rangelands. In 2010, the number was estimated to
be 33,700. As populations grew in the 1970s, amendments to the law in 1976, 1978, 1996,
and 2004 mandated management of herds in such a way as to sustain their habitats and author-
ized humane removal of excess animals from the public lands. The 1976 Federal Land Policy
and Management Act included feral horse management as part of BLM’s and the Forest
Service’s multiple-use missions that require managing public lands for recreation, livestock
grazing, mineral and energy production, and the conservation of natural, historical, and cul-
tural resources. Feral horses are considered part of our national heritage.
Appropriate Management Levels (AMLs) have been determined for the 201 Herd
Management Areas designated by the BLM and the 37 that it comanages with the U.S. Forest
Service. Herd reductions to achieve these population sizes and to maintain them have relied
largely upon roundups and selecting excess animals to be removed and put up for adoption.
The roundups use helicopters and cowboys on horseback, and are expensive and dangerous
to both the horses and their captors. The Adopt-A-Horse program began as a way to transition
feral horses back to private ownership and domestic status. Since 1971, BLM has placed more
than 225,000 horses and burros in private care. Horses that prove impossible to adopt are held
in short-term holding pens and on long-term pastures. In May 2010, 10,700 feral horses (and
burros) were in corrals and another 24,400 maintained on pastures in the Midwest. An esti-
mated 33,700 were on BLM-managed range, whereas the total AML has been set at 26,582.
Fertility control is now a viable practice to reduce herd growth rates. The administration
of porcine zona pellucida vaccine (PZP), an immunocontraceptive, by dart guns in the field
has proved to be a safe and effective means of population control in conjunction with
removal and natural mortality. Currently, booster shots are required annually.
In June 2010 the secretary of the interior proposed new solutions to wild horse manage-
ment, which he deemed unsustainable for the horses, the habitat, and the taxpayer in its cur-
rent form. In part, he advocated more aggressive use of fertility control measures and the
establishment of new wild horse preserves across the country in order to showcase these ani-
mals to the American public and to enhance local economies through increased tourism.

Selected References
“Colonial Spanish Horse.” American Livestock Breeds Conservancy, n.d. http://www.albc-usa.org/cpl/
colonialspanish.html.
Sponenberg, D. P. “The Colonial Spanish Horse in the USA: History and Current Status.” Archivos de
zootecnia 41(extra): 335–48, 1992.
“Wild Horse and Burro Quick Facts.” U.S. Bureau of Land Management, 2010. http://www.blm.gov/
wo/st/en/prog/wild_horse_and_burro.html.
“Wild Horse: Equus caballus.” Enature.com, 2007. http://www.enature.com/fieldguides/detail.asp
?recnum=MA0169.
“Wild Horses.” Theodore Roosevelt National Park, n.d. http://www.theodore.roosevelt.national
-park.com/nat.htm#wil.

n Feral Pig
Also known as: Wild pigs, feral hog, feral swine, Pineywoods rooter, razorback; Eurasian or
European wild boar, Russian wild boar
Scientific name: Sus scrofa
Family: Suidae
276 n VERTEBRATES (MAMMALS)

Native Range. Europe. Feral hogs are descendents of domestic breeds originally brought to
the United States from Europe. Recent genetic analysis suggests that these breeds arose from
wild boars native to Europe. In addition to domestic breeds, some truly wild boars from
Europe were deliberately released in several states as game animals.
Distribution in the United States. Feral hogs are found primarily in the southern tier of states
from California to North Carolina. Isolated populations are reported as far north as Wisconsin and
New Hampshire. The greatest
numbers occur in California,
Florida, and Texas. Feral pigs
inhabit all of the major islands of
Hawai’i and occur on St. John in
the Virgin Islands.
Description. Three types of
free-roaming pig occur in the
United States: feral hogs derived
from escaped or released domes-
tic stock, European wild boars,
and hybrids between them.
Most are feral livestock. These
medium-size, cloven-hoofed
mammals have long, pointed
heads and stocky bodies. Males
and females are similar in
appearance. Reflecting their
domestic origins, feral pigs dis-
play a variety of coat colors and
patterns, from solid black,
brown, white, or red, to spotted
with several colors, or belted
with a wide pale band across
the shoulders. Their coat is
coarser; bristles tend to be
longer than those of domestic
pigs, but shorter than those of
wild boars or hybrids. The tail
is straighter than a domestic
pig’s and never coiled. The flat-
tened snout is flexible and elon-
Top: The wild boar, ancestor of domestic pigs, is widespread in Eurasia. gate. Total body length is 3.6–
However, the domestic breeds that were brought to the United States and 4.9 ft. (1.1–1.5 m). The average
that contributed to the feral pig gene pool seem to have descended only adult has a shoulder height of
from European populations. (Range of wild boar adapted from about 3 ft. (1 m); males (boars)
Altaileopard, http://en.wikipedia.org/wiki/File:Sus_scrofa_range_map.jpg. weigh on average 130 lbs.
Source of domestic breeds based on Larson et al., “Worldwide (60 kg) and females (sows)
Phylogeography of Wild Boar Reveals Multiple Centers of Pig
about 110 lbs. (50 kg).
Domestication.” Science 307: 1618–21, 2005.) Bottom: In the United
States, feral pigs are found primarily in southern states and on all the major Maximum weight is probably
islands of Hawai’i. (Adapted from “Feral/Wild Pigs: Potential Problems for about 300 lbs. (135 kg). Males
Farmers and Hunters.” USDA Agricultural Bulletin No. 799, 2005.) have four very sharp tusks that
 FERAL PIG n 277

continue to grow throughout the lifetime of the boar, although they usually break or
wear down to lengths of approximately 5 in. (12.5 cm). The lower tusks are continually
sharpened as they rub against the upper pair. Tusks are used to establish dominance
among males and to defend against predators and other males. The shoulder skin of males
thickens as they age into a shield, a product of both aging and fighting; it consists of tough
scar tissue and cartilage. Some but not all feral piglets are striped, much like the piglets of
wild boars.
European wild boars are usually light brown or black with cream-colored tips on the bris-
tles. The ends of these long stiff hairs are often frayed. The belly hairs are lighter and the legs,
ears, and tail darker than the rest of the body. Long side whiskers develop and hairs on the
back of the neck give them the razorback appearance. Purebreds have longer legs and snouts
and relatively larger heads than a feral pig; they have little fat. Piglets are reddish with black,
lengthwise stripes.
Hybrids show characteristics of both parents. The length of the bristles is longer than in
feral pigs, but shorter than in wild boars. The diameter of the bristle shaft, however, is
smaller than either feral or wild pigs.
Occasionally, truly enormous pigs are captured in the wild. The infamous “Hogzilla,”
shot in Georgia in 2004 and estimated to have weighed some 800 lbs. (363 kg) and have
had a total body length of 8 ft. (2.4 m), was determined by National Geographic research-
ers to be a hybrid between a domestic pig and a wild boar. Others are likely escaped or
abandoned domestic pigs, which are often bred and fattened to weigh over 1,000 lbs.
(454 kg).
The presence of feral pigs can be determined by a variety of signs. Hog tracks are square
with rounded, splayed toes. Their rooting is often extensive; it can reach depths of 3 ft. (1 m)
and may leave a field looking like it has been plowed. Wallows are depressions formed when
pigs roll in mud to protect their skin from sun and insects; they often fill with water. Rubs
develop where pigs scratch against trees, fence posts, and rocks to remove external parasites,
dried mud, and dead hair. Telltale bristles are often left behind.
Related or Similar Species. In the southwestern states of Arizona, New Mexico, and
southern Texas, there is a native pig-like animal, the collared peccary or javelina (Tayassu
tajuca). These are not true pigs and not closely related to feral swine, but belong in a New
World family of mammals, Tayassuidae. Peccaries are gray with a narrow white band around
the shoulders, and smaller than a feral hog. They congregate in herds.
Introduction History. The first domesticated pigs came to North America with early
Spanish explorers in the sixteenth century. Two hundred pigs arrived in Florida in 1539 with
Hernando de Soto as a walking larder to accompany his expedition through the Southeast.
Native Americans as well as later European settlers in the Southeast used the descendents of
these hogs, often keeping them under free-range conditions. In the early nineteenth century,
Spanish hogs accompanied the southern Native American groups resettled by the U.S.
government in the Oklahoma Territory. (The small, wattled Choctaw hog, with its fused toes
that look like a mule’s hoof, is a pure Spanish breed and is still managed today as a free-
range animal in Oklahoma.) Undoubtedly, many pigs escaped to live in a wild state from the
time of de Soto onward.
Hernando Cortes brought pigs to Mexico, and they spread north with colonizers, prob-
ably reaching Texas some 300 years ago. Many may have been abandoned when people left
Texas during its war for independence from Mexico (1834–1835). In Hawai’i, small pigs of
Asian descent were originally introduced by Polynesians around 400 AD, but were replaced
by European breeds about 200 years ago. English settlers brought their own breeds to
278 n VERTEBRATES (MAMMALS)

A. The coat and bristles of feral pigs tend to be longer and coarser than those of the domestic pigs from which they
descend. Their piglets sometimes are reddish with lengthwise black stripes like those of wild boars. (Laurie L.
Snidow/Shutterstock.) B. Some free-roaming pigs descend from wild European boars introduced as game ani-
mals. They have longer legs and snouts than a truly feral pig. (Alan Lucas/Shutterstock.)

North America beginning in the seventeenth century, and Americans developed new breeds,
such as Duroc, Hereford, and Poland China. All these stock interbred in the free state to
form the common, genetically mixed feral hog populations that roam the United States
today.
Free-roaming feral pigs have occurred in southern states for a long time. However, range
expansions into Colorado, Illinois, Indiana, Kansas, Kentucky, Missouri, Nebraska, Nevada,
Ohio, Oregon, and West Virginia occurred mostly in the last decade of the twentieth cen-
tury. Many of these were deliberate introductions by hunting clubs or private landowners
for sport hunting. In all locations, they have been successful invaders because of their high
reproductive potential, improved disease control in domestic livestock, pasture improve-
ments related to modern livestock grazing practices, and development of watering sources
for domestic stock in arid lands.
Pure European or Russian boars were first released in the late 1800s and early 1900s for
sport hunting in New Hampshire, North Carolina, Missouri, Arkansas, and Tennessee. They
were introduced to Texas with the same purpose in the 1930s.
Habitat. Feral pigs are habitat generalists but prefer densely vegetated moist forests and
bottomlands where they can find shelter and make wallows.
Diet. Pigs are opportunistic omnivores. Their diet depends on seasonal and geographic
availability and is dominated by plant matter. Pigs will consume grasses, forbs, roots and
tubers, fruits, acorns, and nuts. They also eat fungi and live earthworms, mollusks, amphib-
ians, reptiles, and birds as well as eggs, and also scavenge carrion.
Life History. Feral pigs display several characteristics that reflect ancestral domestic
breeds that were purposefully selected for fast growth and early maturation. After a gestation
period of 115 days, feral sows generally farrow litters of 4–6 but, under prime conditions,
may give birth to 10–12 young. (Wild boars usually have smaller litters, with 3–8 piglets
being typical.) Some scientists consider pigs to be the most prolific large mammal on the
planet. As a rule, a 1:1 sex ratio exists at birth. Some females have two litters a year, but most
have only one. Although young may be born throughout the year, peak production occurs
in early spring. Piglets weigh 1–2 lbs. (0.45–0.91 kg) at birth. Young are weaned at
3–4 months. Family groups or sounders composed of 1–3 sows and their offspring may
 FERAL PIG n 279

contain three generations and 50 or more individuals; they are maintained until the young
mature. Females may stay in the family group or leave with their sisters to form a new group.
Males always leave the family group by 16 months of age. Adult males are solitary, traveling
among sounders to breed. Feral pigs are sexually mature at 6 months, but many females do
not mate until 10–12 months of age. Young males may be prevented from breeding by more
dominant boars. Adult stature is attained by age 3. Average lifespan is 4–5 years, with a
maximum of 8 years. Hunting is the primary cause of death in adults.
Groups rather than individuals have home ranges, and they vary in size from several hun-
dred to a few thousand acres depending upon habitat quality. Pigs tend to occupy different
parts of the home range seasonally. The need to cope with heat dictates the location of
summer home ranges near water and can result in nocturnal activity. Neither individuals
nor sounders maintain territories within the home range.
Impacts. Due to their rooting and trampling of vegetation, feral pigs and wild boars may
be the most extreme vertebrate modifier of natural ecosystems in the United States. They
disturb soil horizons and compact soil, decreasing water infiltration and increasing ero-
sion. Soil erosion, along with contamination by bacteria from fecal deposits, can increase
sedimentation and pollution in waterways; pig activity has been implicated in declining
freshwater mussel and aquatic insect populations in some areas. Their feeding can alter
plant species composition and plant community structure. Disturbance of the soil and
herb layer allows invasive plants to spread by creating habitat in which weedy exotics gen-
erally outcompete native plants. Feral pigs certainly compete with native animals for food
and other resources. This is particularly true of mast-eaters such as white-tailed deer
(Odocoileus virginianus), Wild Turkey (Meleagris gallopavo), and squirrels. They may also
negatively affect wildlife populations through predation, habitat destruction, and the
spread of parasites and disease. On the Atlantic beaches of the Southeast, feral pigs are
threatening the nesting success of several endangered sea turtles, including loggerhead
(Caretta caretta), leatherback (Dermochelys coriacea), hawksbill (Eretmochelys imbricata),
green (Chelonia mydas), and Kemp’s ridley (Lepidochelys kempii) turtles. In Great Smoky
Mountains National Park, their destruction of the leaf litter reduces the habitat of red-
backed voles (Myodes gapperi) and short-tailed shrews (Blarina brevicauda), while direct
predation reduces populations of threatened red-cheek salamanders (Plethodon jordani)
and Jones middle-toothed snail (Mesodon jonesianus). Tree ferns are a major part of the diet
of feral pigs in Hawai’i, where they also eat other native trees and epiphytes and reduce
vegetative cover. Feral hogs also facilitate the spread of the invasive strawberry guava
(Psidium cattleianum; see Volume 2, Trees, Strawberry Guava), a plant threatening the sur-
vival of several native Hawaiian plants and animals.
Ecological and economic damage to crops, timber, and pasture can also be significant; crop
depredations alone have been estimated at $1.5 billion a year in the United States. The greatest
impact is to hay, small grains, corn, and peanuts, although tree fruits, cotton, vegetables, and
conifer seedlings are not immune. Feral pigs are known to prey upon lambs, kids, and newborn
calves, attracted to birthing areas by afterbirth. Predation on animals is difficult to document,
since feral pigs also scavenge carcasses left by other animals. When they do hunt for themselves,
they typically kill their prey by biting and crushing the skull. However, since the entire carcass
is usually eaten, little or no evidence is left to point to the actual killer.
Feral pigs carry several parasites and disease. Among those that could affect humans are
trichinosis, leptospirosis, toxoplasmosis, and brucellosis. Hunters should be careful han-
dling live animals and carcasses, and all pig meat should be well cooked. Livestock diseases
280 n VERTEBRATES (MAMMALS)

Ossabaw Island Hog

T he feral hogs on Osssabaw Island, Georgia, have been isolated long enough to
develop unique characteristics but also are the closest representative, geneti-
cally, of original Spanish stock. They are a rare resource for science and for future
animal breeding. These small pigs are able to store huge amounts of fat to survive
lean seasons, a biochemical adaptation analogous to non-insulin-dependent dia-
betes in humans, which makes them an excellent model for medical research.
Natural selection has also left Ossabaw Island hogs tolerant of high amounts of salt
in their diet. Animals removed from the island have served as founding stock for a
newly domesticated Ossabaw hog breed, which appears well adapted to sustain-
able or pastured pork production. Feral hogs may no longer be taken from the
island because some individuals have tested positive for pseudorabies and porcine
vesicular stomatitis (PSV); their future as a feral population depends on manage-
ment goals of the state of Georgia.
Source: “Ossabaw Island Hog.” American Livestock Breeds Conservancy, n.d. http://
www.albc-usa.org/cpl/Ossabaw.html.

are also potentially harbored in feral pigs. Those of greatest concern currently are swine bru-
cellosis (Brucella suis) and pseudorabies, a herpes virus not related to true rabies. Were foot-
and-mouth disease to be reintroduced to the United States, feral pigs could become a reser-
voir for this highly infectious virus (Aphtae epizooticae) and make eradication difficult if not
impossible. Swine brucellois is a disease of the reproductive tract that causes spontaneous
abortions, stillbirths, and infertility in both male and female pigs. Pseudorabies, also known
as “mad itch,” causes abortion and mummified fetuses in pregnant domestic sows and can be
fatal in piglets less than a month old. In other livestock, pseudorabies is an infection of the
central nervous system that causes loss of appetite, staggering, and spasms, and is almost
always fatal. Its early symptoms include scratching and rubbing and biting that can result
in self mutilation. The virus spreads from wild pigs to domestic pigs through venereal
contact.
The Invasive Species Specialist Group (ISSG) of the IUCN has nominated the feral pig as
one of “100 of the World’s Worst” invasive species.
Management. Management of feral hogs can be controversial when the interests of hunt-
ers conflict with those of agriculturalists and environmentalists. It is also extremely difficult
to eradicate established populations because they are so prolific and so mobile.
Hunting is the best means of controlling populations. Using specially trained tracking
and catch dogs or mules and dogs are popular practices. Live-trapping is also used.
Regulations for hunting feral hogs, operating hunting facilities, and importing feral hogs
vary from state to state.

Selected References
“Feral Pig Hunting Information.” Wisconsin Department of Natural Resources, 2008. http://dnr.wi.gov/
org/land/wildlife/HUNT/Pig/Pig_Hunting.htm.
IUCN/SSC Invasive Species Specialist Group (ISSG). “Sus scrofa (mammal).” ISSG Global Invasive
Species Database, 2008. http://www.issg.org/database/species/ecology.asp?si=73&fr=1&sts.
 HOUSE MOUSE n 281

Stevens, Russell. “The Feral Hog in Oklahoma.” Samuel Roberts Noble Foundation, Inc., 1999. http://
www.noble.org/ag/wildlife/feralhogs/.
Taylor, Rick. “The Feral Hog in Texas.” Texas Parks and Wildlife, 2003. http://www.tpwd.state.tx.us/
huntwild/wild/nuisance/feral_hogs/.
West, B. C., A. L. Cooper, and J. B. Armstrong. “Managing Wild Pigs: A Technical Guide.” Human-
Wildlife Interactions Monograph 1: 1–55, 2009. Berryman Institute. http://www.berryman
institute.org/pdf/managing-feral-pigs.pdf.

n House Mouse
Scientific name: Mus musculus
Family: Muridae
Native Range. Most sources state Asia or, more specifically, Central Asia as the place of
origin of this species, but recent genetic evidence suggests that the Indian subcontinent
may have been the earliest center of radiation.
Distribution in the United States. Throughout, including all Hawaiian islands, but
absent from most of Alaska.
Description. These small Old World mice have relatively large round ears and prominent
black eyes. The fur is gray to brown, with the undersides either a lighter shade than the back
or a buffy white. The feet are a drab buff color, and the tips of the toes are white. The long
grayish-brown tail, close to half the total body length, is essentially hairless; although lighter
on the underside, it is not distinctly bicolor. Unlike New World mice, the incisors are not
grooved. Adult head-body length is about 3 in. (65–95 mm); the tail is 3–4 in. (60–
105 mm) long. Average adult weight is 0.5–0.8 oz. (17–25 g).
The presence of their droppings often alerts people to their presence. Fecal pellets are
0.25 in. (6 mm) long and have longitudinal ridges and square ends.
Related or Similar Species. New World mice that also may occupy dwellings and other
buildings include deer mice (Peromyscus maniculatus) and white-footed mice (P. leucopus).
Both of these rodents have white bellies, sharply demarcated from the darker back, and tails
that are covered with hair and distinctly bicolor. The tail of the white-footed mouse is
shorter than its head-body length. The incisors of both are grooved.
Introduction History. The house mouse has long been associated with humans. The ear-
liest evidence of its commensal relationship with people goes back to a Neolithic site in
Turkey dated at 8,000 BP. It was able to spread from settlement to settlement across Asia
with expanding Neolithic populations and later along trade routes into Europe, where two
subspecies, M. m. musculus and M. m. domesticus, were found by 4,000 years ago. It was
likely accidentally introduced into Florida by Spanish explorers in the early 1500s and came
to what is now the northern United States with French fur traders and English settlers in the
early 1600s. This small rodent easily stows away in tiny spaces and in grain and other food
stores on ships. Ships carrying goods to English colonists in the Pacific spread house mice to
remote islands.
Habitat. House mice have had a commensal relationship with humans for at least
8,000 years. Thus, they are usually found in close association with human dwellings and
other structures such as granaries, barns, and stores. They may live in stone walls, fence-
rows, cultivated fields, and other areas of dense cover close to buildings during warmer sea-
sons and retreat to buildings during winter. They do not hibernate. House mice nest in
cracks and crevices in stone walls, in woodpiles, behind rafters, or in other snug places near
food, rarely moving more than 50 ft. (16 m) from these secure spots to feed. In dense grass,
282 n VERTEBRATES (MAMMALS)

they will make their own run-

ways or share those made by
native mice. House mice are
rarely encountered in undis-
turbed or natural habitats.
Diet. House mice prefer
cereal grains, but consume a
variety of other plant material
including fleshy roots, leaves,
and stems. They also eat insects
and sometimes meat.
In houses, they will nibble on
any type of human food as well
as glue, paste, and soap. They
are physiologically capable of
extracting much of the water
they need from their food and
concentrate their urine, so they
may live without free water if
the food supply available to
them allows.
Life History. House mice
have a very high reproductive
potential and may, in mild cli-
mates, breed all year. Free-living
populations outside of human
habitations tend to have a sea-
sonal breeding pattern and may
only breed from early summer
into fall. A female produces 15–
150 young in a year, depending
Top: The house mouse is of Asian origin, and recent genetic analysis on conditions. Individual mice
supports the Indian subcontinent as its likely source. However, many will make their own nests even
have long thought of Central Asia as the starting place for this rodent, though they live in groups and
which has traveled around the world with humans. Bottom: The house share escape holes, latrines, and
mouse occurs throughout the United States, except for most of Alaska.

House Mice in the Laboratory

D omesticated house mice are common laboratory animals in medical and genetic
research. Many inbred strains have been developed since Clarence Cook Little
developed the first such population in 1909. Mutant strains have been since created
by normal breeding, by insertion of foreign genes, and by gene knockout, a process
that makes selected genes inoperable. Cloning mice has been possible since 1998.
In 2002, sequencing the mouse genome, which has many human homologues that
make it such a valuable model for research on human diseases, was finalized.
 HOUSE MOUSE n 283

feeding areas. The nests are con-

structed of finely shredded paper
and other soft fibrous materials.
After a gestation period of
19–21 days, a female gives birth
to a litter of 5 or 6 (range 2–13)
blind, hairless young whose ears
and eyes are closed. Young mice
are fully furred at 10 days of
age, open their eyes at 14 days,
and are weaned at 21 days.
After weaning, they leave their
mother’s territory. Females are
sexually mature when 35 days
old, males when 60 days old. A
female may have 5–10 litters in The house mouse has relatively large rounded ears and prominent black
a year. The average lifespan is eyes. Neither the dark back nor the upper tail is sharply demarcated
12–18 months. from paler undersides. (Gertjan Hooijer/Shutterstock.)
Impacts. House mice have
not contributed to the extinc-
tion of any species in the United States, nor do they compete with native mice. They do
not carry the hantavirus that can be problematic with infestations of native mice in the
western states. They also have not caused the serious health problems associated with Old
World rats (see Mammals, Black Rat, and Norway Rat), although they may carry some of
the same pathogens. However, they are major agricultural pests in some areas because they
consume grains and other feedstuffs and contaminate food with their urine and droppings.
In buildings, they gnaw and can destroy electric wiring (creating a fire hazard), insulation,
woodwork, furniture, upholstery, and clothing.
Management. House mice can be controlled with poisons, fumigants, traps, and repel-
lants. To mouse-proof a structure, all openings greater than 0.25 in. (6 mm) must be blocked.
All grain, pet food, and human food should be cleaned up and stored in rodent-proof contain-
ers. Outside, removing debris and cutting and thinning dense vegetation can discourage house
mice. On those islands where mice have been eradicated, an anticoagulant poison was
administered.

Selected References
Ballenger, L. “Mus musculus.” Animal Diversity Web, University of Michigan Museum of Zoology, 1999.
http://animaldiversity.ummz.umich.edu/site/accounts/information/Mus_musculus.html.
Barwell, Ezra. “Mus musculus (mammal).” ICUN/SSC Invasive Species Group (ISSG), ISSG Global
Invasive Species Database, 2006. http://www.issg.org/database/species/ecology.asp?fr=1&si=97.
Davis, William B., and David J. Schmidly. “House Mouse.” Mammals of Texas—Online version. Texas
Tech University, 1997. http://www.nsrl.ttu.edu/tmot1/mus_musc.htm.
“House Mouse.” Wikipedia, 2009. http://en.wikipedia.org/wiki/Mus_musculus#Mice_as
_an_invasive_species.
Timm, R. M. “House Mouse.” Pests of Homes, Structures, People, and Pets. Pest Notes, Publication
7483, University of California, Division of Agriculture and Natural Resources, 2006. http://
www.ipm.ucdavis.edu/PMG/PESTNOTES/pn7483.html.
284 n VERTEBRATES (MAMMALS)

n Indian Mongoose
Also known as: small Indian mongoose, small Asian mongoose
Scientific name: Herpestes javanicus
Family: Herpestidae
Native Range. Southwest and South Asia, from Iran through India and Myanmar
(Burma) to the Malay Peninsula
and Java. Populations in the
West Indies and Hawai’i derive
from animals originating in
eastern India that were brought
first to Jamaica.
Distribution in the United
States. Hawai’i (Hawai’i, Maui,
Moloka’i, and O’ahu); Puerto
Rico; U.S. Virgin Islands (St.
Croix and St. John).
Description. Mongooses are
small, weasel-like carnivores
with slender bodies and short
legs. The long tail, equal to
about 40 percent of head-body
length, is robust and muscular
at the base and tapers gradually
toward the tip. The narrow head
has a pointed snout and short
ears. The short, soft, brown fur
has golden flecks and is paler
on the undersides than on the
back. Their eyes are amber.
Adults have head-body lengths
of 20–26 in. (500–650 mm)
and weigh 0.6–2 lbs. (300–900
g). Males are larger than females
and have wider heads and more
robust bodies.
Introduction History. The
first successful introduction of
Top: The Indian mongoose is native to southern Asia from Iran to mongooses to the Caribbean
Southeast Asia. It was brought to Jamaica to control rats and from there region occurred in 1872, when
transported to other Caribbean islands and to Hawai’i. Bottom: The four males and five females from
Indian mongoose is well established on Puerto Rico and several of the Kolkatta (Calcutta) were released
Hawaiian Islands. (Both maps adapted from Yamada, F., and K. in Jamaica. They were inten-
Sugimura. “Negative Impact of an Invasive Small Indian Mongoose
tionally brought to the islands
Herpestes javanicus on Native Wildlife Species and Evaluation of a
Control Project in Amami-Ohshima and Okinawa Islands, Japan.” to control rats that were des-
Global Environmental Research 8: 117–24, 2004. http://www.aires.or.jp/ troying the sugarcane crop.
publication/ger/pdf/08-02-02.pdf.) From Jamaica, mongooses were
 INDIAN MONGOOSE n 285

A. The Indian mongoose is a weasel-like animal with a robust tail. (Bill Hubick Photography.) B. The head is
narrow with a pointed snout and amber eyes. (Bill Hubick Photography.)

deliberately taken to other Caribbean islands including, between 1877 and 1879, Puerto Rico,
St. Croix, and St. John for the same rat-control purpose. In 1883, Jamaican mongooses were
imported by sugar growers on the Big Island of Hawai’i. Later they were taken to Mau’i,
Moloka’i, and O’ahu.
Habitat. In Puerto Rico and the U.S. Virgin Islands, mongooses are most abundant in
drier habitats, but in Hawai’i, they inhabit rainforest. They may also be found near human
settlements.
Diet. Mongooses feed primarily on invertebrates, but will take small vertebrates and
fruits. On O’ahu and Moloka’i, cockroaches are a major part of the diet. Studies in the
West Indies and on the Hawaiian islands indicate that the main mammals consumed are
introduced rats and house mice. Birds and reptiles usually comprise minor parts of the diet,
although consumption of the eggs of ground-nesting birds and of sea turtles is well docu-
mented. They are strictly diurnal hunters.
Life History. The time and length of the breeding season varies with latitude. In Hawai’i
and the U.S. Virgin Islands, pregnant females have been trapped from February through
August. Gestation lasts 49 days. Litter size ranges from one to five pups, but usually is
two. At six weeks of age, young mongooses begin to hunt with their mothers, and they stay
with her until they reach sexual maturity at 4–6 months. Females may produce 2–3 litters a
year. Life expectancy is 4–5 years.
Impacts. Although the IUCN lists the Indian mongoose as one of the world’s 100 worst
invaders, its damage to the native fauna of Caribbean islands and Hawai’i may be exagger-
ated. In the early twentieth century in Puerto Rico, for example, mongooses were blamed
for the decline of five ground-nesting birds (Key West Quail-dove [Geotrygon chrysia],
Bridled Quail-dove [G. mystacea], Black Rail [Laterallus jamaicensis], Short-eared Owl [Asio
flammeus], and Puerto Rican Nightjar [Caprimulgis moctitherus]). The nightjar was believed
to have gone extinct, but was rediscovered in 1961 in areas with and without mongoose
populations. With the exception of the Black Rail, populations of all these birds have
rebounded, suggesting a balance may have developed between the native birds and exotic
predator. By the 1980s, the Bridled Quail-dove had shifted to nesting in trees and was
common on St. Croix.
286 n VERTEBRATES (MAMMALS)

In Hawai’i also, the mongoose is blamed for the reduction or extirpation of several
endemic birds, but documentation of predation is difficult and rare. Among the birds
listed as threatened by mongooses are eight on the federal list of endangered species: the
Hawaiian Goose or Nēnē (Branta sandvicensis), the Hawaiian Crow or ‘Alalā (Corvus hawai-
iaensis), the Hawaiian Duck or Koloa (Anas wyvilliana), the Hawaiian Coot or Alae ke’ko-
ke’o (Fulica alai), the Hawaiian Stilt or A’eo (Himantopus mexicanus knudseni), the
Hawaiian Gallinule or Alae ‘ula (Gallinula chloropus sandvicensis), the Hawaiian Petrel or
‘Ua’u (Pterodroma sandwichensis), and the Newell Shearwater or ‘A’o (Puffinus auricularis
newelli).
Contrary to popular stories, mongooses are not responsible for black rats nesting in roofs
and trees; this is the rats’ natural behavior. Mongooses, however, do seem to have shifted the
relative abundance of introduced rats in Hawai’i in favor of black rats to the detriment of
Norway rats and Polynesian rats (Rattus exulans), both ground-nesters. A similar pattern
has been noted on St. Croix, where there are more black rats than Norway rats in mongoose
habitat. In Puerto Rico, Norway rats exist only in areas free of mongooses, but black rats
co-occur with mongooses.
Mongooses were not introduced to St. Croix or Nevis until well after the disappearance of
two native snakes, the Saint Croix racer (Alsophis sanctaecrucis) and orange-bellied racer (A.
rufiventris), so blaming mongooses for the reptiles’ extinctions is erroneous. However, they
are implicated in the loss of the Saint Croix ground lizard (Ameiva polops) and in reduced
populations of lizards and amphibians elsewhere. Mongooses are known to prey upon the
eggs and hatchlings of four sea turtle species on Caribbean islands: the hawksbill sea turtle
(Eretmochelys imbricata), leatherback turtle (Dermochelys coriacea), green sea turtle
(Chelonia mydas), and loggerhead sea turtles (Caretta caretta).
Since mongooses do prey on reptiles and birds, they represent one more danger to
already threatened species on islands and are considered a major deterrent to the recovery
or reestablishment of island endemics.
Management. Mongooses are often naturalized members of ecosystems into which they
were introduced and may not need management. Trapping is the most common tool used
in sensitive areas such as nesting grounds of seabirds and sea turtles. However, trapping is
expensive and labor intensive, and has only temporary results, so it is not generally
employed. The use of diphacinone, an anticoagulant, has had positive results in trials.
Prevention of introduction into areas where mongooses do not already exist is the preferred
course of action. In Hawai’i, state law forbids any person to introduce, keep, or breed mon-
gooses without permits from the Hawai’i Department of Agriculture, which will not issue
them for mongoose-free islands.

Selected References
Hays, Warren S. T., and Sheila Conant. “Biology and Impacts of Pacific Island Invasive Species. 1. A
Worldwide Review of Effects of the Small Indian Mongoose, Herpestes javanicus (Carnivora:
Herpestidae).” Pacific Science 61 (1): 3–16, 2007.
“Mongoose (Herpestes javanicus).” Hawai’i Invasive Species Partnership, 2008. http://www
.hawaiiinvasivespecies.org/pests/mongoose.html.
Roy, Sugoto. “Herpestes javanicus (Mammal).” IUCN/SSC Invasive Species Specialist Group, ISSG
Global Invasive Species Database, 2006. http://www.issg.org/database/species/ecology.asp?si=86.
 NORWAY RAT n 287

n Norway Rat
Also known as: Brown rat, common rat, sewer rat, wharf rat
Scientific name: Rattus norvegicus
Family: Muridae
Native Range. Northern China and Mongolia, perhaps originally living along stream banks,
but for millennia living in close association with human settlement. (This rodent did not origi-
nate in Norway, as its name mistakenly implies. Indeed, it was unknown in Europe until the
medieval period and did not make its way into Western Europe until the early 1700s.)
Distribution in the United States. Found throughout the United States.
Description. Norway rats
have coarse brown or dark-gray
fur dorsally with underparts a
lighter shade of the general body
color. The tail is hairless and
scaly, and shorter than the head-
body length. The ears are promi-
nent but relatively short and bald.
Adult head-body length averages
about 10 in. (25 cm), total length
about 15.5 in. (39 cm). Adult
body weight ranges from 0.5 to
1.0 lb. (200–400 g); males are
larger than females.
The presence of these noctur-
nal animals is often indicated by
their droppings, the effects of
gnawing and rubbing, and
sounds such as scratching and
squeaking in the walls. Drop-
pings occur in runways, feeding
areas, and near sites of shelter;
single pellets can reach 0.75 in.
(2 cm) in length and 0.25 in.
(0.6 cm) in diameter.
Related or Similar Species.
The smaller black rat (Rattus rat-
tus), another invasive species
(see Mammals, Black Rat), has a
tail that is longer than its head-
body length. Native woodrats
(Neotoma spp.) have white under-
sides. The black rat is much
Top: The Norway rat originated in northern China and Mongolia and not
rarer in the United States than Norway, as its common and scientific names imply. Bottom: The Norway
the Norway, having been largely rat arrived in the United States at the time of the American Revolution and
replaced by it after the Norway largely replaced alien black rats. Today it is found in all 50 states and
rat arrived in this country. Puerto Rico.
288 n VERTEBRATES (MAMMALS)

Introduction History. The

early history of the rat as a
human commensal is still to be
unraveled, but it appears to have
spread along routes of human
migration and trade out of
northern China and Mongolia
during the Middle Ages. Stow-
aways on ships, Norway rats
were present in England by
1730 and soon were reported in
other European countries, reach-
ing Spain by 1800. The first rats
showed up in North America
prior to or during the American
Revolution (1770s), according
to some accounts, arriving in
Norway rats have coarse brown or gray fur on the back; the underparts grain stores brought in with
are lighter. The hairless scaly tail is shorter than head-body length. Hessian soldiers fighting with
(S. Cooper Digital/Shutterstock.)
the British against the American
colonists. Rats then moved from
port to port as accidental hitchhikers on ships. On land, their prodigious ability to reproduce,
their adaptability to a wide range of habitat and food types, their propensity to live with
humans, and their ability to climb, swim, jump, and dig allowed them to expand their range
and invade farmstead, village, town, and city. When overcrowding occurs, they will undertake
mass migrations to new areas.
The rats on Rat Island, Alaska, finally eradicated in 2009, derived from the wreck of a
Japanese ship in 1780.
Habitat. The Norway rat occurs in close association with humans, occupying such man-
made environments as cities, towns, farms, garbage dumps, and sewers. They may also be found
in disturbed and human-modified habitats such as marshes, vacant lots, and open fields and
may dig complex systems of burrows in the banks of streams and canals. They are excellent
swimmers. Away from water, they are ground dwellers, and in buildings, they prefer cellars
and lower floors, although they can climb. They may also burrow under foundations and side-
walks as well as roads and railroad tracks.
Diet. True omnivores, Norway rats eat everything humans eat and more. Experimental
studies show that, if available, they will select a well-balanced, nutritious diet of fresh foods
including cereals, meat, fish, nuts, and fruits. Household garbage usually provides adequate
moisture and limits their need for drinking water. They forage mainly at night, led by their
excellent sense of smell. Cautious about new food, they avoid unfamiliar items at first and
then test small amounts to see if it will make them sick. They can detect very low levels of
contaminants, making poisoning a problematic strategy for rat control.
Life History. Norway rats construct nests below ground and line them with leaves, twigs,
shredded paper, or other fibrous materials. Each female gives birth to a litter of about eight
pups after a gestation period lasting 22–24 days. Several females may use the same commu-
nal nest, and all females care for all of the young. The small, hairless pups do not open their
eyes for 14–17 days and will not be weaned until 3–4 weeks old, at which time they leave
the nest. Females come into estrous about 18 hours after giving birth and will mate.
 NORWAY RAT n 289

Good Rats

R ats are often associated with filth and disease, but domestic descendents of
the Norway rat have had a positive influence on humanity. They are commonly
used in research and are familiar to undergraduates as the albino laboratory rat
that runs through mazes and is subjected to a variety of behavioral experiments
in psychology classes. Domestication has also produced the fancy or pet rat, a
clean and intelligent companion animal that comes in a variety of colors and
markings.
Most American laboratory rats trace their origins to the Wistar Institute in
Philadelphia, where in 1906 an albino strain was developed by Helen Dean King
for use in biological research. Later strains include the Sprague-Dawley rat, a multi-
purpose research animal prized for its docility; the black-hooded Long Evans rat
used in behavioral and obesity studies; Zucker rats, lean and obese forms of which
are used in the understanding the genetics of obesity; and hairless rats that serve as
models for research on immune-deficiency diseases and genetic kidney diseases.
Domestic rats may go back to the eighteenth and nineteenth centuries, when
rat-catchers were employed throughout Europe to trap rats. Many of the captives
ended up as the bait in a bloodsport in which terriers competed (and people made
wagers) to see how fast they could kill all the rats in a pit. Some naturally occurring
albinos or oddly colored and marked rodents were apparently kept and the tamest
bred and sold as pets. Rat fancy first became a formal hobby in England when the
National Mouse Club accepted rats at an exhibition in 1901. In 1976 the National
Fancy Rat Society was established in England, and in 1983 the American Fancy Rat
and Mouse Association was founded in California.
Sources: “Fancy Rat.” Wikipedia. http://en.wikipedia.org/wiki/Fancy_rat.
Hanson, Anne. “History of the Norway rat (Rattus norvegicus)” Rat behavior and biology.
2003, 2004. http://www.ratbehavior.org/history.htm.
“Laboratory rat.” n.d. Wikipedia. http://en.wikipedia.org/wiki/Laboratory_rat.

Breeding occurs year round, although mating peaks in spring and fall in the temperate cli-
mates of the United States. An individual female may breed as many as seven times and pro-
duce some 60 offspring a year. Young males reach reproductive maturity at three months of
age, females at four months. Normal lifespan for a wild rat is two years.
Impacts. Worldwide, especially on islands, Norway rats have been implicated in the
extinction or range reduction of native species through both competition and predation.
On Rat Island in the western Aleutians of Alaska, for example, rat predation on eggs and
chicks led to the extirpation of such burrow-nesting seabirds as Cassin’s Auklet
(Ptychorampus aleuticus), Tufted Puffin (Fratercula cirrhata), and Storm Petrels
(Oceanodroma spp.), and likely contributed to population losses among several other
ground- and crevice-nesting shorebirds. In most of the United States, they are mainly an
agricultural pest and urban menace. Rats consume crops and contaminate stored food with
their feces and urine. They eat bird eggs and kill young poultry. They gnaw and dig their
way into buildings, where they can cause structural damage to walls and floors, destroy
insulation as they nest in and burrow in walls, and gnaw electrical wires to cause short cir-
cuits and fires. Their burrows may undermine roads, bridges, canals, and levees.
290 n VERTEBRATES (MAMMALS)

The fleas and lice on rats carry infectious bacterial diseases to both human and livestock,
including murine typhus and bubonic plague, although the latter has been more closely
associated with black rats throughout history. Flies, mosquitoes, and ticks transmit tula-
remia from rats to humans; their ticks also carry spotted fever. Contact with the urine of rats
in contaminated soil, water, or plants can transmit the bacterium Leptospira interrogans, the
cause of infectious jaundice or leptospirosis, to livestock and humans. Food poisoning (sal-
monellosis) from contaminated feed is particularly dangerous to horses. Contaminated food
may also harbor the nematodes that cause trichinosis. Bites can transmit rat bite fever.
Norway rats are responsible for millions of dollars of damage to crops and buildings in the
United States every year.
Management. On islands, poisoning rats with anticoagulants has been successful in
eradicating populations. Less slow-acting poisons are quickly learned and avoided by these
highly intelligent rodents. Other effective control methods include rat-proofing buildings by
blocking entry points, storing food in rat-proof containers, and removing potential shelter
sites such as trash heaps, overgrown vegetation, and woodpiles.

Selected References
Hanson, Anne. “History of the Norway Rat (Rattus norvegicus).” Rat Behavior and Biology, 2004. http://
www.ratbehavior.org/history.htm.
ICUN/SSC Invasive Species Group (ISSG). “Rattus norvegicus (mammal).” ISSG Global Invasive Species
Database, 2006. http://www.issg.org/database/species/ecology.asp?si=159.
Myers, P., and D. Armitage. “Rattus norvegicus.” Animal Diversity Web, University of Michigan Museum
of Zoology, 2004. http://animaldiversity.ummz.umich.edu/site/accounts/information/Rattus
_norvegicus.html.
“Norway Rat, Rattus norvegicus.” eNature.com, 2007. http://www.enature.com/fieldguides/detail.asp
?recnum=MA0095.
“Norway Rats.” Internet Center for Wildlife Damage Management, Cornell University, Clemson
University, University of Nebraska–Lincoln, and Utah State University, 2005. http://icwdm.org/
handbook/rodents/NorwayRats.asp.

n Nutria
Also known as: Coypu, swamp beaver
Scientific name: Myocastor coypus
Family: Myocastoridae
Native Range. Subtropical South America: Argentina, southern Brazil, Bolivia, Chile,
Uruguay, and Paraguay. These large, semiaquatic rodents are found chiefly in the lowlands,
but may range to elevations of 4,000 ft. (1,190 m) in the Andes.
Distribution in the United States. Alabama, Arkansas, Colorado, Delaware, Florida,
Georgia, Idaho, Louisiana, Maryland, Mississippi, New Mexico, North Carolina,
Oklahoma, Oregon, Tennessee, Texas, Virginia, and Washington.
Description. This is a fairly large, stocky rodent with a large, nearly triangular head and short
neck. The eyes and ears are small and positioned high on the head. The nostrils and mouth can
be sealed to prevent the intake of water when it dives or feeds underwater. The prominent front
teeth range in color from yellow to dark orange. The tail, which comprises roughly one-third of
total body length, is round and scaly with a sparse cover of bristles. Four toes on their hind feet
are webbed, but the outermost, fifth toe is free. The much smaller, black forefeet have four
 NUTRIA n 291

unwebbed toes that provide dex-

terity in digging and manipulat-
ing food items. The 4–5 pairs of
teats on the female are located
high on the flank so that young
may nurse while their mother is
in the water or lying on her
abdomen. The fur that made
them so valuable in the past is
actually a dense, velvety slate-
gray undercoat concealed
beneath long, coarse, but glossy
yellowish-brown guard hairs.
White hairs cover the chin.
Head and body length ranges
from 20 to 25 in.; tail length
from 10–17 in. (25–43 cm).
Nutria may weigh as much as
15–20 lbs. (6.8–9.0 kg); males
are larger than females.
Nutria droppings are cylindri-
cal in shape and as much as 3 in.
(7.6 cm) long. Fine, length-wise
grooves distinguish them from
the fecal pellets of other animals.
Droppings will be seen floating
on the water, along the shore,
and at feeding sites.
Nutria are nocturnal and
most commonly seen at twi-
light, usually when they are
swimming. A distinguishing Top: Nutria are native to the wetlands of South America, south of the
factor, then, is the narrow tail Tropic of Capricorn. (Adapted from map at http://en.wikipedia.org/
snaking behind them or arched wiki/File:Nutria-SouthAmerica.gif. ) Bottom: Nutria being raised on fur
out of the water. farms escaped captivity and established populations in numerous states,
Related or Similar Species. where they damage native marsh communities and can become agricul-
Nutria are similar to native bea- tural pests. (Adapted from Fuller, Pam. “Myocastor coypus.” USGS Non-
ingidenous Aquatic Species Database, Gainesville, FL, 2005. http://
ver (Castor canadensis) and
nas.er.usgs.gov/queries/factsheet.aspx?speciesID=1089.)
muskrat (Ondatra zibethica),
but intermediate between the
two in size and appearance. The nutria’s head resembles that of a beaver, but its tail is more
like that of a muskrat. Beavers have flat, paddle-shaped tails, whereas muskrat tails are slen-
der and flattened from side to side. Nutria are twice as large as muskrats and about a third the
size of beavers. The tracks of nutria can be confused with those of beavers, which have five
webbed toes. Nutria slides, slick muddy trails into the water, are much narrower than those
of beavers. Nutria make flattened, circular platforms of coarse emergent vegetation 3–6 ft.
(1–2 m) in diameter, where they feed, loaf, and groom, and sometimes give birth. Multiple
runways radiate out from these platforms, which could be mistaken for muskrat houses.
292 n VERTEBRATES (MAMMALS)

A. The nutria’s prominent teeth range in color from yellow to dark orange. (Sters/Shutterstock.) B. Nutria are
large, stocky rodents. Their small eyes and ears are set high on their large heads. (Bodil 1955/Shutterstock.)

Introduction History. The first nutria were deliberately introduced into the United States
from South America in 1899 in an effort to establish new ventures in fur farming. The first
attempt was made in Elizabeth Lake, California, but the animals failed to reproduce.
Between 1899 and 1940, nutria ranches were started in Washington, Oregon, Michigan,
New Mexico, Louisiana, Ohio, and Utah. The boom years for production were in the
1930s, after which time the industry collapsed due to low fur prices during World War II,
competition from beaver pelts, and poor reproduction. Some ranchers released their nutria,
while others simply failed to round up any that escaped during storms and floods, or
because of poor confinement structures. Since these animals had all been bred on ranches,
the free-roaming animals now are considered feral.
For a time, nutria were viewed as weed cutters to control noxious vegetation or as ways to
increase the take of fur trappers. Both state and federal agencies, as well as individuals, intro-
duced nutria into Alabama, Arkansas, Georgia, Kentucky, Maryland, Mississippi,
Oklahoma, Louisiana, and Texas.
California’s small feral population was eradicated in the 1970s. Nutria were eradicated or
simply failed to survive in Idaho, Indiana, Kansas, Kentucky, Michigan, Minnesota,
Missouri, Montana, Nebraska, Ohio, and Utah.
Maryland received nutria in 1943 as part of a federal program to establish an ill-fated
experimental fur station in Blackwater National Wildlife Refuge. The first feral animals were
reported in 1952, after the fur station proved unprofitable. Early attempts to eradicate nutria
failed, and the population exploded and expanded their range into Delaware. Only recently
have they finally been eradicated from Blackwater NWR, but small populations remain in
Maryland. Elsewhere in the Chesapeake Bay region, a small population in Virginia is
believed to have moved in from North Carolina.
In the early 1930s, nutria were introduced into Louisiana near New Orleans, where they
were quickly trapped out. They were introduced again in 1938, this time for fur farming,
and many escaped captivity during a 1940 hurricane. In the next 60 years, feral populations
dispersed into West Texas (1941), Arkansas (early 1960s) and Mississippi, reaching
Tennessee in 1996.
In Oregon and Washington, nutria were first imported in the late 1930s for fur farming.
By the 1950s, some 600 farms had been established in the two states. Storms and flooding
damaged holding pens and allowed animals to escape. Viable feral populations continue to
 NUTRIA n 293

exist in the Pacific Northwest. In Washington State, after a series of warm winters, nutria
seem to be expanding their distribution area.
Small feral populations survive in Colorado, Florida, New Mexico, North Carolina,
Oklahoma, and Texas.
Habitat. Nutria remain close to permanent water and may be associated with fresh and
brackish marshes, swamps, and the shores of rivers, bayous, lakes, and drainage canals.
They shelter in burrows that they excavate themselves, but they will also occupy the burrows
of other species. Their burrows may be simple tunnels or a complex network of passages and
chambers up to 50 ft. (15 m) long.
Diet. Nutria are herbivores and consume a wide variety of terrestrial and aquatic plants,
including the bark of trees. In the United States, favored foods include cordgrasses (Spartina
spp.), bulrushes (Scirpus spp), spikerushes (Eleocharis spp.), flatsedges (Cyperus spp.) pickerel-
weeds (Pontederia spp.), arrowheads (Sagittaria spp.), and cattails (Typha spp.). They also eat
baldcypress (Taxodium distichum) seedlings, duckweeds, and waterhyacinth (Eichornia cras-
sipes; see Volume 2, Aquatic Plants, Waterhyacinth), an invasive plant species. Often nutria will
cut a plant off at the waterline and carry it to a feeding platform that they have constructed in
shallow water. Nutria also dig beneath the marsh surface and feed on the root mat.
Life History. Nutria construct nests of coarse plant material in dens within their burrows.
They reproduce throughout the year, producing two or litters annually. The female will
come into estrus every 24–26 days and stay in heat 1–4 days. Gestation is about 130 days.
Typically, 4–6 precocial young are born, although the original number of embryos may have
been much higher. Prenatal embryo losses are common, especially in cold weather and
among poorly nourished females; miscarriage rates are high. Estrus begins soon after the
female miscarries or gives birth. She will mate with several males each time she comes into
estrus.
Young are born with full coats of fur and open eyes. Within a day, they swim and feed on
plants. They will be weaned in 5–8 weeks. Nutria are sexually mature in 4–6 months. Adults
usually live in pairs, but large numbers may congregate in favorable habitat. Individuals
occupy only a small area throughout their lives, rarely traveling more than 600 ft. (180 m)
from their dens each day. Wild nutria rarely live more than three years. Young may be taken
by coyotes, foxes, raccoons, alligators, and owls; humans are the only predators of adults.
Impacts. The damage nutria do to native ecosystems is related to both their feeding and
their burrowing. Animals feeding directly on the root mat of marsh plants produce what is
known as an “eat out.” The mat binding the marsh substrate together is weakened and soil
is washed away, pockmarking the marsh with holes. In Louisiana some “eat outs” are 500 ac.
(202 ha) in size.
Deep swimming channels come to fragment the marsh and, in coastal areas, allow the
influx of salt water. The channels or “runs” create more edge habitat exposed to erosion by
wind and water. Productive wetlands can be turned into barren mudflats or open water,
destroying habitat for marshland invertebrates, fish, and birds. Nutria feeding on the rhi-
zomes of sea oat on the barrier islands of Mississippi has exposed the sand dunes to erosion.
Large burrows made by nutria weaken river banks, levees, dams, and the sides of drain-
age canals and can lead to cave-ins and erosion during heavy rains.
In man-made environments, nutria may girdle orchard trees as well as ornamental trees
and shrubs. In the 1950s, expanding populations in Louisiana became pests in rice and
sugar cane fields.
Nutria has been nominated by the IUCN as one of “100 of the World’s Worst” invasive
species.
294 n VERTEBRATES (MAMMALS)

Management. Trapping and shooting have been the most effective ways of controlling
feral populations of nutria, and high fur prices once made these viable strategies. With the
fall of prices, Louisiana established a nutria control program that included a bounty system
to encourage the harvest of 400,000 nutria a year from coastal areas. Part of that program
involves promoting nutria as a source of lowfat, high-protein meat.
The legal status of nutria varies from state to state, and influences control methods. In
Delaware, they are considered furbearers and regulated as such. In Maryland, they are listed
as “unprotected” and can be controlled as wildlife species when they cause damage to eco-
logical or economic resources. In Virginia, nutria are “nuisance” species; there is open season
all year for trapping them. In Washington State, nutria are classified as “prohibited,” mean-
ing they may not be released into the wild and may not be transported without a permit from
the Washington Department of Fish and Wildlife.

Selected References
Bertolino, S. “Myocastor coypus (Mammal).” IUCN/SSC Invasive Species Specialist Group, ISSG Global
Invasive Species Database, 2005. http://www.invasivespecies.net/database/species/ecology.asp
?si=99&fr=1&sts=.
“Exotic Aquatics of the Gulf Coast: Nutria (Myocastor coypus).” La Mer, Louisiana Sea Grant, n.d. http://
www.lamer.lsu.edu/invasivespecies/nutria/index.html.
“Invasive Species in the Chesapeake Watershed.” Chesapeake Bay Program and Maryland Sea Grant,
2002. http://www.mdsg.umd.edu/issues/restoration/non-natives/workshop/nutria.html.
Link, Russell. “Living with Wildlife: Nutria.” Washington Department of Fish and Wildlife, 2006.
http://wdfw.wa.gov/wlm/living/nutria.pdf.
“Worldwide Distribution, Spread of, and Efforts to Eradicate the Nutria (Myocastor coypus).” National
Wetlands Research Center, U.S. Geological Survey, 2008. http://www.nwrc.usgs.gov/special/
nutria/namerica.htm.
n State-by-State Occurrences of Invasive
Microorganisms, Fungi, and Animals

Includes only the species featured in the Encyclopedia.

Alabama

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease
Invertebrates
Cnidarian: Australian spotted jellyfish
Mollusks: Asian clam, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Argentine ant, Asian tiger mosquito, common bed bug, Formosan subterranean
termite, Japanese beetle, multicolored Asian lady beetle, red imported fire ant
Vertebrates
Fish: Brown trout, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Monk Parakeet, Rock Pigeon
Mammals: Black rat, feral cat, feral pig, house mouse, Norway rat, nutria

Alaska

Microorganisms
Fungi: Chytrid frog fungus
Invertebrates
Tunicates: Chain tunicate
Insect: Common bed bug
Vertebrates
Fish: Rainbow trout
Birds: Cattle Egret, European Starling, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

Arizona

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, New Zealand mud snail, quagga mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insect: Africanized honey bee, Argentine ant, Common bed bug, Formosan subterranean
termite, multicolored Asian lady beetle
Vertebrates
Fish: Bighead carp, brown trout, gizzard shad, mosquitofish, rainbow trout
Amphibians: African clawed frog, American bullfrog
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Feral burro, feral cat, feral horse, feral pig, house mouse, Norway rat

(Continued )
296 n STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS

Arkansas

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, Argentine ant, Asian tiger mosquito, common bed bug,
Formosan subterranean termite, Japanese beetle, multicolored Asian lady beetle, red
imported fire ant
Vertebrates
Fish: Bighead carp, brown trout, grass carp, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat, nutria

California

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Chytrid frog fungus, sudden oak death, white pine blister rust
Invertebrates
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, golden apple snail, New Zealand mud snail,
quagga mussel, zebra mussel
Crustaceans: Chinese mitten crab, green crab
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, Argentine ant, Brown marmorated stink bug, common
bed bug, Formosan subterranean termite, glassy-winged sharpshooter, multicolored
Asian lady beetle, red imported fire ant
Vertebrates
Fish: Bighead carp, brown trout, mosquitofish, rainbow trout, spotted tilapia
Amphibians: African clawed frog, American bullfrog
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Sparrow, Rock
Pigeon
Mammals: Black rat, feral burro, feral cat, feral goat, feral horse, feral pig, house mouse,
Norway rat

Colorado

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, New Zealand mud snail, quagga mussel,
zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Alewife, bighead carp, brown trout, gizzard shad, mosquitofish, rainbow trout
Amphibians: American bullfrog:
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Sparrow,
Rock Pigeon
Mammals: Feral cat, feral horse, feral pig, house mouse, Norway rat, nutria
STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS n 297

Connecticut

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Bryozoan: Lacy crust bryozoan
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, common periwinkle, zebra mussel
Crustaceans: Green crab, rusty crayfish
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, gypsy moth, hemlock woolly adelgid, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, mosquitofish, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Monk Parakeet, Mute
Swan, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

Delaware

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease
Invertebrates
Tunicates: Chain tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, common periwinkle
Crustaceans: Green crab
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian tiger mosquito, brown marmorated stink bug, common bed bug, gypsy
moth, hemlock woolly adelgid, Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Monk Parakeet, Mute
Swan, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat, nutria

Florida

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease
Invertebrates
Cnidarian: Australian spotted jellyfish
Mollusks: Asian clam, Asian green mussel, golden apple snail
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, Argentine ant, Asian tiger mosquito, common bed bug,
Formosan subterranean termite, multicolored Asian lady beetle, red imported fire ant
Vertebrates
Fish: Asian swamp eel, bighead carp, brown trout, lionfish, mosquitofish, rainbow trout,
spotted tilapia, walking catfish
Amphibians: Coqui (noninvasive), Cuban treefrog
Reptiles: Brown anole, Burmese python, green iguana, Nile monitor
Birds: Cattle Egret, Common Myna, Eurasian Collared-Dove, European Starling, House
Finch, House Sparrow, Monk Parakeet, Rock Pigeon
Mammals: Black rat, feral cat, feral pig, house mouse, Norway rat, nutria

(Continued )
298 n STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS

Georgia

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Asian green mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Argentine ant, Asian tiger mosquito, common bed bug, Formosan subterranean
termite, hemlock woolly adelgid, Japanese beetle, multicolored Asian lady beetle, red
imported fire ant
Vertebrates
Fish: Alewife, Asian swamp eel, brown trout, lionfish, rainbow trout
Reptiles: Brown anole
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Black rat, feral cat, feral horse, feral pig, house mouse, Norway rat, nutria

Hawai’i

Microorganisms: Avian malaria

Fungi: Chytrid frog fungus
Invertebrates
Cnidarian: Australian spotted jellyfish
Mollusks: Asian clam, giant African snail, golden apple snail, zebra mussel
Insects: Argentine ant, Asian tiger mosquito, common bed bug, Formosan subterranean
termite
Vertebrates
Fish: Asian swamp eel, bighead carp, brown trout, mosquitofish, rainbow trout
Amphibians: American bullfrog, coqui
Reptiles: Green iguana
Birds: Cattle Egret, Common Myna, European Starling, House Finch, House Sparrow,
Japanese White-eye, Rock Pigeon
Mammals: Black rat, feral cat, feral goat, feral pig, house mouse, Indian mongoose,
Norway rat

Idaho

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, New Zealand mud snail
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, mosquitofish, rainbow trout
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat, nutria

Illinois

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, quagga mussel, zebra mussel
STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS n 299

Arachnids: Honeybee tracheal mite, varroa mite

Insects: Argentine ant, Asian longhorned beetle, Asian tiger mosquito, common bed bug,
emerald ash borer, gypsy moth, Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Alewife, brown trout, gizzard shad, mosquitofish, rainbow trout, round goby, sea
lamprey, silver carp
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Monk Parakeet, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

Indiana

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian tiger mosquito, common bed bug, emerald ash borer, gypsy moth,
Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Alewife, bighead carp, brown trout, gizzard shad, mosquitofish, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

Iowa

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, quagga mussel, zebra mussel
Crustaceans: Rusty crayfish
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Bighead carp, brown trout, mosquitofish, rainbow trout
Amphibians: American bullfrog (in DeSoto NWR)
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat
Kansas

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease
Invertebrates
Mollusks: Asian clam, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian tiger mosquito, common bed bug, Japanese beetle, multicolored Asian lady
beetle
Vertebrates
Fish: Bighead carp, brown trout, gizzard shad, mosquitofish, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

(Continued )
300 n STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS

Kentucky

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, quagga mussel, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, Asian tiger mosquito, common bed bug, emerald ash
borer, hemlock woolly adelgid, Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Alewife, bighead carp, brown trout, gizzard shad, grass carp, mosquitofish, rainbow
trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

Louisiana

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease
Invertebrates
Cnidarian: Australian spotted jellyfish
Annelid worms: European earthworms
Mollusks: Zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, Argentine ant, Asian tiger mosquito, common bed bug,
Formosan subterranean termite, multicolored Asian lady beetle, red imported fire ant
Vertebrates
Fish: Bighead carp, grass carp, rainbow trout, silver carp
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Monk Parakeet, Rock Pigeon
Mammals: Black rat, feral cat, feral pig, house mouse, Norway rat, nutria

Maine

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine
blister rust
Invertebrates
Bryozoan: Lacy crust bryozoan
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, common periwinkle
Crustaceans: Green crab, rusty crayfish
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, gypsy moth, hemlock woolly adelgid, multicolored Asian lady
beetle
Vertebrates
Fish: Alewife, brown trout, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Mute Swan, Rock
Pigeon
Mammals: Feral cat, house mouse, Norway rat

Maryland

Microorganisms: Lyme disease bacterium, West Nile virus

STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS n 301

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Tunicates: Chain tunicate
Annelid worms: European earthworms
Mollusks: Asian clam
Crustaceans: Green crab, rusty crayfish
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Argentine ant, Asian tiger mosquito, brown marmorated stink bug, common bed
bug, emerald ash borer, gypsy moth, hemlock woolly adelgid, Japanese beetle,
multicolored Asian lady beetle
Vertebrates
Fish: Alewife, brown trout, mosquitofish, northern snakehead, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Mute Swan, Rock
Pigeon
Mammals: Feral cat, house mouse, Norway rat, nutria

Massachusetts

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Bryozoan: Lacy crust bryozoan
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, common periwinkle, zebra mussel
Crustaceans: Green crab, rusty crayfish
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian longhorned beetle, common bed bug, gypsy moth, hemlock woolly adelgid,
Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Alewife, brown trout, mosquitofish, rainbow trout
Amphibians: American bullfrog (on Nantucket, Martha’s Vineyard,
and Cape Cod)
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Mute Swan, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

Michigan

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, New Zealand mud snail, quagga mussel,
zebra mussel
Crustaceans: Rusty crayfish, spiny water flea
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, emerald ash borer, gypsy moth, Japanese beetle, multicolored
Asian lady beetle
Vertebrates
Fish: Alewife, brown trout, gizzard shad, mosquitofish, rainbow trout, round goby, sea
lamprey
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Mute Swan, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

(Continued )
302 n STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS

Minnesota

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, New Zealand mud snail, quagga mussel,
zebra mussel
Crustaceans: Rusty crayfish, spiny water flea
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian tiger mosquito, common bed bug, emerald ash borer, Japanese beetle,
multicolored Asian lady beetle
Vertebrates
Fish: Alewife, brown trout, mosquitofish, rainbow trout, round goby, sea lamprey
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

Mississippi

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease
Invertebrates
Cnidarian: Australian spotted jellyfish
Annelid worms: European earthworms
Mollusks: Asian clam, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, Argentine ant, Asian tiger mosquito, common bed bug,
Formosan subterranean termite, Japanese beetle, multicolored Asian lady beetle, red
imported fire ant
Vertebrates
Fish: Bighead carp, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Black rat, feral cat, feral pig, house mouse, Norway rat, nutria

Missouri

Microorganisms: West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, quagga mussel, zebra mussel
Crustaceans: Rusty crayfish
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Argentine ant, Asian tiger mosquito, common bed bug, emerald ash borer,
Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, grass carp, mosquitofish, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

Montana

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease, white pine blister rust
STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS n 303

Invertebrates
Annelid worms: European earthworms
Mollusks: New Zealand mud snail
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, mosquitofish, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Sparrow, Rock
Pigeon
Mammals: Feral cat, feral horse, house mouse, Norway rat

Nebraska

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease
Invertebrates
Mollusks: Asian clam, Chinese mystery snail, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian tiger mosquito, common bed bug, Japanese beetle, multicolored Asian lady
beetle
Vertebrates
Fish: Alewife, bighead carp, brown trout, gizzard shad, mosquitofish, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

Nevada

Microorganisms: West Nile virus

Fungi: chytrid frog fungus, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, New Zealand mud snail, quagga mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, Argentine ant, common bed bug, multicolored Asian lady
beetle
Vertebrates
Fish: Brown trout, mosquitofish, rainbow trout, spotted tilapia
Amphibians: American bullfrog
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Feral burro, feral cat, feral horse, feral pig, house mouse, Norway rat

New Hampshire

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Bryozoan: Lacy crust bryozoan
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Chinese mystery snail
Crustaceans: Green crab, rusty crayfish
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, gypsy moth, hemlock woolly adelgid, Japanese beetle

(Continued )
304 n STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS

Vertebrates
Fish: Alewife, brown trout, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

New Jersey

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, common periwinkle
Crustaceans: Green crab, rusty crayfish
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian longhorned beetle, Asian tiger mosquito, brown marmorated stink bug,
common bed bug, gypsy moth, hemlock woolly adelgid, Japanese beetle, multicolored
Asian lady beetle
Vertebrates
Fish: Asian swamp eel, brown trout, mosquitofish, rainbow trout
Amphibians: American bullfrog (Cape May NWR)
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Monk Parakeet, Mute
Swan, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

New Mexico

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, common bed bug, Formosan subterranean termite,
multicolored Asian lady beetle, red imported fire ant
Vertebrates
Fish: Brown trout, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Sparrow, Rock
Pigeon
Mammals: Feral cat, feral horse, house mouse, Norway rat, nutria

New York

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, common periwinkle, quagga mussel, zebra
mussel
Crustaceans: Green crab, rusty crayfish, spiny water flea
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian longhorned beetle, brown marmorated stink bug,
common bed bug, emerald ash borer, gypsy moth, hemlock woolly adelgid, Japanese
beetle, multicolored Asian lady beetle
STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS n 305

Vertebrates
Fish: Alewife, brown trout, mosquitofish, rainbow trout, round goby
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Monk Parakeet, Mute
Swan, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

North Carolina

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Argentine ant, Asian tiger mosquito, common bed bug, Formosan
subterranean termite, gypsy moth, hemlock woolly adelgid, Japanese beetle,
multicolored Asian lady beetle, red imported fire ant
Vertebrates
Fish: Alewife, brown trout, lionfish, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Mute Swan, Rock Pigeon
Mammals: Feral cat, feral horse, feral pig, house mouse, Norway rat, nutria

North Dakota

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease
Invertebrates
Annelid worms: European earthworms
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, rainbow trout
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Black rat, feral cat, feral horse, house mouse, Norway rat

Ohio

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, quagga mussel, zebra mussel
Crustaceans: Spiny water flea
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian tiger mosquito, common bed bug, emerald ash borer, gypsy moth,
Japanese beetle, multicolored Asian lady beetle
Vertebrates
Fish: Alewife, brown trout, mosquitofish, rainbow trout, round goby, sea lamprey
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

Oklahoma

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease

(Continued )
306 n STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS

Invertebrates
Crustaceans: Zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insect: Africanized honey bee, Argentine ant, Asian tiger mosquito, multicolored Asian
lady beetle, red imported fire ant
Vertebrates
Fish: Bighead carp, brown trout, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat, nutria

Oregon

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, sudden oak death, white pine blister rust
Invertebrates
Tunicates: Chain tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, New Zealand mud snail
Crustaceans: Green crab
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Argentine ant, brown marmorated stink bug, common bed bug, multicolored
Asian lady beetle
Vertebrates
Fish: Brown trout, mosquitofish, rainbow trout
Amphibians: American bullfrog
Birds: Cattle Egret, European Starling, House Sparrow, Monk Parakeet, Rock Pigeon
Mammals: Black rat, feral burro, feral cat, feral horse, feral pig, house mouse, Norway rat,
nutria

Pennsylvania

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, quagga mussel, zebra mussel
Crustaceans: Rusty crayfish, spiny water flea
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian tiger mosquito, brown marmorated stink bug, common bed bug, Japanese
beetle, multicolored Asian lady beetle
Vertebrates
Fish: Alewife, brown trout, gizzard shad, mosquitofish, northern snakehead, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

Rhode Island

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Bryozoan: Lacy crust bryozoan
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Chinese mystery snail, common periwinkle
Crustaceans: Green crab
STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS n 307

Arachnids: Honeybee tracheal mite, varroa mite

Insects: Common bed bug, gypsy moth, hemlock woolly adelgid, Japanese beetle,
multicolored Asian lady beetle.
Vertebrates
Fish: Brown trout, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Monk Parakeet, Mute
Swan, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

South Carolina

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Asian green mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Argentine ant, Asian tiger mosquito, common bed bug, Formosan
subterranean termite, hemlock woolly adelgid, Japanese beetle, multicolored Asian lady
beetle, red imported fire ant
Vertebrates
Fish: Alewife, brown trout, lionfish, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Mute Swan, Rock Pigeon
Mammals: Black rat, feral cat, feral pig, house mouse, Norway rat

South Dakota

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Mollusks: Zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, multicolored Asian lady beetle
Vertebrates
Fish: Bighead carp, brown trout, rainbow trout
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Feral burro, feral cat, house mouse, Norway rat

Tennessee

Microorganisms: West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Argentine ant, Asian tiger mosquito, brown marmorated stink bug, common bed
bug, Formosan subterranean termite, hemlock woolly adelgid, Japanese beetle,
multicolored Asian lady beetle, red imported fire ant
Vertebrates
Fish: Alewife, bighead carp, brown trout, grass carp, mosquitofish, rainbow trout
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Finch, House
Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat, nutria

(Continued )
308 n STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS

Texas

Microorganisms: West Nile virus

Fungi: Chestnut blight fungus, chytrid frog fungus, Dutch elm disease
Invertebrates
Mollusks: Asian clam, Chinese mystery snail, golden apple snail, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, Argentine ant, Asian tiger mosquito, common bed bug,
Formosan subterranean termite, multicolored Asian lady beetle, red imported fire ant
Vertebrates
Fish: Bighead carp, brown trout, grass carp, mosquitofish, rainbow trout
Reptiles: Green iguana
Birds: Cattle Egret, Eurasian Collared-Dove, European Starling, House Sparrow, Monk
Parakeet, Rock Pigeon
Mammals: Black rat, feral cat, feral pig, house mouse, Norway rat, nutria

Utah

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, New Zealand mud snail, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Africanized honey bee, common bed bug, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, gizzard shad, mosquitofish, rainbow trout
Amphibians: American bullfrog
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Feral burro, feral cat, feral horse, house mouse, Norway rat

Vermont

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, gypsy moth, hemlock woolly adelgid, Japanese beetle,
multicolored Asian lady beetle
Vertebrates
Fish: Alewife, brown trout, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Rock Pigeon
Mammals: Feral cat, house mouse, Norway rat

Virginia

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Tunicates: Chain tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, veined rapa whelk, zebra mussel
Crustaceans: Green crab
STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS n 309

Arachnids: Honeybee tracheal mite, varroa mite

Insects: Asian tiger mosquito, brown marmorated stink bug, common bed bug, emerald
ash borer, Formosan subterranean termite, gypsy moth, hemlock woolly adelgid,
Japanese beetle, multicolored Asian lady beetle, red imported fire ant
Vertebrates
Fish: Alewife, bighead carp, brown trout, mosquitofish, northern snakehead, rainbow
trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Monk Parakeet, Mute
Swan, Rock Pigeon
Mammals: Black rat, feral cat, feral horse, feral pig, house mouse, Norway rat, nutria

Washington

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, white pine blister rust
Invertebrates
Tunicates: Chain tunicate, colonial tunicate
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, New Zealand mud snail
Crustaceans: Green crab
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, rainbow trout
Amphibians: American bullfrog
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Black rat, feral cat, house mouse, Norway rat, nutria

West Virginia

Microorganisms: West Nile virus

Fungi: Bat white-nose syndrome, chestnut blight fungus, chytrid frog fungus, Dutch elm disease,
white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, zebra mussel
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Asian tiger mosquito, brown marmorated stink bug, common bed bug, emerald
ash borer, gypsy moth, hemlock woolly adelgid, Japanese beetle, multicolored Asian
lady beetle
Vertebrates
Fish: Alewife, bighead carp, brown trout, mosquitofish, rainbow trout
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

Wisconsin

Microorganisms: Lyme disease bacterium, West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: Asian clam, Chinese mystery snail, New Zealand mud snail, zebra mussel
Crustaceans: Rusty crayfish, spiny water flea
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, emerald ash borer, gypsy moth, Japanese beetle, multicolored
Asian lady beetle

(Continued )
310 n STATE-BY-STATE OCCURRENCES OF INVASIVE MICROORGANISMS, FUNGI, AND ANIMALS

Vertebrates
Fish: Alewife, brown trout, mosquitofish, rainbow trout, sea lamprey
Birds: Cattle Egret, European Starling, House Finch, House Sparrow, Rock Pigeon
Mammals: Feral cat, feral pig, house mouse, Norway rat

Wyoming

Microorganisms: West Nile virus

Fungi: Chytrid frog fungus, Dutch elm disease, white pine blister rust
Invertebrates
Annelid worms: European earthworms
Mollusks: New Zealand mud snail
Arachnids: Honeybee tracheal mite, varroa mite
Insects: Common bed bug, multicolored Asian lady beetle
Vertebrates
Fish: Brown trout, mosquitofish, rainbow trout
Birds: Cattle Egret, European Starling, House Sparrow, Rock Pigeon
Mammals: Feral cat, feral horse, house mouse, Norway rat

Commonwealth of Puerto Rico

Fungi: Chytrid frog fungus

Invertebrates
Cnidarian: Australian spotted jellyfish
Mollusks: Asian clam
Insects: Red imported fire ant
Vertebrates
Fish: Brown trout, gizzard shad, mosquitofish, rainbow trout?
Amphibians: Cuban treefrog
Reptiles: Burmese python, green iguana
Birds: Cattle Egret, European Starling, House Sparrow, Monk Parakeet, Rock Pigeon
Mammals: Feral cat, feral goat, house mouse, Indian mongoose, Norway rat
n Glossary
Achene. A small, dry, hard one-seed fruit.
Adventive. Refers to an introduced species that has arrived in a new habitat or environment
without the aid of humans and that has not established a self-replacing population.
Aeciospore. A fungal spore produced in an aecium. Each spore has two nuclei and is part of
a chain of spores.
Aecium. The cuplike fruiting body of some rust fungi.
Aerenchyma. Pithy respiratory tissue, common in stems of some aquatic plant species.
Agnathan. Member of the class Agnatha, the jawless fish.
Alate. Winged reproductive adult of a social insect, such as ants and termites.
Alien (species). A nonnative species. A species found beyond its normal range limits.
Synonyms: exotic, nonindigenous.
Allee effect. The consequences of low population density when the presence of too few
individuals greatly reduces reproductive success.
Alleleopathy. Condition in which one plant or species exudes chemicals that prevent the
growth of other plants in the immediate vicinity.
Altricial. Refers to recently hatched birds or other newborn animals that have closed eyes
and little or no down or fur, and that are unable to leave the nest and therefore must depend
upon the parents for food.
Anadromous. Refers to fish that spend most of their lives in salt water but ascend freshwater
streams to spawn.
Anecic. Refers to deep-burrowing earthworms that inhabit the lower layers of the soil.
Annelid worm. Any member of the phylum Annelida, the segmented worms.
Annual. A plant that germinates from seed, matures, and dies in one season.
Apical (snail). The tip of a spiraling shell.
Apomictic. Refers to a flower than does not require pollination to produce seed.
Aquatic. Refers to a plant growing primarily or entirely in water, either rooted or free-floating.
Aril. The fleshy coating around a seed.
Arthropod. Member of the phylum Arthropoda, invertebrates with exoskeletons, segmented
bodies, and jointed appendages. The phylum includes arachnids, insects, and crustaceans.
Ascospore. A type of spore bearing a single copy of each chromosome formed by sexual
reproduction in fungi in the Division/Phylum Ascomycetes.
Asexual reproduction. The multiplication of individuals without the fusion of gametes. Can
occur in fungi and animals through cell splitting, budding, cloning, or sporation. In plants,
formation of new plants without the transfer of pollen. In some plants, new individuals can
be generated vegetatively from parts of the parent plant.
Auricle. Earlike appendage at the base of some leaves, which clasps the stem.
312 n GLOSSARY

Awn. A bristle-shaped appendage on a grass.

Axis. The central line of any organ, such as a stem.
Barbel. Whiskerlike tactile organ in catfish and carp that houses taste glands and helps them
to find food in murky water.
Basidiospore. A spore bearing a single copy of each chromosome and found in fungi of the
Division/Phylum Basidiomycetes.
Beak (bivalve). The highest raised part of each valve, which is generally pointed and located
near the hinge.
Benthos. A collective term referring to organisms living on the seabed.
Bergmann’s Rule. An ecogeographic pattern wherein the higher the latitude or colder the
climate, the larger the body size of warmed-blooded animals compared to close relatives
living at lower latitudes and/or in warmer climates.
Biennial. A plant that lives for two years, usually flowering and setting seed in the second year.
Bilabiate. Refers to a corolla, two-lipped.
Biodiversity. The total variation and variability of life found in genes, species, communities,
ecosystems, and landscapes.
Biogeography. The science that studies the distribution patterns of species and the pro-
cesses that determine those patterns.
Biotype. A subset of a species with a particular set of genetic features.
Bivalve. A mollusk, such as a clam or mussel, that has its body covered by two rigid shells
joined by a hinge.
Blade. The portion of the leaf that extends from the leaf sheath, flat, folded, or with rolled
margins.
Bolt. Rapid growth of flower stalk.
Bract. A small scale-like leaf, usually associated with a flower.
Bulbil. Small bulb, usually growing from leaf axils.
Byssal threads. Filaments that some mollusks produce and use to fasten themselves to hard
surfaces.
Calyx. The leaf-like sepals that enclose the petals of a flower.
Canker. A localized area of dead tissue on the trunk or branch of a woody plant.
Cardinal teeth (bivalve). Ridges and grooves on the inner surfaces of both valves of a
bivalve near the front end of the hinge that help hold the shells in alignment.
Carton. Material made of undigested cellulose, mud, and termite saliva.
Catadromous. Refers to fish that spend most of their lives in freshwater but migrate to the
sea to breed.
Chasmogamous. Refers to flowers that open to allow cross-pollination.
Chitin. A strong, semitransparent, horny substance that is the main material composing the
exoskeletons of arthropods and the internal structures of certain other invertebrates.
Chlamydospore. Large, thick-walled resting spore of several kinds of certain fungi. It is the
part of the life cycle that allows survival during unfavorable conditions, such as excessive
drought or heat.
 GLOSSARY n 313

Cilia. Hairlike structures used by some cells to move themselves or to move food particles.
Clambering. Refers to shrubs or vines with stems that climb onto and over other plants.
Cleistogamous. Refers to flowers that do not open and are self-pollinated as buds.
Clitellum. Thickened, saddlelike section of the body of earthworms that secretes a viscous
fluid in which the worm’s eggs are deposited.
Colubrid. Any snake of the large and poorly defined group of nonvenomous snakes placed
in the family Colubridae.
Columella (snail). The central structural spine of coiled snail and whelk shells.
Community (ecological). All species living in the same area or a subset of them, such as the
bird community or the plant community.
Compound. Refers to leaves that are divided into leaflets.
Conidia. Asexual, nonmotile spores of a fungus such as chestnut blight.
Contact. Refers to herbicides that kill only the plant portions contacted.
Coppicing. Refers to trees that sprout many shoots from a cut stump.
Corm. A type of bulb.
Corolla. The petals of a flower.
Corona. A distinct circular growth between the corolla and the stamens, especially in the
milkweed family.
Culm. Stem of a grass, usually hollow.
Cuticle (insect). The exoskeleton, composed mostly of chitin.
Cuticle (plant). A protective waxy coating produced by the outermost cells of a leaf or other
aerial part of a plant.
Cyme. A broad and flat-topped determinate flower cluster, with central flowers opening
first.
DBH (Diameter at breast height). Standard way of expressing the diameter of a living tree.
There is no universal standard, however, as to what breast height is. In the United States,
DBH is usually measured at a height of 1.4 m (about 4.5 ft.) above ground.
Decumbent. Refers to a stem that is reclining or lying on the ground, but with the tip
upright.
Dehiscent. Refers to a seed capsule that opens, sometimes explosively.
Determinate. Refers to when a branch or stem ceases to grow after flowering.
Detritus. Organic debris composed of parts of plants, the remains of animals, and waste
products that accumulates on the ground or moves into water bodies from surrounding
terrestrial areas.
Diapause. A suspension of development in response to adverse environmental conditions.
Dioecious. Refers to male and female flowers being on different plants.
Disjunct. A distribution pattern in which parts of the range are noncontiguous, i.e., separated
geographically.
Drupe. A fleshy, one-seeded fruit. Druplet is one part of a berry fruit.
Ecology. The interrelationships among organisms and the nonliving aspects of the environ-
ment in which they live; the science that studies such interrelationships.
314 n GLOSSARY

Ecosystem. The totality of living and nonliving elements in a given area that function as a
unit to cycle nutrients and maintain a flow of energy.
Ecotype. A population that is adapted to a particular environment and displays characteris-
tics that set it apart from related populations but that has not evolved into a distinct species.
Emergent. Refers to aquatic plants that grow primarily above the water surface.
Entire. Refers to leaf margins that are smooth, not toothed or serrated.
Epiphyte. A plant that physically lives on another but obtains no nutrients from the host.
Bromeliads and tropical orchids are frequently epiphytic. The leaves of some epiphytic bro-
meliads fuse to form a tank in which water collects, creating prime breeding grounds for
mosquitoes and some treefrogs.
Erect. Growing upright, not sprawling or trailing.
Established (species). A species not native to a geographic area and with a self-replacing
population.
Exotic (species). Any nonnative species. Synonyms: alien, nonnative, nonindigenous.
Fasciated. Abnormal growth of a plant part, such as an inflorescence, causing it to be
twisted or incurved. Also called crested.
Floret. The individual flower of a grass, comprised of two bracts, the lemma and the palea,
and the pistil and stamens. Also the individual flower of a composite.
Follicle. A dry, dehiscent fruit or seed pod that splits open on the front.
Forb. A broad-leaved, green-stemmed, nonwoody plant. One type of herb.
Fouling (organism). Any organism that accumulates on solid surfaces in an aquatic envi-
ronment and impedes the normal mechanical functioning of the equipment or host on
which it resides.
Frass. Fine, powdery material that wood-eating insects produce as waste after digesting
plant matter.
Fruiting body. Multicellular structure of fungi that carries spore-forming bodies. When the
sexual stages of the life cycle are aerial, they are usually visible to the naked eye.
Gamete. A mature sexual reproductive cell, either sperm, pollen, or egg, that fuses with
another cell during fertilization to form a new organism.
Genotype. The total complement of genes in an individual or an entire species.
Gill rakes. Bony or cartilaginous, finger-like projections off the gill arch of fish that allow
filter-feeders to retain food particles and keep solids from entering the gill cavity. Also called
gill rakers.
Glabrous. Smooth; not rough, fuzzy, or hairy.
Glaucous. Covered with a bloom, a whitish substance that rubs off.
Gloger’s Rule. An ecogeographic pattern in which warm-blooded animals in humid envi-
ronments tend to have darker pigments in skin, feathers, or hair than close relatives living
in drier environments.
Glume. Bract on a grass that does not have associated flowers.
Gonopodium. An anal fin that on some male live-bearing fish has been modified to allow
passage of sperm and internal fertilization.
Graminoid. Herbaceous plant that includes grasses, reeds, rushes, and sedges.
 GLOSSARY n 315

Granivorous. Feeding on seeds.

Gravid. Refers to a female carrying eggs or developing young; pregnant.
Habitat. The place in which an organism lives and the physical attributes of that place.
Halophyte. Plants adapted to salty conditions.
Hammock. Slightly raised tree islands surrounded by other vegetation, usually sawgrass, in
the Everglades.
Harborage (insects). Shelter or refuge.
Haustoria. The root-like, absorbing organs of a parasitic plant.
Herb. A plant with no persistent woody stem above ground. Herbaceous.
Hibernaculum. The wintering place that shelters hibernating bats.
Holoparasite. A parasitic plant that can obtain nutrients and water in no way other than
from a host plant.
Host specific. Refers to a biological control that affects only the intended plant.
Hyphae. Long, branching threads that are the main vegetative structural feature of many
fungi. See also Mycelium.
Indehiscent. Refers to a fruit that does not split open at maturity to release seeds.
Indeterminate. Refers to when a branch or stem continues to grow after flowering.
Inflorescence. Flower stalk and how the flowers are arranged.
Injurious wildlife. Species of wild mammals, wild birds, fish, mollusks, crustaceans,
amphibians, and reptiles listed under the auspices of the Lacey Act that the secretary of
the interior has determined may be harmful to the health and welfare of humans; the inter-
ests of agriculture, horticulture, or forestry; and the welfare and survival of wildlife resources
of the United States. Such species require a permit in order to be imported or trans-
ported between states.
Instar. A developmental stage in larval insects that begins and ends with a molt until the
individual is sexually mature. Often, but not always, morphological changes occur from
one instar to the next.
Introduced species. A species that has been transported, either deliberately or unintention-
ally, by humans to a location it had not previously occupied.
Introduction. The transport and release into a free-living state of a nonnative species.
Invasive species. (1) A nonnative species that is currently spreading rapidly or that has
done so in the past; (2) “An alien species whose introduction does or is likely to cause eco-
nomic or environmental harm or harm to human health” (Executive Order 13112).
Involucre. Whorl of small leaves or bracts beneath a flower or flower cluster, especially
thistles.
Irruption. A sudden, rapid increase in numbers in an animal population, usually accompa-
nied by the migration of many individuals.
Keratin. A fibrous structural material composed of protein in skin, hair, nails, feathers, and
beaks of vertebrates.
Lateral line. A sense organ in aquatic organisms such as fish and amphibians that is used to
detect movement and vibration. Commonly visible as a faint line running lengthwise down
each side.
316 n GLOSSARY

Lateral teeth (bivalve). Elongated, interlocking projections along the hinge line of a shell
that prevent the two valves from sliding against each other when the shell is closed.
Lemma. The lower of the two bracts that enclose the flower in a grass.
Lenticel. Corky cells in the bark of trees that allow air to penetrate into the interior.
Ligule. Ring on the inside of a grass leaf where the blade meets the sheath.
Lore (bird). The area between the eye and the bill on the side of the head
Macroalgae. Large, multicellular algae. Seaweed.
Macrophyte. Any plant large enough to be visible to the naked eye.
Margins. The edges of leaves or leaflets.
Membranous. Thin, parchment-like texture.
Meristem. The growing point of a plant.
Monocarpic. Refers to a plant in which the growing point of the plant becomes the flower-
ing stem, and the plant dies after flowering.
Monoecious. Refers to both female and male flowers on the same plant.
Monospecific. Consisting of one species.
Monotypic. Consisting of one genotype or ecotype, such as a clone.
Mucilaginous. Soft, moist, sticky, or gel-like.
Mycelium. A mass of branching, filamentous hyphae through which a fungus absorbs
nutrients and decomposes plant material.
Naı̈ve. Previously unexposed to a pathogen and therefore having no natural immunity.
Native. In this encyclopedia, describes species, habitats, or ecosystems known to have
existed in North America prior to European colonization. Considered by many to be the
natural elements of a continent’s biodiversity that would occur even if humans had not set-
tled the region.
Native transplant. A species that is native to the country or region in question but has been
transported beyond its natural range limits.
Naturalized (species). Refers to a nonindigenous species that is able to sustain itself repro-
ductively in the wild outside of cultivation, and has become a functioning member of a
native ecosystem.
Nitrogen fixer. A plant that, with the help of certain soil bacteria that form nodules on its
roots, can utilize atmospheric nitrogen.
Node. Joint in a stem, usually where the leaves grow.
Nonindigenous (species). A species that is not native to the place in which it now occurs.
Nonnative (species). An alien, exotic, or nonindigenous species.
Noxious (weed). A plant specified by law as being especially undesirable, troublesome, and
difficult to control.
Nuisance (species). According to the Nonindigenous Nuisance Aquatic Prevention and
Control Act of 1990, an alien aquatic species that “threatens the diversity or abundance of
native species or the ecological stability of infested waters, or commercial, agricultural or
recreational activities dependent upon such waters.”
 GLOSSARY n 317

Nymph (insect). The immature form of insects that undergo a gradual and incomplete
metamorphosis before reaching the adult stage. A nymph resembles the adult form and
never enters a pupal stage. It becomes an adult after the final molt.
Opercle. A bony plate that supports the gill covers of fishes, especially the most posterior
one.
Organelle. Any of the distinct structures within a cell that performs a specific and vital
function.
Outcompete. When a plant or animal displaces another plant or animal by being a better
competitor for some resource.
Palate (plant). A bulge in the lower lip of a figwort (Scrophulariaceae) flower that closes off
the throat.
Palea. The upper of the two bracts that enclose the flower in a grass.
Palpus. A jointed organ for touching or tasting attached to a mouthpart in arthropods.
Panicle. A loose, irregularly compound inflorescence with flowers on pedicels.
Pantropical. Found throughout the tropics.
Pappus. Appendage to a flower in the sunflower family (Asteraceaceae), such as a thistle,
which may remain attached to the fruit; may be bristled, plume-like, or scaly.
Paradioecious. Refers to male and female flowers occurring on separate plants, but any
individual can develop flowers of either gender.
Parietal callus. In some snails, a thickened deposit on the margin of the aperture and the
wall of the body whorl closest to the central spine (columella). It is often smooth and glossy
and may be adorned with raised ribs or wrinkles.
Parthenogenesis. A form of asexual reproduction in which growth and development of
embryos occurs without fertilization of the ovum by sperm.
Pathway. The means by which a species arrives at a new region.
Pedicel. Stalk that supports an individual flower or fruit.
Peduncle. Stalk that supports a flower cluster.
Perennial. A plant that lives for more than one season, although aerial parts may die back.
Perigynium. The inflated sac, which encloses the ovary in Carex species.
Petiole. A leaf stalk.
pH. The measure of acidity or alkalinity of soil or water. Using a logarithmic scale, it
describes the amount of hydrogen ions in the solution.
Pharyngeal teeth (fish). Teeth in the throat located at the back of a fish’s head.
Pheromone. Chemicals released by an organism into its environment to communicate with
other members of its own species. Some pheromones are alarm signals, while others attract
individuals to food or to a mate.
Phloem. Plant tissues that conduct foods made in the leaves to all other parts of the plant.
Photoperiod. The number of hours of daylight.
Phreatophyte. Refers to plants with roots that extend into the water table.
Phytoplankter. A tiny, usually microscopic plant that is part of the plankton.
318 n GLOSSARY

Pinna. The primary division, or branch, of a pinnate leaf. Leaflets are on the pinna. Plural,
pinnae.
Pinnate. Describes compound leaves that have pairs of leaflets on either side of a stalk.
Evenly pinnate leaves have an even number of paired leaflets and terminate in a pair.
Oddly pinnate leaves have an uneven number of leaflets and terminate in a single leaflet.
Pinnatifid. Describes leaves that resemble pinnately compound leaves, but with lobes that
do not reach the midrib of the leaf.
Plankton. A collective term referring to all organisms that drift in open water unable to
move under their own power against tides and currents.
Pollard. Describes the method of severely pruning tree limbs back to the trunk or to a main
branch.
Polychaete. A member of the Polychaete class of annelid worms characterized by having
bristles on each body segment. Also called bristle worms and lugworms.
Postemergent. Describes a herbicide that affects growing plants.
Precocial. Referring to hatchlings or newborns that are born with their eyes open, fully
feathered, or furred, and that leave the nest a short time after birth or hatching,
Preemergent. Describes a herbicide that prevents seeds from germinating.
Pronotum. In insects, the upper surface of the first segment of the thorax.
Propagule. In animals, the minimum number of individuals of a species capable of coloniz-
ing a new area. This may be fertilized eggs, a mated female, a single male and a single female,
or a whole group of organisms, depending upon the biological and behavioral requirements
of the species. In plants, a propagule is whatever structure functions to reproduce the spe-
cies, such as a seed, spore, stem, or root cutting.
Protist. A microorganism that is either single-celled or multicellular, but lacking specialized
tissues, and has the genetic information carried in a cell nucleus.
Pubescence. Describes plant parts covered with soft, fine hairs.
Pupa. In the development of those insects that undergo complete metamorphosis, the life
stage that immediately precedes the adult stage. Some pupae remain inside the exoskeleton
of the final larval instar, but others are encased in a cocoon or chrysalis.
Pustule. A blister-like spot.
Pycnia. A flask-shaped or conical fruiting body of a rust fungus that develops below the
epidermis of the host and bears pycniospores.
Pycniospore. A spore produced in a pycnia of a rust fungus. It fuses with a hypha of the
opposite mating type to produce the sexual generation.
Raceme. Inflorescence with all the individual flowers on a single axis.
Rachilla. A small or secondary axis or rachis, especially the axis that bears florets in sedges
and grasses.
Rachis. An axis bearing flowers or leaflets.
Recurved. Bent backward.
Rhizoid (fungi). A structure that functions like a root to anchor the fungus and absorb
nutrients. Rhizoid also releases enzymes that break down organic matter.
 GLOSSARY n 319

Rhizome. A root structure below the soil surface, distinguished from a root by having nodes;
can grow shoots that produce new plants. Also called a rootstock.
Rootcrown. Top portion of a root, often containing dormant buds that sprout.
Rosette. Arrangement of leaves radiating from a central point.
Ruderal. Waste places.
Samara. An indehiscent winged fruit.
Saprotroph. Any organism that gains energy and nutrients from dead organic material.
Savanna. Grassland with scattered trees.
Scrambling. Sprawling or climbing over other plants.
Sebaceous gland. A gland in the skin that secretes an oily substance to lubricate the skin
and hair.
Semievergreen. Remaining green only in warm climates or sheltered locations.
Senesce. To grow old, turn brown. Senescence.
Sepal. A leaf-like bract that encloses a flower or flower bud. Sepals form the calyx.
Sessile (plant). Refers to flowers or leaves attached directly to stems, without pedicels or
petioles.
Settle (mollusks and crustaceans). The process by which larvae leave the plankton stage of
their life and attach to a substrate.
Sexual reproduction. The formation of new individuals from the union of two gametes, an
ovum and a sperm. In the higher plants it takes places with the transfer of pollen from a male
flower to a female flower.
Sheath. A leaf structure that surrounds and encloses a grass stem.
Shrub. Woody perennial smaller than a tree, usually with several stems.
Silicle. A small seed pod in the Mustard family.
Silique. A seed pod in the Mustard family.
Simple. Refers to leaves that are not divided or compound.
Species. A group of individuals of the same kind that can interbreed and produce viable
offspring.
Spike. A simple inflorescence with sessile flowers on a single axis. Also branch of a grass
inflorescence.
Spikelet. Secondary spike, especially a grass structure that includes glumes and florets, the
cluster of grass flowers.
Spirochete. A bacterium of the phylum Spirochaete, distinguished by its spirally twisted form.
Sporangium. Structure in which spores are formed; spore case. Plural, sporangia.
Sporation. Spore formation.
Stellate. Refers to plant hairs, star-shaped.
Stipule. The basal appendage of a petiole.
Stolon. Root stem on the soil surface that roots at nodes; may produce new plants from
sprouts; also called runners.
Stromata. The connective tissue framework or support of cells or organisms.
320 n GLOSSARY

Subdioecious. Male and female flowers usually restricted to separate plants.

Submergent. Refers to aquatic plants, or parts, that grow completely underwater.
Submersed. Refers to aquatic plants that grow primarily underwater. Flowering parts may
be at or slightly above the water surface. May be free-floating or rooted.
Subshrub. A shrub in which the upper branches die back during the unfavorable seasons.
Substrate. Substance in which plants are rooted; can be soil, sand, alluvium, mud, or rock.
Surfactant. Substance added to a herbicide to help the chemicals adhere to the foliage.
Systemic. Refers to herbicides that are absorbed into plant tissues and translocated through-
out the plant.
Talus. Cone- or fan-shaped slope of loose rocks at the base of a cliff.
Telium. The pimplelike cluster of spore cases that is produced by rust fungi.
Ternate. In sets of three.
Thorax (insects). The central of three main segments of an insect’s body: the segment
between the head and the abdomen.
Tree. A woody plant with one main trunk.
Turion. A scaly, young shoot or sucker on a root or tuber.
Two-ranked. Referring to alternate arrangement of leaves; leaves are on opposite sides of the
stem, in the same geometric plane.
Umbel. Often flat-topped inflorescence resembling an umbrella, with individual pedicels
rising from a common point.
Uredinia. A reddish, pimplelike structure on the tissue of a plant infected by a rust fungus.
Vegetative reproduction. Formation of new plants from pieces of the parent plant, such as
stems, leaves, rhizomes, and stolons. Also called asexual reproduction.
Vent (reptile). Cloaca. The common cavity into which the intestinal, genital, and urinary
tracts end.
Vine. Plant whose stem requires support; can be trailing on the ground or climbing by twin-
ing, tendrils, or other means.
Whorl. Arrangement of leaves in a circle around the stem, three or more leaves at one node.
Xylem. Tissue that conducts water and dissolved minerals from the roots to all other parts of
a plant, provides mechanical support, and forms the wood of trees and shrubs.
Zooanthellae. Protozoans that live symbiotically in some jellyfishes as well as corals and
other marine organisms.
Zooid. One of the individual organisms composing a colonial animal, such as a bryozoan.
Zooplankter. Any animal, single-celled or multicelled, that is part of the plankton.
Zoospore. An asexual spore produced by some fungi that can move around by using a
tail-like appendage (flagellum).
Zygomorphic. In plants, irregular corollas that can be equally divided into mirror-image
halves in only one plane, such as pea or orchid flowers.
n Index
Page numbers in boldfaced type refer to a main entry in the encyclopedia and “t” indicates table.
Acarapis woodi. See Honeybee Tracheal Mite
Acclimatization societies, 238, 245
Acentria ephemerella, biological control (plants)
Eurasian watermilfoil, 324–25
Aceria imperata, biological control (plants)
cogongrass, 447
Aceria malherbae, biological control (plants)
field bindweed, 609
Aceria salsolae, biological control (plants)
prickly Russian thistle, 413
Achatina fulica. See Giant African Snail
Acidotheres tristis. See Common Myna
Acremonium zonaatum, biological control (plants)
waterhyacinth, 342–43
Adedarach species. See Chinaberry
Adelges tsugae. See Hemlock Woolly Adelgid
Adoretus sinicus. See Chinese rose beetle
Adventive species, xiv (v. 1)
Aecidium mori var. broussonetia, biological
control (plants)
paper mulberry, 564
Aedes albopictus. See Asian Tiger Mosquito
Aedes eagypti. See Yellow fever mosquito
Aegopodium podagraria. See Goutweed
African Clawed Frog, xxivt (v. 1), xxvt (v. 1), 18,
19, 201–5
state-by-state occurrences, 295, 296
African feathergrass, 460–61
noxious designation, 653, 665, 666, 668,
669, 670, 671, 691
African fountain grass. See Crimson
Fountain Grass
African foxtail grass. See Buffelgrass
Africanized Honey Bee, xix (v. 1), xxvt (v. 1),
106–10
as varroa mite host, 103, 108
African pyle. See Giant Salvinia
Agonopterix nervosa, biological control (plants)
brooms, 501
Agreement on the Application of Sanitary and
Phytosanitary Measures, 703
Agrilus aurichalceus. See rose stemgirdler
Agrilus hyperici, biological control (plants)
common St. Johnswort, 361
Agrilus planipennis. See Emerald Ash Borer
Agropyron repens. See Quackgrass
AHB. See Africanized Honey Bee
Ailanthus. See Tree of Heaven
Ailanthus altissima. See Tree of Heaven
Ailanthus glandulosa. See Tree of Heaven
Ailanthus peregrine. See Tree of Heaven
Air potato, 607, 660
Aizoaceae, stone plant family, 383
Akala, 536, 538, 657
Akalakala, 536, 656
Akebia quinata. See Chocolate Vine
Akepa, as affected by avian malaria, 248
Alabama jumper (earthworm), 49
ALB. See Asian Longhorned Beetle
Albizia julibrissin. See Silk Tree
Albizzia julibrissin. See Silk Tree
Albonia peregrina. See Tree of Heaven
Albugo. See white leaf rust
Alelaila tree. See Chinaberry
Aleppo grass. See Johnsongrass
Alewife, xviii (v. 1), xxiiit (v. 1), 157–60,
190, 193
state-by-state occurrences, 296, 298–310
Alfalfa dodder, 611, 660
Alfalfa dwarf (disease), Glassy-Winged
Sharpshooter as vector, 137
Alien Animals (Laycock), xxxi (v. 1)
Alien, definition, xiii (v. 1)
Alkali bulrush. See cosmopolitan bulrush
Allegheny blackberry, 524, 656
Alleleopathy
Australian pine, 542
chinaberry, 553
Chinese lespedeza, 352
dyer’s woad, 364
exotic bush honeysuckles, 506
Johnsongrass, 471
kikuyugrass, 480
lantana, 521
quackgrass, 488
tree of heaven, 588
yellow starthistle, 429
Alley cat. See Feral Cat
I-2 n INDEX
Alliaria alliaria. See Garlic Mustard
Alliaria officinalis. See Garlic Mustard
Alliaria petiolata. See Garlic Mustard
Allorhogas species, biological control (plants)
velvet tree, 592
Alosa pseudoharengus. See Alewife
Alternaria solani, tropical soda apple
as host, 533
Altica carduorum, biological control (plants)
Canada thistle, 348
Alvars, 639
Amberique bean, 623, 660
American alligator, 663
as potentially affected by Nile monitor, 228
and West Indian marsh grass, 491
American barberry, 512, 656
American beach grass, 432, 435, 653
American bittersweet, 630–31, 632, 660
American Bullfrog, xiii (v. 1), xviii (v. 1),
xxvt (v. 1), xxvi (v. 1), 205–8
as chytrid frog fungus vector, 19
ISSG 100 worst invaders, 207, 711
state-by-state occurrences, 295, 296, 298,
299, 301, 303, 304, 306, 308, 309
American bumblebee, as varroa mite host, 103
American climbing fern, 598–99, 660
American crocodile, 663
as potentially affected by Nile monitor, 228
and West Indian Marshgrass, 491
American cupscale, 489, 653
American eel, 160, 183, 191
American elm, 23
cultivars, 25
American hogpeanut, 623, 660
American Robin, 661
and exotic bush honeysuckles, 506
and multiflora rose, 525
American wisteria, 645, 660
Aminopyralid, chemical control (plants)
common St. Johnswort, 361
Ampelopsis brevipedunculata.
See Porcelainberry
Ampelopsis glandulosa var. brevipedunculata.
See Porcelainberry
Ampelopsis glandulosa ‘Elegans,’ 633, 634
Ampelopsis heterophylla. See Porcelainberry
Amphibians, 201–14
ISSG 100 worst invaders, 711
Amur Honeysuckle 502–8, 614
noxious designation, 666, 668, 671, 692.
See also Exotic Bush Honeysuckles
Amur peppervine. See Porcelainberry
Amynthas agrestis. See Alabama jumper
Anabasis glomerata. See Halogeton
Anacardaceae. See sumac family
Anchored waterhyacinth, 339, 649
noxious designation, 665, 666, 668, 669,
670, 671, 689
Andean pampas grass. See Jubata Grass
Andes grass. See Jubata Grass
Andropogon vimineum. See Japanese Stilt Grass
Anecic worms, 50
Angle worm, 49. See European Earthworms
Anisantha tectorum. See Cheatgrass
Anitimicrobial chemicals, African Clawed
Frog, 204
Anjan grass. See Buffelgrass
Annelid worms, 48–53, 216
Anolis distichus. See Bark anole
Anolis equestris. See Knight anole
Anolis garmani. See Jamaican giant anole
Anoplophora glabripennis. See Asian Longhorned
Beetle
Antelope bitterbrush, 442, 656
Anthonomus tenebrosus, biological control
(plants), tropical soda apple, 534
Antiblemma acclinalis, biological control (plants)
Koster’s curse, 518
Antitoxicum rossicum. See Pale Swallow-Wort
Aphalara itadori, biological control (plants)
Japanese knotweed, 390
Aphids, biological control (plants), 663
Canada thistle, 347
exotic bush honeysuckle, 507
giant reed, 466
Japanese hops, 621
kikuyugrass, 481
tropical soda apple, 634
Aphtae epizooticae, Wild Pig as host, 280
Apiaceae. See carrot family
Apion fuscirostre, biological control (plants)
brooms, 501
Apion species, biological control (plants)
velvet tree, 592
Apion ulicis, biological control (plants)
gorse, 511
Apis mellifera scutellata. See Africanized
Honey Bee
Aplocera plagiata, biological control (plants)
common St. Johnswort, 361
Aporrectodea caliginosa. See European
Earthworms
Applesnail. See Golden Applesnail
Apthona species, biological control (plants)
leafy spurge, 398
Aquarium watermoss. See Giant Salvinia
Aquatic invertebrates, ISSG 100 worst
invaders, 711

Aquatic plants, 321–43
American species invasive abroad, 696
ISSG 100 worst invaders, 710
noxious designation, 689
Aquatic soda apple. See wetlands nightshade
Aquatic weeds, grass carp control of, 173, 174
Arachnids, 99–106
Araliacaeae. See ginseng family
Archanara geminipuncta, biological
control (plants)
common reed, 451
Archips asiaticus, biological control (plants)
chocolate vine, 596
Argentine Ant, xxii (v. 1), xxvt (v. 1), 110–13
ISSG 100 worst invaders, 711
state-by-state occurrences, 295–310
Aristolochia. See pipevine and wooly
Duchman’s pipe
Arizona cottontop, 436–37, 653
Arizona wheatgrass, 483, 653
Artimpaza argenteonota, biological
control (plants)
Asiatic colubrina, 496
Artipus floridanus, biological control (plants)
Australian pine, 543
Arundo. See Giant Reed
Arundo donax. See Giant Reed
Arundo selloana. See Pampas Grass
Arundo versicolor. See Giant Reed
Asclepiadaceae. See milkweed family
Asian applesnail. See Chinese Mystery Snail
Asian bittersweet. See Oriental Bittersweet
Asian Clam, xxiiit (v. 1), xxix (v. 1),
53–56, 77
state-by-state occurrences, 295–310
Asian euonymus scale, biological
control (plants)
winter creeper, 643
Asian Green Mussel, xxvt (v. 1), 56–58
state-by-state occurrences, 297, 298, 307
Asian gypsy moth, 138
Asian honey bee, as varroa mite host, 102
Asian lady beetle. See Multicolored Asian
Lady Beetle
Asian Longhorned Beetle, xix (v. 1), xxvt (v. 1),
113–16
ISSG 100 worst invaders, 711
state-by-state occurrences, 299, 301, 304
Asian nakedwood. See Asiatic Colubrina
Asian rapa whelk. See Veined Rapa Whelk
Asian snakeroot. See Asiatic Colubrina
Asian Swamp Eel, 160–62
state-by-state occurrences, 297, 298, 304
Asian tapeworm, grass carp as host, 174
INDEX n I-3
Asian Tiger Mosquito, xxiv (v. 1), xxvt (v. 1),
116–20
ISSG 100 worst invaders, 711
state-by-state occurrences, 295–310
Asian water monitor, 226
Asiatic bittersweet. See Oriental Bittersweet
Asiatic clam. See Asian Clam
Asiatic Colubrina, 493–96
impacts, 683
noxious designation, 666, 692
pathways of introduction, 675, 677
uses of, 496
Asiatic Sand Sedge, 432–35
impacts, 683
noxious designation, 666, 668, 691
pathways of introduction, 433, 675, 676
Asiatic tear-thumb. See Mile-A-Minute
Asteraceae. See aster family; sunflower family
Athel tamarisk, 581, 583, 658
Atlantic cordgrass. See Cordgrasses and Their
Hybrids
Atlantic ivy, 603–4, 660
Atlantic shipworm. See Naval Shipworm
Atlantic wisteria, 645, 660
Atomacera petroa, biological control (plants)
velvet tree, 592
Atrazine, chemical control (plants)
cheatgrass, 443
medusahead, 485
mile-a-minute, 628
Australian Pine, 540–44
and Asiatic colubrina, 495
impacts, 681, 682, 684
noxious designation, 666, 692
pathways of introduction, 541, 675
uses of, 543
Australian pine borer, biological
control (plants)
Australian pine, 543
Australian river oak, 540, 541, 542, 658
Australian Spotted Jellyfish, 45–48
state-by-state occurrences, 295, 297, 298,
300, 302, 310
Austromusotima camptozonale, biological control
(plants)
climbing ferns, 601
Autumn olive, 568, 570
noxious designation, 570, 668, 669, 672, 692
Avian Malaria, xix (v. 1), xxvii (v. 1),
1–3, 24, 248
bird reservoir, 234
state-by-state occurrences, 298
Awapuhi kahili. See Kahili Ginger
Azolla species, 327
I-4 n INDEX

Bacterial leaf scorch (Xylalla fastidiosa), 22, 137 kudzu, 625

Bacterial leaf scorch, and Dutch Elm Disease, 22 leafy spurge, 398–99
English Ivy as host, 605 melaleuca, 561
Bagous affinis, biological control (plants) purple loosestrife, 417
hydrilla, 334 rattlebox, 529
Bagpod, 528, 650 tamarisk, 585
Bahamian brown anole. See Brown Anole toadflax, 426
Bald brome, 440, 653 tropical soda apple, 534
noxious designation, 666, 691 velvet tree, 592
Ball nut. See Water Chestnut water chestnut, 338
Banded mystery snail, 59 Begomovirus. See tobacco leaf curl
Bangasternus orientalis, biological Bell’s honeysuckle, 502–8
control (plants) noxious designation, 666, 668, 669, 691,
yellow starthistle, 430 692. See also Exotic Bush Honeysuckles
Banker horse. See Feral Horse Belut eel. See Asian Swamp Eel
Barberry family, 512 Berberidaceae. See barberry family
Barbwire Russian thistle, 411, 650 Berberis japonica. See Japanese Barberry
noxious designation, 665, 666, 690 Berberis sinensis. See Japanese Barberry
Bark anole, 215 Berberis thunbergia var. atropurpurea.
Barley, 483, 653 See Japanese Barberry
Basal bark application, herbicides, xx (v. 2) Berberis thunbergii. See Japanese Barberry
Basiliscus vittatus. See Brown basilisk Berberis x ottawensis, 512
Bat nut. See devil pod Bergmann’s Rule, xxxi (v. 1), 312, 714
Batrachochytrium dendrobatidis. See Chytrid Bermuda grass, 479, 654
Frog Fungus Big Cypress National Preserve
Bat White-Nose Syndrome Fungus, xix (v. 1), West Indian marsh grass, 490
11–14 melaleuca, 559
state-by-state occurrences, 297, 301–2, 304, Big Cypress National Preserve, spotted
306, 308, 309 tilapia, 197
Bay cedar, 495, 658 Burmese python, 218
Beach clustervine, 546, 660 melaleuca, 559
Beach layia, 386, 650 Big eye herring. See Alewife
Beach panic grass, 432, 435, 653 Bighead. See hyperparathyroidism
Beach she-oak. See Australian pine Bighead Carp, xxvt (v. 1), 163–66, 195
Beachstar, 546, 654 state-by-state occurrences, 295–300,
Bead tree. See Chinaberry 302–3, 306–9
Bean aphid, Canada thistle host, 347 Bignonia tomentosa. See Princess Tree
Bean stalk borer, Canada thistle host, 347 Big sage. See Lantana
Bearberry honeysuckle, 505, 656 Big taper. See Common Mullein
Beardless wheatgrass, 487, 654 Biogeography, xiii (v. 1), xxxii (v. 1)
Bedbug. See Common Bed Bug Biological control (plants). See species entries
Bee colonies destruction, by varroa mite, 104–5 Birds, 228–59
Beefwood. See Australian pine Bishop’s goutweed. See Goutweed
Beetles, biological control (plants) Bishop’s weed. See Goutweed
Asiatic colubrina, 496 Bitter panicum, 432
Brazilian peppertree, 547–48 Black acacia. See Rattlebox
Canada thistle, 348 Black carp, 172
climbing ferns, 601 Black dog-strangle vine. See Black Swallow-Wort
common St. Johnswort, 361 Blackfin cisco, 193
garlic mustard, 372 Black-hooded Parakeet. See Nanday Conure
hydrilla, 334 Black imported fire ant, 152
Japanese hops, 621 Black-legged tick
Japanese knotwood, 390 as Lyme disease vector, 4, 5, 6
Koster’s curse, 518 Black mangrove cichlid. See Spotted Tilapia

Black Rat, xviii (v. 1), xxvit (v. 1), 259–62, 287
ISSG 100 worst invaders, 261, 712
state-by-state occurrences, 295–310
Blacksage. See Lantana
Black snail. See Chinese Mystery Snail
Black spinytail iguana, 222–23
Black stem grain rust, 513
Black Swallow-Wort, 636–40
impacts, 678, 680, 684
noxious designation, 666, 668, 669, 671,
693. See also Swallow-Worts
Bladderpod. See bagpod
Blady grass. See Cogongrass
Blue toadflax. See Canada toadflax
BMSB. See Brown Marmorated Stink Bug
Bohemian knotweed. See giant knotweed
Boiga irregularis. See Brown tree snake
Bootanelleus orientalis, biological control (plants)
Australian pine, 543
Boreioglycaspis melaleucae, biological
control (plants)
melaleuca, 561
Borrelia burgdorferi. See Lyme Disease bacterium
Boston ivy, 603, 633, 660
Bothriocephalus opsarichthydis, 174
Botrylloides diagensis, 40
Botrylloides perspicuum, 40
Botrylloides violaceus. See Chain Tunicate
Botryllus schlosseri, 40
Botrytis cinerea, biological control (plants)
fire tree, 557
Botrytis cinerea, fire tree control, 557
Bowfin, 182, 183
Branch herring. See Alewife
Brachypterolus pulicarius, biological
control (plants)
toadflax, 426
Branched tearthumb, 600, 650
Brassicaceae. See mustard family
Brazilian elodea. See Brazilian waterweed
Brazilian oak, 540, 541–42, 543, 658
Brazilian Peppertree, 495, 544–48, 560
impacts, 682, 684, 685
ISSG 100 worst invaders, 710
noxious designation, 666, 671, 692
pathways of introduction, 673
uses of, 547
Brazilian satintail, 445, 654
noxious designation, 665, 666, 668, 669,
670, 671, 691
Brazilian waterhyacinth, 440, 649
Brazilian waterweed, 332, 334, 649
Breea arvensis. See Canada Thistle
Breea incana. See Canada Thistle
INDEX n I-5
Bridal broom, 498, 656
Bristletips, 515, 589–90, 656
British Petroleum oil spill (2010), 329
Broadleaf toadflax, 424, 650
Broadleaved pepperweed. See Perennial
Pepperweed
Broad-leaved toadflax. See Dalmatian
Toadflax
Broghammerus reticulates, 220
Bromacil, chemical control (plants)
Brazilian peppertree, 547
Bromeliads, 495, 660, 661
Bromus tectorum. See Cheatgrass
Broncograss. See Cheatgrass
Broomleaf toadflax, 424, 650
Brooms, 496–502, 509
impacts, 679, 681, 683
noxious designation, 666, 667,
670, 671, 692
pathways of introduction, 673, 675, 676
uses of, 502
Brotogeris chiriri. See Yellow-chevroned
Parakeet
Brotogeris versicolurus. See White-winged
Parakeet
Broussonetia papyrifera. See Paper Mulberry
Brown Anole, xviii (v. 1), 214–17
state-by-state occurrences, 297, 298
Brown basilisk, 223
Brown Marmorated Stink Bug, xix (v. 1),
xxvt (v. 1), xxx (v. 1), 120–23
state-by-state occurrences, 296, 297, 304–6,
307, 309
Brown rat. See Norway Rat
Brown tree snake, xxiv (v. 1), xxvii (v. 1), 210
Brown Trout, xxivt (v. 1), 159, 166–68, 185
ISSG 100 worst invaders, 168, 711
state-by-state occurrences, 295–310
Bruchus atronotatus, biological control (plants)
Brazilian peppertree, 548
Bryozoans, 36–38
Bubulcus ibis. See Cattle Egret
Buckthorn family, 493
Buckwheat family, 387, 626
Budgerigars, 250
Buffalobur, 533, 650
noxious designation, 667, 670, 671, 690
Buff-backed heron. See Cattle Egret
Buffelgrass, 435–39
impacts, 679, 683
noxious designation, 665, 691
pathways of introduction, 675
Bulb panicgrass, 470, 654
Bulbous buttercup. See Fig Buttercup
I-6 n INDEX

Bull thistle, 345–46, 348–49, 401, 403, 650
noxious designation, 666, 667, 668, 669,
670, 671, 690
Burbot, 182, 183
Bur cucumber, 619, 660
Bureau of Land Management (BLM)
feral burros, 262, 264–65
feral horses, 273, 274–75
Burgdorfer, Dr. Willy, 5
Burmese Python, xviii (v. 1), xxivt (v. 1),
217–21
state-by-state occurrences, 297, 310
Burning, physical control (plants)
Australian pine, 543
Canada thistle, 348
cheatgrass, 442–43
Chinese lespedeza, 352
common reed, 450–51
English ivy, 605
exotic bush honeysuckles, 506
garlic mustard, 371
giant reed, 465
gorse, 511
Japanese barberry, 514
Japanese dodder, 613
Japanese stilt grass, 469
medusahead, 484
quackgrass, 488
tamarisk, 584
tree of heaven, 588
yellow starthistle, 430
Burning bush, 380, 641, 643, 650, 656
noxious designation, 666, 660, 671, 690
pathways of introduction, 673
Bursting heart, 641, 643, 656
Bush currant. See Velvet Tree
Bush honeysuckles. See Exotic Bush
Honeysuckles
Bush morning glory. See western
morning glory
Bush muhly, 437, 654
Bushy white solanum. See turkey berry
Butter and eggs. See Yellow Toadflax
Buttercup family, 366
Butterflies
and Canada thistle, 348
and garlic mustard, 371
and Japanese stiltgrass, 468,
and swallow-worts, 639
Butterflies, biological control (plants)
fire tree, 557
garlic mustard, 371
Bythotrephes cederstroemi, 96
Bythotrephes longimanus. See Spiny Water Flea
California broom, 498
California Clapper Rail, 661
and cordgrass, 457
California cordgrass, 453, 454–55, 456, 457,
654. See also Cordgrasses and Their
Hybrids
California peppertree. See Peruvian peppertree
California satintail, 445, 654
noxious designation, 666, 691
Californian thistle. See Canada Thistle
Calophasia lunula, biological control (plants)
toadflax, 426
Caloptilla sp. nr. schinella, biological
control (plants)
fire tree, 557
Camasey. See Koster’s Curse
Canada germander, 414, 650
Canada Thistle, 344–49, 401
impacts, 678, 679, 682, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 677
Canada thistle stem weevil, xxiv (v. 1), 348
Canada toadflax, 424, 650
Canadian honeysuckles, 505, 656
Canadian waterweed, 332, 649
Canary Island St. Johnswort, 359, 656
Canby’s mountain-lover, 643, 656
Candleberry myrtle. See Fire Tree
Cane. See Common Reed
Cane ti, 515, 589–90, 656
Cane tibouchina. See cane ti
Cannabidaceae. See hemp family
Caper spurge, 396, 650
Capra hircus. See Feral Goat
Caprifoliaceae. See Honeysuckle family
Carcinus maenas. See Green Crab
Carderia draba. See Hoary Cress
Cardaria latifolia. See Perennial Pepperweed
Carduus acanthoides. See plumeless thistle
Carduus arvensis. See Canada Thistle
Carduus nutans. See Musk Thistle
Carduus pycnocephalus. See Italian thistle
Carduus species, 344, 345, 399–404, 651, 652
noxious designation, 605, 607
pathways of introduction, 677. See also Musk
Thistle
Carduus tenuflorus. See slender-flowered thistle
Carex kobomugi. See Asiatic Sand Sedge
Caribbean fruit fly, 663
strawberry guava as host, 578, 579
Carolina horsenettle, 533, 650
noxious designation, 665, 666, 667,
669, 690

Carolina Parakeet, xxix (v. 1), 251
Carpobrotus edulis. See Ice Plant
Carpodacus mexicanus. See House Finch
Carposina bullata, biological control (plants)
Koster’s curse, 518
Carrizo. See Giant Reed
Carrot family, 372, 376
Carrot weed. See Carrotwood
Carrotwood, 548–51
impacts, 684
noxious designation, 666, 692
pathways of introduction, 673, 676
Cartwheel-flower. See Giant Hogweed
Cassida rubiginosa, biological control (plants)
Canada thistle, 348
Casuarina. See Australian Pine
Casuarinaceae. See casuarina family
Casuarina equisetifolia. See Australian Pine
Casuarina litorea. See Australian Pine
Casuarina littorea. See Australian Pine
Casuarina family, 540
Catalpa, 566, 659
Caterpillars, biological control (plants)
giant reed, 466
Japanese stiltgrass, 468
lantana, 521
Cat facing, fruit deformation, 122
Cattle
forage for, 437, 441, 445, 470, 479,
480, 489
Johnsongrass, 472
leafy spurge, 398
plants toxic to, common St. Johnswort, 360
toadflax, 425, 521
Cattle Egret, xiv (v. 1), xxi (v. 1), 228–32
state-by-state occurrences, 295–310
Cattley guava, 576
Cayuga Lake, and Eurasian
watermilfoil, 325
Celandine, 367, 650
Celandine poppy, 367, 650
Celastraceae. See staff-tree family;
staff-vine family
Celastrus articulatus. See Oriental Bittersweet
Celastrus orbiculatus. See Oriental Bittersweet
Celastrus sepiarius. See Asiatic Colubrina
Cenchrus ciliaris. See Buffelgrass
Cenchrus glaucus. See Buffelgrass
Cenchrus setaceus. See Crimson Fountain Grass
Centaurea biebersteinii. See Spotted Knapweed
Centaurea maculosa. See Spotted Knapweed
Centaurea solstitialis. See Yellow Starthistle
Centaurea stoebe. See Spotted Knapweed
Ceonothus asiaticus. See Asiatic Colubrina
INDEX n I-7
Ceonothus capsularis. See Asiatic Colubrina
Cercopagis pengoi. See Fishhook water flea
Cercospora rodmanii, biological control (plants)
waterhyacinth, 343
Ceutrhynchus litura. See Canada thistle
stem weevil
Ceutrhynchus species, biological control (plants)
garlic mustard, 372
Ceutrhynchus trimaculatus, biological
control (plants)
musk thistle, 403
Chaetococcus phragmitis. See legless red mealybug
Chaetorellia species, biological control (plants)
yellow starthistle, 431
Chain sea squirt. See Chain Tunicate
Chain Tunicate, xxiv (v. 1), xxvt (v. 1),
39–41, 42
state-by-state occurrences, 295–97, 300,
301, 303, 304, 306, 308, 309
Chamaespecia species. See clearwing moths
Channa argus. See Northern Snakehead
Channeled apple snail. See Golden Apple Snail
Chaparral false bindweed. See western
morning glory
Character displacement, feeding adaptation, 243
Charru mussel, 56
Cheatgrass, 439–43
impacts, 678, 679, 685
and medusahead, 484
noxious designation, 666, 691
pathways of introduction, 676, 677
uses of, 443
Cheeseberry. See Yellow Himalayan Raspberry
Cheilosia corydon, biological control (plants)
musk thistle, 403
Chemical control (plants). See species entries
Chenopodiaceae. See goosefoot family
Cherokee rose, 523, 656
pathways of introduction, 673
Cherry guava. See Strawbery Guava
Chestnut bark disease fungus. See Chestnut
Blight Fungus
Chestnut Blight Fungus, xxvit (v. 1), xxviii (v. 1),
14–17, 34
state-by-state occurrences, 295–310
Chewing disease. See nigropalallidal
encephalomalacia
Chinche. See Common Bed Bug
Chilean iceplant. See sea fig
Chilo phragmitella, biological control (plants)
common reed, 451
Chinaberry, 551–54
impacts, 681, 684
noxious designation, 692
I-8 n INDEX
pathways of introduction, 673
uses of, 554
China tree. See Chinaberry
Chinese bush clover. See Chinese Lespedeza
Chinese-glysine. See Wisteria
Chinese honeysuckle, 614, 660. See also
Japanese Honeysuckle
Chinese Lespedeza, 349–53
impacts, 680, 682
noxious designation, 666, 667, 690
pathways of introduction, 676
Chinese Mitten Crab, xxiiit (v. 1), 86–89
ISSG 100 worst invaders, 711
state-by-state occurrences, 296
Chinese Mystery Snail, xxiiit (v. 1), 58–61
state-by-state occurrences, 295–97, 299–310
Chinese packing grass. See Japanese Stilt Grass
Chinese rose beetle, biological control (plants)
velvet tree, 592
Chinese sumac. See Tree of Heaven
Chinese tamarisk. See Tamarisk
Chinese turtledove, and lantana, 520, 661
Chinese wisteria. See Wisteria. See also Rattlebox
Chinese yam, 603, 607, 660
Chlopyralid, chemical control (plants)
wisteria, 647
Chlorflurenol, chemical control (plants)
ice plant and crystalline ice plant, 386
Chlorsulfuron, chemical control (plants)
Canada thistle, 348
dyer’s woad, 365
pepperweed and hoary cress, 409
Chocolate Vine, 594–97
impacts, 684
pathways of introduction, 673
uses of, 597
Christmasberry. See Brazilian Peppertree
Chukar Partridge, 661
and cheatgrass, 443
and Medusahead, 484
Chrysobothris tranquebarica. See Australian
pine borer
Chrysolina species, biological control (plants)
common St. Johnswort, 361
Chytonix segregate, biological control (plants)
Japanese hops, 621
Chytrid fungus. See Chytrid Frog Fungus
Chytrid Frog Fungus, xv (v. 1), 13, 18–21
African clawed frog as host, 204
American bullfrog as host, 207
state-by-state occurrences, 295–10
Chytridomycosis, 13, 204, 207
Cicadallid cotton pest, and Japanese
honeysuckle, 617
Cichla ocellaris. See Peacock cichlid
Cimex lectularius. See Common Bed Bug
Cimex hemipterus, 124
Cinnamon vine, 603, 660
Cipangopaludina chinensis malleata. See Chinese
Mystery Snail
Cipangopaludina japonica. See Japanese
mystery snail
Cirsium arvense. See Canada Thistle
Cirsium setosum. See Canada Thistle
Cirsium species, 401
Cirsium vulgare. See bull thistle
Cissus brevipedunculata. See Porcelainberry
Clarias batrachus. See Walking Catfish
Clastoptera undula, biological control (plants)
Australian pine, 543
Clearwing moths, biological control (plants)
leafy spurge, 398
Cleonis pigra, Canada thistle control, 349
Clethodim, chemical control (plants)
quackgrass, 488
Cletus schmidti, biological control (plants)
mile-a-minute, 629
Clidemia. See Koster’s Curse
Clidemia crenata. See Koster’s Curse
Clidemia elegans. See Koster’s Curse
Clidemia hirta. See Koster’s Curse
Climbing euonymus. See Winter Creeper
Climbing fern family, 597
Climbing ferns, 597–602
impacts, 680, 684
noxious designation, 665, 667, 693
pathways of introduction, 673
Climbing milkweed. See Black Swallow-Wort
Climbing nightshade, 532, 637, 660
Climbing prairie rose, 524, 656
Climbing spindleberry. See Oriental Bittersweet
Clitocybe tabescens, biological control (plants)
Australian pine, 543
Clopyralid chemical control (plants)
Canada thistle, 348
Chinese lespedeza, 352
common mullein, 352
spotted knapweed, 421
yellow strarthistle, 430
Clusiaceae. See mangosteen family
Cnidarians, 45–48
Coachella Valley Preserve
tamarisk eradication, 584
Coastal sand spurge, 434, 650
Coast she-oak. See Australian Pine
Cobicula fluminea. See Asian Clam
Coccinella septempunctata. See Seven-spotted
lady beetle

Cochlearia draba. See Hoary Cress
Cockroach berry, 532–33, 656
Cocostroma myconae, biological control (plants)
velvet tree, 592
Codium fragile tomentosoides. See Dead man’s
fingers
Coffee colubrina, 493, 656
Cogon grass. See Cogongrass
Cogongrass, 443–47
impacts, 679, 683
pathways of introduction, 675
ISSG 100 worst invaders, 710
noxious designation, 665, 666, 667, 668,
669, 670, 671, 691
Colchis ivy, 604, 660
Coleophora klimeschiella, biological control
(plants)
prickly Russian thistle, 413
Coleophora parthenica, biological control (plants)
halogeton, 383
prickly Russian thistle, 413
Colletotrichum gloesporioides, biological control
(plants)
Koster’s curse, 518
kudzu, 625
Colletotrichum gloesporioides f. sp. miconiae,
biological control (plants)
velvet tree, 592
Colonial ascidian. See Colonial Tunicate
Colonial sea squirt. See Colonial Tunicate
Colonial Tunicate, xxiv (v. 1), xxvt (v. 1),
42–45
state-by-state occurrences, 296, 300, 303,
304, 306, 309
Colorado potato beetle, and tropical soda apple
as host, 533–34, 663
Colubrina asiatica. See Asiatic Colubrina
Columba livia. See Rock Pigeon
Columnar cactus, 495, 658
Common barberry, 512–13, 656
noxious designation, 666, 668, 669, 692
Common Bed Bug, xix (v. 1), xxvt (v. 1),
xxvt (v. 1), 123–27
state-by-state occurrences, 295–310
Common colubrina. See Asiatic Colubrina
Common coqui. See Coqui
Common cordgrass. See Cordgrasses and Their
Hybrids; noxious designation, 670, 671,
691; pathways of introduction, 675
Common dodder, 611, 660
Common elderberry, 374, 551, 656
Common goatweed. See Common St. Johnswort
Common gorse. See Gorse
Common guava, 577, 578, 579, 659
INDEX n I-9
Common hops, 619, 621, 660
Common hornwort. See coontail
Common ice plant. See Crystalline Ice Plant
Common iguana. See Green Iguana
Common Mullein, xiii (v. 2), 353–57
impacts, 682, 685
pathways of introduction, 674, 675
noxious designation, 666, 667, 690
uses of, 357
Common Myna, xxiiit (v. 1), 1, 232–34
ISSG 100 worst invaders, 234, 712
lantana, 520
state-by-state occurrences, 297
strawberry guava, 578
and velvet tree, 591
Common parsnip, 373, 650
Common Periwinkle, 61–63
state-by-state occurrences, 297, 300, 301,
304, 306
Common pigeon. See Rock Pigeon
Common platanna. See African Clawed Frog
Common Reed, 447–51
and Giant Reed, 463
impacts, 683
noxious designation, 665, 666, 668, 670,
671, 691
pathways of introduction, 677
uses of, 451
Common saltwort. See Prickly Russian Thistle
Common salvinia, 327, 649, 328, 329
Common St. Johnswort, 358–62
impacts, 678, 679, 680, 682
in medicine, 361
noxious designation, 666, 668, 669, 670,
671, 672
pathways of introduction, 675
Common toadflax. See Yellow Toadflax
Contact herbicides, xx (v. 2)
Convolvulaceae. See morning glory family
Convolvulus ambigens. See Field Bindweed
Convolvulus arvensis. See Field Bindweed
Convolvulus incanus. See Field Bindweed
Coontail, 322, 649
Copal. See Brazilian Peppertree
Copal-tree. See Tree of Heaven
Copper chelate, chemical control (plants)
waterhyacinth, 342
Copper sulfate biological control (plants)
waterhyacinth, 342
Coptotermes formosanus. See Formosan
Subterranean Termite
Coqui, xxvi (v. 1), xxx (v. 1), 208–11
ISSG 100 worst invaders, 210, 711
state-by-state occurrences, 297, 298
I-10 n INDEX

Coquı́ común. See Coqui Crystalline Ice Plant, 383–87

Coral honeysuckle, 615, 641, 660 impacts, 682
Coralberry, 505, 656 noxious designation, 690
Cordgrass. See Cordgrasses and Their Hybrids pathways of introduction, 677
Cordgrasses and Their Hybrids, 63, 274, 293, Ctenopharyngodon idella. See Grass Carp
452–56 Ctenosaurus similis. See Black spinytail iguana
American species invasive abroad, 696 Cuban brown anole. See Brown Anole
impacts, 682, 683 Cuban nakedwood, 493, 657
noxious designation, 670, 671, 691, 692 Cuban Treefrog, xx (v. 1), xxviii (v. 1), xxxii (v. 1),
pathways of introduction, 675, 676, 677 211–14, 227
Corn bind. See Field Bindweed state-by-state occurrences, 297, 310
Corn thistle. See Canada Thistle Cucullia verbasci. See mullein moth
Cortaderia. See Jubata Grass Cucumber, 619
Cortaderia argentea. See Pampas Grass Culex quinquefasciatus. See Southern house
Cortaderia atacamensis. See Jubata Grass mosquito
Cortaderia dioica. See Pampas Grass Cultivation, physical control (plants). See tilling
Cortaderia jubata. See Jubata Grass Cupania anacardioides. See Carrotwood
Cortaderia selloana. See Pampas Grass Cupania anacardioides var. parvifolia. See
Cosmopolitan bulrush, 455, 654 Carrotwood
Cotton States Exposition (1884), Waterhyacinth Cupaniopsis anacardioides. See Carrotwood
introduction, 340 Currants, White Pine Blister Rust alternate host,
Couch grass. See Quackgrass 30, 32, 34
Council of Europe, and Water Chestnut, 338 Curvulara lunata, biological control (plants)
Cow parsnip. See common parsnip West Indian marsh grass, 492
Coypu. See Nutria Cuscutaceae family. See morning glory family
Crasimorpha infuscate, biological control (plants) Cuscuta japonica. See Japanese Dodder
Brazilian peppertree, 548 Cutleaf toothwart, 371, 650
Crassostrea gigas. See Japanese oyster Cutting or mowing, physical control (plants)
Crater Lake National Park, and white pine brooms, 501
blister rust, 34 cheatgrass, 442
Crawdad. See Rusty Crayfish chocolate vine, 596
Crawfish. See Rusty Crayfish dyer’s woad, 365
Cream lily. See yellow ginger English ivy, 605
Creeper. See Porcelainberry giant reed, 465
Creeping Charlie. See field bindweed; ground ivy gorse, 511
Creeping Jenny. See field bindweed Japanese barberry, 514
Creeping thistle. See Canada Thistle Japanese dodder, 613
Creeping wild rye. See Quackgrass Japanese hops, 621
Crested wheatgrass, 485, 487, 654 Japanese knotweed, 390
Cricotopus myriophylli. See milfoil midge Japanese stilt grass, 468–69
Crimson beauty. See Japanese Knotweed Johnsongrass, 472
Crimson bottlebrush, 558, 657 kudzu, 625
Crimson Fountain Grass, 458–62 mile-a-minute, 628
noxious designation, 667, 691 multiflora rose, 525
pathways of introduction, 673 oriental bittersweet, 632
Crinkleroot, 371, 650 rattlebox, 529
Cristulariella pyramidalis. See zonate leafspot swallow-worts, 639
Cronartium ribicola. See White Pine Blister Rust toadflax, 426
Crustaceans, 86–99 winter creeper, 643
American species invasive abroad, 696 wisteria, 647
Cryphonectria parasitica. See Chestnut Blight yellow starthistle, 430
Fungus Cyanophyllum magnificum. See Velvet Tree
Cryophytum crystallinum. See Crystalline Cygnus olor. See Mute Swan
Ice Plant Cynanchum medium. See Pale Swallow-Wort
 INDEX n I-11

Cynanchum rossicum. See Pale Swallow-Wort tamarisk, 585

Cyperaceae. See sedge family
Cyphocleonus achates, biological control (plants)
spotted knapweed, 421
Cypress spurge, 396, 650
Cytisus monspessulana. See French Broom
Cytisus scoparius. See Scotch Broom
Cytisus striatus. See Portuguese Broom
Crytobagous salviniae, biological
control (plants)
giant salvinia, 330
Dalmatian toadflax, 421–26
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 674
Dark-green white-eye, 246
Daubentonia punicea. See Rattlebox
Davis Pond Freshwater Diversion Project
giant salvinia, 329
Dead man’s fingers (green algae), 37
Deer, and leafy spurge, 397
Deer mice, 281, 662
Deer tick, as Lyme disease vector, 4, 5, 6
Deformed wing virus (DWV), bee virus, 105
Dendrobaena octaedra. See European Earthworms
Dendryphiella broussonetiae, biological
control (plants)
paper mulberry, 564
Dengue fever, xix (v. 1), xx (v. 1), 7, 119
Dense-flowered cordgrass, noxious designation,
671, 691
pathways of introduction, 676, 677. See also
Cordgrasses and Their Hybrids
Depressed shrubverbena. See pineland lantana
Devil firefish, 175, 176, 176
Devil pod, 336, 649
Devil’s apple. See cockroach berry
Devil’s fig. See turkey berry
Devils-grass. See Quackgrass
Devil’s guts. See Field Bindweed
Devil’s hair. See Japanese Dodder
Devil’s tail tear-thumb. See Mile-A-Minute
Diatraea succharalis, biological control (plants)
giant reed, 466
Dicamba, chemical control (plants)
Australian pine, 543
gorse, 511
Japanese honeysuckle, 617
multiflora rose, 525
musk thistle, 403
prickly Russian thistle, 413
spotted knapweed, 421
strawberry guava, 578
tropical soda apple, 534
Didemnid. See Colonial Tunicate
Didemnum vexillum. See Colonial Tunicate
Diffuse knapweed, 419, 650
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
Digging out, hand pulling, or bulldozing,
physical control (plants)
Asiatic colubrina, 495
buffelgrass, 438
cordgrasses, 457
dyer’s woad, 364
giant hogweed, 375
ice plant, 386
Japanese barberry, 514
Japanese hops, 621
Japanese stilt grass, 468
Kahili ginger, 394
mile-a-minute, 628
porcelainberry, 635
tamarisk, 584
velvet tree, 592
Diorhabda elongate. See saltcedar leaf beetle
Dioscorea species, 607, 660
Diplodia natalensis, biological control (plants)
Australian pine, 543
Diquat, chemical control (plants)
giant salvinia, 329
Dittander. See Perennial Pepperweed
Dodder, 386. See also Japanese Dodder
Dog grass. See Quackgrass
Dog rose, 523–24, 657
Dog-strangling vine. See Pale Swallow-Wort
Dolichos hirsutus. See Kudzu
Dolichos lobatus. See Kudzu
Dollar leaf plant. See prostrate tickrefoil
Domesticated livestock, xviii (v. 1), xxiv (v. 1)
Dorosoma cepedianum. See Gizzard Shad
Downy brome. See Cheatgrass
Downy chess. See Cheatgrass
Drake, J. A., xxxi (v. 1)
Dreissena polymorpha. See Zebra Mussel
Dreissena rostriformis bugensis. See Quagga mussel
Drooping brome. See Cheatgrass
Drummond rattlebox, 528–29, 657
Dutch Elm Disease Fungi, xix (v. 1), xxvi (v. 1),
21–25, 115, 130
state-by-state occurrences, 295–310
Dutchman’s pipe, 603, 660
Dwarf euonymus. See Winter Creeper
Dwarf gorse, 509, 657
Dwarf honeysuckle. See European fly
honeysuckle
I-12 n INDEX
Dwarf St. Johnswort, 359, 650
Dyer’s Woad, 362–66
impacts, 678, 680, 682
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 675, 677
uses of, 365
EAB. See Emerald Ash Borer
Early chess. See Cheatgrass
Early saxifrage, 370, 650
Eastern mosquito fish. See Mosquitofish
Ebola, xix (v. 1)
Ecological impacts, xxvi–xxvii (v. 1)
in natural and semi-natural ecosystems,
xxvii–xxix (v. 1). See also species entries
Ecology of Biological Invasions of North American
and Hawaii (Mooney and Drake),
xxxi (v. 1)
“Ecology of Biological Invasions” (ICSU),
xxxi (v. 1)
Ecology of Invasions by Animals and Plants,
The (Elton), xxx (v. 1)
Economic impacts, xxix–xxxi (v. 1).
See also species entries
Edible Periwinkle. See Common
Periwinkle
Eleutherodactylus coqui. See Coqui
Eleutherodactylus planirostris.
See Greenhouse frog
Eggleaf spurge, 396, 650
Eggs and bacon. See Yellow Toadflax
Eichhornia crassipes. See Waterhyacinth
Eichhornia speciosa. See Waterhyacinth
Elaeagnaceae. See oleaster family
Elaeagnus augustifolia. See Russian Olive
Elaeagnus hortensis. See Russian Olive
Elaeagnus iliensis. See Russian Olive
Elaeagnus umbellate. See autumn olive
Elaeodendron fortunei. See Winter Creeper
Eleocharus dulcis, 335
Elephant grass, 459, 460, 654
noxious designation, 691
Elm yellows, 22
Elongate paulownia, 566, 568, 659
Elton, Charles S., xx (v. 1)
Elymus caput-medusae. See Medusahead
Elymus repens. See Quackgrass
Elytrigia repens. See Quackgrass
Elytrigia vaillantiana. See Quackgrass
Emerald Ash Borer, xxvit (v. 1), xxix (v. 1),
127–31
state-by-state occurrences, 299–302, 305,
309, 310
Emerging infectious diseases, xix (v. 1)
Empoasca biguttula, Japanese honeysuckle as
host, 617
Empress tree. See Princess Tree
Enchanted Lake (O’ahu), giant salvinia, 329
Endothall, chemical control (plants)
hydrilla, 334
“English House Sparrow Has Arrived in Death
Valley, The An Experiment in Nature”
(Grinnell), xxxi (v. 1)
English ivy, 602–6
impacts, 684, 685
noxious designation, 670, 671, 693
pathways of introduction, 673, 681
and winter creeper, 641
English Sparrow. See House Sparrow
Epigeic worms, 51
Epirrhoe sepergressa, biological control (plants)
Japanese hops, 621
Episimus utilis, biological control (plants)
Brazilian peppertree, 548
Equus asinus. See Feral Burro
Equus caballus. See Feral Horse
Eriocheir japonicus. See Japanese mitten crab
Eriocheir sinensis. See Chinese Mitten Crab
Erophora cardui, biological control (plants)
Canada thistle control, 348
Erysiphe cichoracearum, biological
control (plants)
common mullein, 357
Erysium alliaria. See Garlic Mustard
Eteobalea species, biological control (plants)
toadflax, 426
Eucerocoris suspectus, biological
control (plants)
melaleuca, 561
Eucalyptus family, 557
Eucerocoris suspectus. See leaf-blotching bug
Euhrychiopsis lecontei, biological
control (plants)
Eurasian watermilfoil, 324–25
Eulalia. See Japanese Stilt Grass
Eulalia viminea. See Japanese Stilt Grass
Eunectes murinus, 220
Eunectes notaeus, 220
Euonymus. See Winter Creeper
Euonymus fortunei. See Winter Creeper
Euonymus species. See Winter Creeper
Euphorbiaceae. See spurge family
Euphorbia esula. See Leafy Spurge
Euphorbia virgata. See Leafy Spurge
Eurasian Collared-Dove, 234–37
state-by-state occurrences, 295, 296,
297–300, 302–8

Eurasian Watermilfoil, 173, 321–26, 334, 336
grass carp, 173, 325
impacts, 680, 681, 682, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671, 689
pathways of introduction, 677
Eurasian wild boar. See Feral Pig
European bindweed. See Field Bindweed
European Earthworms, 48–53
state-by-state occurrences, 296–310
European fly honeysuckle, 504–5, 614, 657
European green crab. See Green Crab
European gypsy moth. See Gypsy Moth
European honey bee, xix (vol 1), 99, 102, 103,
105, 106, 107, 108, 109
European rose chalicid, biological control
(plants)
multiflora rose control, 526
European shore crab. See Green Crab
European Starling, xviii (v. 1), xxiiit (v. 1),
xxx (v. 1), 237–40
ISSG 100 worst invaders, 240, 712
and oriental bittersweet, 631
state-by-state occurrences, 295–310
European swallow-wort. See Pale Swallow-Wort
European Tree Sparrow, 244
European wand loosestrife, 414, 657
European water chestnut. See Water Chestnut
European wild boar. See Feral Pig
Eustenopus villosus, biological control (plants)
yellow starthistle, 431
Evecliptopera decurrens, biological control (plants)
chocolate vine, 596
Everglades
threats from invasive species,
xvi (v. 1), 162, 197, 199, 218, 219,
220, 223, 490
Australian pine, 541, 542–43
Brazilian peppertree, 545, 546
climbing ferns, 599, 600
melaleuca, 558, 559, 560
West Indian marsh grass, 490
Everglade Snail Kite, 661
and waterhyacinth, 342,
and West Indian marshgrass, 491
Everglades National Park, threats from invasive
species, xvi (v. 1), xviii (v. 1)
Asian swamp eel, 162
Burmese python, 218, 219, 220–21
green iguana 223
spotted tilapia, 197
walking catfish, 199
Executive Order 13112 (February 3, 1999),
xiii (v. 1), xv (v. 1), 703
INDEX n I-13
Exotic Bush Honeysuckles, 121, 502–8
impacts, 681, 683
and Japanese honeysuckle, 614
noxious designation, 666, 668, 669,
671, 692
pathways of introduction, 673, 674,
675, 676
Fabaceae. See pea family
Fallopia japonica. See Japanese Knotweed
False peacock fly, biological control (plants)
yellow starthistle, 431
False poinciana. See Rattlebox
Faya bush. See Fire Tree
Fayatree. See Fire Tree
Feathertop, 460, 654
Feathery pennisetum. See missiongrass
Federal Noxious Weed Act, xv (v. 1), 701
Felis silvestris catus. See Feral Cat
Feral Burro, 262–65
state-by-state occurrences, 295, 296, 303,
306–8
Feral Cat, xxivt (v. 1), xxxv (v. 1), 265–68
ISSG 100 worst invaders, 268, 712
state-by-state occurrences, 295–310
Feral Goat, xviii (v. 1), xxiiit (v. 1), 268–71
ISSG 100 worst invaders, 712
state-by-state occurrences, 296, 298
Feral hog/swine, 275
and tropical soda apple, 533
Feral Horse, xxiiit (v. 1), 271–75
state-by-state occurrences, 295, 296, 298,
303–10
Feral Pig, xviii (v. 1), xxiiit (v. 1), 275–81
fire tree, 556
ISSG 100 worst invaders, 280, 712
state-by-state occurrences, 295, 296,
297–300, 302, 303, 305, 306, 307, 308,
309, 310
and strawberry guava, 578
Feral pigeon. See Rock Pigeon
Fergusonina species, biological control (plants)
melaleuca, 561
Ficaria ranunculoides. See Fig Buttercup
Ficaria verna. See Fig Buttercup
Field Bindweed, 606–10
impacts, 678, 679, 684, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 693
pathways of introduction, 673, 675
Field morning glory. See Field Bindweed
Field thistle. See Canada thistle
Fig Buttercup, 366–68
impacts, 682
I-14 n INDEX

noxious designation, 668, 690 Floating fern. See Giant Salvinia

pathways of introduction, 673 Floating fern family, 326
Figwort family, 421, 422, 565, 719 Floating waterhyacinth. See Waterhyacinth
Fire, physical control (plants). See burning Florida apple snail, 68
Firebush. See Fire Tree Florida elodea. See Hydrilla
Fire hazard Florida holly. See Brazilian Peppertree
brooms, 500 Florida Keys National Marine Sanctuary, 178
climbing ferns, 600 Florida thatch palm, 659
cogongrass, 446 and Asiatic colubrina, 495
common reed, 450 Flowered sage. See Lantana
crimson fountain grass, 462 Fluazifop, chemical control (plants)
giant reed, 465 cogongrass, 447
gorse, 511 jubata grass, 477
Johnsongrass, 472 pampas grass, 477
jubata grass, 477 quackgrass, 488
kikuyugrass, 480 Fluridone, chemical control (plants)
lantana, 521 Eurasian watermilfoil, 324
medusahead, 484 giant salvinia, 329
melaleuca, 560 hydrilla, 334
prickly Russian thistle, 412 Fluroxypry, chemical control (plants)
Russian olive, 572 lantana, 521
Firetree. See Fire Tree Flying carp. See Silver Carp
Fire Tree, 248, 554–57 Forbs, 344–431
ISSG 100 worst invaders, 710 American species invasive abroad, 695
noxious designation, 666, 667, 693 ISSG 100 worst invaders, 710–11
pathways of introduction, 673, 676, noxious designation, 690–91
Fireweed, 414, 651 Formosan Subterranean Termite, xix (v. 1),
Fish, 157–201 xxiv (v. 1), xxvt (v. 1), 131–34
American species invasive ISSG 100 worst invaders, 134, 711
abroad, 697 state-by-state occurrences, 295–98, 300,
ISSG 100 worst invaders, 711 302, 305, 307, 308
Fishhook water flea, 97, 711 Fosamine, chemical control (plants)
Five-leaf akebia. See Chocolate Vine multiflora rose, 525
Five-stamen tamarisk, 579, 580, 581. Fountain grass, impacts, 679, 683
See also Tamarisk noxious designations, 667, 691.
Flannel leaf. See Common Mullein See also Crimson Fountain Grass
Flannel mullein. See Common Mullein Fragrant honeysuckle. See winter honeysuckle
Flaxweed. See Yellow Toadflax French broom, 496, 498, 499
Flies, biological control (plants) noxious designation, 666, 667, 670, 692.
Brazilian peppertree, 548 See also Brooms
Canada thistle, 348 French tamarisk, 579, 580, 582.
climbing ferns, 601 See also Tamarisk
common reed, 451 Freshwater herring, 157
hydrilla, 334 Fringecup, 370, 651
lantana, 521 Frog’s-bit family, 331
melaleuca, 561 FST. See Formosan Subterranean Termite
mile-a-minute, 629 Fuitour’s-grass. See Leafy Spurge
musk thistle, 403 Fund for Animals, animal welfare group, 271
Russian knapweed, 421 Fungi, 11–35
strawberry guava, 578 American species invasive abroad, 697
velvet tree, 592 Fungi, biological control (plants)
waterhyacinth, 342 Asiatic sand sedge, 439
yellow Himalayan raspberry, 539 Australian pine, 543
yellow starthistle, 431 buffelgrass, 439
 INDEX n I-15

Canada thistle, 349 Giant African land snail. See Giant African Snail
cheatgrass, 443 Giant African Snail, 64–67
chocolate vine, 596 state-by-state occurrences, 298
cogongrass, 447 Giant air plant, 600, 651
common mullein, 357 Giant Asian dodder. See Japanese Dodder
common reed, 447 Giant cow-parsnip. See Giant Hogweed
cordgrasses, 458 Giant Hogweed, 372–76
dyer’s woad, 365 impacts, 678, 681, 682
Eurasian watermilfoil, 325 noxious designation, 665, 666, 668, 669,
fire tree, 557 670, 671, 690
Japanese hops, 621 pathways of introduction, 673, 676
kikuyugrass, 481 Giant knotweed, 388, 651,
Koster’s curse, 518 noxious designation, 666, 670, 671, 690
kudzu, 625 Giant Reed, 449, 462–66
paper mulberry, 564 impacts, 679, 681, 683, 686
silk tree, 575 ISSG 100 worst invaders, 710
tree of heaven, 589 noxious designation, 671, 691
tropical soda apple, 533 pathways of introduction, 673, 675
velvet tree, 592 uses of, 465. See also Common Reed
waterhyacinth, 342–43 Giant reedgrass. See Common Reed
West Indian marshgrass, 492 Giant Salvinia, 326–30
winter creeper, 643–44 impacts, 681, 682, 685
yellow starthistle, 431 noxious designation, 665, 666, 668, 669,
Furze. See Gorse 670, 671
Fusarium nivale, biological control (plants) pathways of introduction, 674, 677
cheatgrass, 443 Giant tree frog. See Cuban Treefrog
Fusarium oxysporum, biological control (plants) Giant whiteweed. See Perennial Pepperweed
tree of heaven, 589 Giant wild pine, 600, 651
Fusarium oxysporum f. perniciosum. Gill-over-the-ground. See ground ivy
See mimosa wilt Ginger family, 391
Ginger-lily. See white ginger
Galarhoeus esula, biological control (plants). Ginseng family, 602
See Leafy Spurge Gizzard Shad, 164, 165, 168–71
Galerucella species, biological control (plants) state-by-state occurrences, 295, 296, 299,
purple loosestrife, 417 300, 302, 303, 306, 308, 310
water chestnut, 338 Glacier National Park, and white pine blister
Gallerucida bifasciata, biological control (plants) rust, 34
Japanese knotweed, 390 Glassy-Winged Sharpshooter, xxvit (v. 1),
Gallina de palo, 221 134–38
Gambusia affinis. See Mosquitofish state-by-state occurrences, 296
Gambusia holbrooki. See Mosquitofish Globe artichoke, 651
Garlic Mustard, 51, 369–72, 507 and Canada thistle, 348
impacts, 682 Gloger’s Rule, xxxi (v. 1)
noxious designation, 665, 666, 668, 669, Glorybush, 515, 590, 657
670, 671, 690 and Koster’s curse, 515
pathways of introduction, 675, 676 Glut herring. See Alewife
Gateway National Recreation Area, 435 Glyphosate, chemical control (plants)
Genista junceum. See Brooms Asiatic sand sedge, 435
Genista monspessulana. See French Broom Australian pine, 543
Geomyces destructans. See Bat White-Nose Brazilian peppertree, 547
Syndrome Fungus brooms, 501
Georgia bully, 600, 657 buffelgrass, 439
Geraldton carnation weed, 396, 651 Canada thistle, 348
German trout. See Brown Trout carrotwood, 550
I-16 n INDEX
cheatgrass, 443
chinaberry, 554
Chinese lespedeza, 352
chocolate vine, 596
climbing ferns, 601
cogongrass, 446
common mullein, 356
common reed, 451
English ivy, 605
exotic bush honeysuckles, 506
field bindweed, 609
fig buttercup, 368
fire tree, 557
garlic mustard, 371
giant hogweed, 375
giant reed, 465
giant salvinia, 329
gorse, 511
goutweed, 379
halogeton, 382
ice plant, 386
Japanese barberry, 514
Japanese honeysuckle, 617
Japanese hops, 621
Japanese knotweed, 390
Japanese stiltgrass, 469
Johnsongrass, 472
jubata grass, 477
kikuyugrass, 481
Koster’s curse, 518
kudzu, 625
lantana, 521
leafy spurge, 398
medusahead, 484
melaleuca, 561
mile-a-minute, 628
multiflora rose, 525
musk thistle, 403
Oriental bittersweet, 632
pampas grass, 477
papermulberry, 564
perennial pepperweed, 409
porcelainberry, 635
prickly Russian thistle, 413
princess tree, 568
purple loosestrife, 417
quackgrass, 488
rattlebox, 529
silk tree, 575
swallow-worts, 640
tamarisk, 584
toadflax, 426
tree of heaven, 588
tropical soda apple, 534
waterhyacinth, 342
West Indian marsh grass, 492
winter creeper, 643
wisteria, 647
yellow Himalayan raspberry, 538
Glysine sinensis. See Chinese Wisteria
Goats
and brooms, 501
and fire tree, 557
and gorse, 511
plants toxic to, 360
Gold clam. See Asian clam
Golden Apple Snail, xxiiit (v. 1), 67–70
ISSG 100 worst invaders, 69, 711
state-by-state occurrences, 296, 297, 298, 308
Golden shad. See Alewife
Golden starthistle. See Yellow Starthistle
Golden star tunicate, 40
Golpar, 374
Gooseberries, White Pine Blister Rust alternate
host, 30, 32, 34
Goosefoot family, 379, 409
Gopher plant. See caper spurge
Gopher tortoise, 228, 663
and Australian pine, 543
and cogon grass, 446
Gorse, 508–12
and brooms, 498
impacts, 680, 683
ISSG 100 worst invaders, 710
noxious designation, 666, 667, 670, 671, 692
pathways of introduction, 673, 675
Goutweed, 376–79
impacts, 682
noxious designation, 666, 668, 671, 690
pathways of introduction, 673
Gracula religiosa, 233
Graminoids, 432–92
American species invasive abroad, 696
herbicides, xx (v. 2)
ISSG 100 worst invaders, 710
noxious designation, 691–92
Grape family, 633
Grape honeysuckle, 505, 657
Grape industry threat, Glassy-Winged
Sharpshooter, 137
Grass Carp, 164, 172–75
Hydrilla control, 334
state-by-state occurrences, 296, 300, 302,
307, 308
Grass Carp, biological control (plants)
Eurasian watermilfoil, 325
hydrilla, 334
waterhyacinth, 342
 INDEX n I-17

Grass family, 435, 439, 443, 447, 452, 458, Green peach aphid, 663
462, 466, 469, 473, 478, 481, and tropical soda apple as host, 533–34
485, 489 Green sea turtle, 286, 543, 663
Grasshoppers, biological control (plants) Grey worms. See European Earthworms
cheatgrass, 443 Grinnell, Joseph, xxxi (v. 1)
Chinese lespedeza, 353 Ground elder. See Goutweed
Gratiana boliviana, biological control (plants) Ground ivy, 367, 651
tropical soda apple, 534 Gully-bean. See turkey berry
Gray, Asa GWSS. See Glassy-winged Sharpshooter
water chestnut cultivation, 336 Gypsy Moth, xix (v. 1), xxivt (v. 1), xxvt (v. 1),
Gray herring. See Alewife 138–42
Gray thistle. See wavyleaf thistle ISSG 100 worst invaders, 711
Grazing, physical control (plants) state-by-state occurrences, 295–97, 299,
brooms, 501 300, 301, 303, 304, 305, 307–10
giant hogweed, 375 Gymnancyla canela, biological control (plants)
kudzu, 625 prickly Russian thistle, 413
medusahead, 484 Gymnaetron tetrum, biological control (plants)
spotted knapweed, 420 common mullein, 357
tree of heaven, 588 Gynerium argentium. See Pampas Grass
yellow starthistle, 430 Gynerium jubatum. See Jubata Grass
Greater celandine. See celandine
Great Lakes, xxiv (v. 1), xxvii (v. 1), xxix (v. 1), Hack-and-squirt, herbicide application,
xxx (v. 1) xx (v. 2)
alewives, 157, 158, 159 Haghighat, Sahar, exotic bush honeysuckle
Eurasian watermilfoil, 325 study, 507
gizzard shad, 169 Hairy lespedeza. See Chinese Lespedeza
impacts of sea lamprey, 193 Hairy wheatgrass, 487, 654
musk thistle, 402 Hairy whitetop, 406, 651
New Zealand mud snail, 73, 74 noxious designation, 665, 666, 667, 670,
quagga mussel, 76, 77, 78 671, 672, 690
round goby, 187, 189, 190, 193 Haleakala National Park, and Argentine ants,
sea lamprey, 191 110, 112
spiny water flea, 96, 97 Halogeton, 379–83
threat from bighead carp, 165, 166 impacts, 678, 680, 682
tubenose goby, 188 noxious designation, 665, 666, 667, 669,
water chestnut, 335, 336, 337 670, 690
zebra mussel, 82, 83, 86 pathways of introduction, 677
Great shipworm. See Naval Shipworm and Russian thistle, 412
Great Smoky Mountains National Park, and Halogeton glomeratus. See Halogeton
hemlock woolly adelgid, 143, 145 Haloragaceae. See watermilfoil family
and feral pigs, 279 Halloween lady beetle. See Multicolored Asian
Green anaconda, 220 Lady Beetle
Green comet milkweed, and white Hall’s honeysuckle, 614, 660
swallow-wort, 637, 651 Halyomorpha halys. See Brown Marmorated
Green Crab, xxxi (v. 1), 90–92 Stink Bug
state-by-state occurrences, 296, 297, 300, Harmonia axyridis. See Multicolored Asian
301, 304, 305, 306, 307, 309 Lady Beetle
Greenheart. See coffee colubrina Harold L. Lyon Arboretum, velvet tree, 590
Greenhouse frog, 208–9 Harris mud crab, 87
Green Iguana, 221–25 Hart’s tongue fern, 639, 651
state-by-state occurrences, 297, 298, Hawaiian blackberry, 536, 657
308, 310 Hawai’i Volcanoes National Park
Green-lipped mussel. See Asian Green Mussel and feral goats, 270
Green mussel. See Asian Green Mussel and fire tree, 555, 556
I-18 n INDEX
and kahili ginger, 392, 393, 394
and yellow Himalayan raspberry, 538
Hawkmoth, biological control (plants)
leafy spurge, 398
Heartleaf horsenettle, 532, 651
Heart-podded hoary cress. See Hoary Cress
Hedera helix. See English Ivy
Hedge false bindweed, 607, 660
Hedge garlic. See Garlic Mustard
Hedychium gardnerianum. See Kahili Ginger
Hemlock Woolly Adelgid, xix (v. 1), xxvit (v. 1),
142–45
state-by-state occurrences, 297, 298, 300,
301, 307–9
Hemp family, 618
Hemp sesbania, 528
noxious designation, 665, 690, 692
Herbicide control. See species entries
Herbicides
application methods, xx (v. 2)
major categories, xx (v. 2)
Heracleum mantegazzianum. See Giant Hogweed
Heracleum species. See Giant Hogweed
Herpestes javanicus. See Indian Mongoose
Herpetogramma licarsicalis, biological control
(plants)
kikuyugrass, 481
Heteranthera formosa. See Waterhyacinth
Heterodera sinensis, biological control (plants)
cogongrass, 447
Heteroperreyia hubrichi, biological control
(plants)
Brazilian peppertree, 548
Hexazinone, chemical control (plants)
Brazilian peppertree, 547
buffelgrass, 439
crimson fountain grass, 462
giant salvinia, 329
Highway ice plant. See Ice Plant
Hildebrand, Dr. William, 233
Hill Myna, 233
Himalayan blackberry, 537, 657
noxious designation, 670, 692
pathways of introduction, 676
Himalayan bush clover. See Chinese Lespedeza
Histoplasma capsulatum, Rock Pigeon
as host, 258
HIV, xix (v. 1)
Hive death, honeybee tracheal mite as
cause, 99, 100
Hoary Cress, 404–9
pathways of introduction, 677
“Hogzilla,” Georgia wild pig, 277
Hoh River, Japanese knotweed, 389
Hojo-e, Buddhist captive animal release
ceremony, 165
Holcus halapensis. See Johnsongrass
Hollyhock bindweed. See mallow bindweed
“Hollywood Finches,” 241
Homalodisca vitripennis. See Glassy-Winged
Sharpshooter
Homogenization, of biota, xxviii–xix (v. 1)
Honey bee, 663
and gorse, 511. See also Asian honey bee
Honeybee Tracheal Mite, xix (v. 1), 99–102
state-by-state occurrences, 295–310
Honeycreepers, Hawaii, xxvii (v. 1), 1–3, 217,
247, 261
Honeylocust, 574, 659
Honeysuckle family, 502, 614
Honeysuckle. See Exotic Bush Honeysuckles;
Japanese Honeysuckle
Honeyvine, 637, 660
Hordeum caput-medusae. See Medusahead
Horn nut. See devil pod
Horned water chestnut. See Water Chestnut
Horses, plants toxic to
buffelgrass, 438
common St. Johnswort, 360
field bindweed, 609
Johnsongrass, 472
rattlebox, 529
yellow starthistle, 430
Horsetail tree. See Australian Pine
Hottentot fig. See Ice Plant
House cat. See Feral Cat
House Finch, xiii (v. 1), 240–43
state-by-state occurrences, 295–310
House Mouse, 281–83
ISSG 100 worst invaders, 712
state-by-state occurrences, 295–10
House Myna. See Common Myna
House rat. See Black Rat
House Sparrow, xviii (v. 1), xxiiit (v. 1),
xxxi (v. 1), 1, 238, 241, 243–46
state-by-state occurrences, 295–310
Human factor in species invasions, xxxii (v. 1)
Humboldt Bay, smooth cordgrass
eradication, 458
Humboldt Bay owl’s clover, 457, 651
Humulus japonicus. See Japanese Hops
Humulus scandens. See Japanese Hops
Hyadaphis tatariacae, biological control
(plants)
exotic bush honeysuckles, 507
Hybridization, impacts, xxviii (v. 1)
Hydrellia species, biological control (plants)
hydrilla, 334
 INDEX n I-19

Hydrilla, xviii (v. 1), xxx (v. 1), 173, pampas grass, 477
331–35, 336 perennial pepperweed, 409
impacts, 681, 682, 685 silk tree, 575
noxious designation, 665, 66, 667, 668, 669, tamarisk, 584
670, 671, 689 tropical soda apple, 534
pathways of introduction, 674, 677 west Indian marshgrass, 492
Hydrilla verticillata. See Hydrilla yellow Himalayan raspberry, 539
Hydrocharitaceae. See frog’s-bit family Imperata arundinaceae. See Cogongrass
Hyles euphorbiae. See hawkmoth Imperata cylindrica. See Cogongrass
Hylobius transversovittatus, biological control Imperata cylindrica var. major. See Cogongrass
(plants) Indian lilac. See Chinaberry
purple loosestrife, 417 Indian Mongoose, xviii (v. 1), xxiiit (v. 1),
Hymenachne acutigluma. See West Indian xxvi (v. 1), 284–86
Marsh Grass ISSG 100 worst invaders, 285, 712
Hymenachne amplexicaulis. See West Indian state-by-state occurrences, 298, 310
Marsh Grass Indian Myna. See Common Myna
Hypena srigata, biological control (plants) Indian snakewood. See Asiatic Colubrina
lantana, 521 Indian star vine. See Hydrilla
Hypericum perforatum. See Common Indigo snake, 446, 663
St. Johnswort Infectious diseases, xxiiit (v. 1)
Hyperparathyroidism disease, in horses, 438 Influenza, xix (v. 1)
Hypophthalmichthys molitrix. See Silver Carp Injurious animal species, definition, xv (v. 1)
Hypophthalmichthys nobilis. See Bighead Carp Insects, 106–56
American species invasive abroad, 696
‘Inia. See Chinaberry Insects, biological control (plants), xviii (v. 2),
Iberian knapweed, 419, 651 xix (v. 2)
noxious designation, 665, 666, 669, Australian pine, 543
670, 690 Brazilian peppertree, 547–48
Iberian starthistle, 428, 651 brooms, 501–2
Ice Plant, 383–87 Canada thistle, 348–49
impacts, 683 climbing ferns, 601
noxious designation, 690 common mullein, 357
pathways of introduction, 675 common reed, 451
Ice plant scale insects, biological control (plants) common St. Johnswort, 361
ice plant, 386 Eurasian watermilfoil, 324–25
Iguana iguana. See Green Iguana field bindweed, 609
Imazapic, chemical control (plants) fire tree, 557
cheatgrass, 443 garlic mustard, 372
dyer’s woad, 365 giant hogweed, 375
Japanese stilt grass, 469 giant reed, 466
Imazapyr, chemical control (plants) giant salvinia, 330
Brazilian peppertree, 547 gorse, 511
chinaberry, 554 halogeton, 383
climbing ferns, 601 hoary cress, 409
cogongrass, 446 hydrilla, 334
common reed, 447 ice plant, 386
cordgrasses, 457 Japanese hops, 621
exotic bush honeysuckles, 506 Japanese knotweed, 390
hoary cress, 409 kikuyugrass, 481
jubata grass, 477 Koster’s curse, 518
kahili ginger, 394 kudzu, 625
lantana, 521 lantana, 521
mile-a-minute, 628 leafy spurge, 398–99
melaleuca, 561 mile-a-minute, 628–29
I-20 n INDEX
melaleuca, 561
multi-flora rose, 526
musk thistle, 403
perennial pepperweed, 409
prickly Russian thistle, 413
purple loosestrife, 417
rattlebox, 529
spotted knapweed, 421
strawberry guava, 578–79
swallow-wort, 640
tamarisk, 585
toadflax, 426
tropical soda apple, 534
waterhyacinth, 342–43
west Indian marsh grass, 492
yellow starthistle, 43–31. See also flies,
beetles, butterflies, midges, mites, wasps,
and weevils
Intentional pathways of introduction,
xxiiit–xxivt (v. 1)
Intermediate wheatgrass, 484, 654
International Biosphere Reserve,
Everglades, 560
International Maritime Organization, 708
International Plant Protection Convention
(IPPC), 700, 707
Introduced species, xiv (v. 1)
Invasion process, xx–xxii (v. 1)
Invasion science, xxx–xxxi (v. 1)
Invasive plants
American species abroad, 695–96
impacts of, xviii (v. 2), 678–86
introduction of, xvii–xviii (v. 2), 673–77
management, xix (v. 2)
noxious designation, 665–71, 689–93
organizations and publications concerning,
687–88
problem of, xviii (v. 2)
reproduction/dispersal, xvii (v. 2)
Invasive species
animal problem extent, xviii–xx (v. 1)
definitions, xiii–xv (v. 1)
ecological impacts, xxvi–xxvii (v. 1)
economic impacts, xxix–xxxi (v. 1)
federal legislation and agreements pertaining
to, 699–706
human factor, xxxii (v. 1)
international agreements and conventions
pertaining to, 707–9
natural ecosystem impacts,
xxvii–xxix (v. 1)
pathways of introduction, xxii–xxvi (v. 1)
plant problem extent, xv–xviii (v. 1)
process, xx–xxii (v. 1)
public health and well-being
impacts, xxx (v. 1)
Invasive Species Specialist Group (ISSG)100
worst invasive alien species, 710–12
Invertebrates, 36–156
American species invasive abroad, 696–97
Annelid Worms, 48–53
Bryozoan, 36–38
Cnidarian, 45–48
ISSG 100 worst invaders, 711
Mollusks, 53–86
Tunicates, 39–45
Ioxodes scapularis. See Black-legged tick
Ipomoea species. See pipevine and
Dutchman’s pipe
Irish furze. See Gorse
Irish ivy. See Atlantic ivy
Ironwood. See Australian Pine
Isatis tinctoria. See Dyer’s Woad
Ischnodemus variegatus. See myakka bug
Island applesnail, 67
Italian thistle, 346, 401, 651
noxious designation, 665, 666, 667, 670,
671, 690
Jack-by-the-hedge. See Garlic Mustard
Jacob’s ladder. See Yellow Toadflax
Jacob’s staff. See Common Mullein
Jamaican giant anole, 223
Japanese arrowroot. See Kudzu
Japanese bamboo. See Japanese Knotweed
Japanese barberry, 51, 512–15
impacts, 683
noxious designation, 666, 668, 692
pathways of introduction, 673, 674
Japanese Beetle, xxvit (v. 1), 146–48
state-by-state occurrences, 295–310
Japanese blood grass. See Cogongrass
Japanese brome, 440, 654, 692
Japanese Climbing Fern, 597–602
noxious designation, 665, 667, 693
pathways of introduction, 673
Japanese creeper. See Boston ivy
Japanese Dodder, 610–14
in medicine, 613
impacts, 678, 684, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671
Japanese fleece flower. See Japanese Knotweed
Japanese Honeysuckle, 614–18, 641–42
impacts, 684, 685
noxious designation, 666, 668, 669, 671, 693
pathways of introduction, 673
Japanese hop. See Japanese Hops

Japanese Hops, 618–21
impacts, 681, 684
noxious designation, 666, 668, 693
pathways of introduction, 673, 675
Japanese Knotweed, 387–91
impacts, 681, 682, 685
ISSG 100 worst invaders, 710
noxious designation, 665, 666, 668, 669,
670, 671, 690
pathways of introduction, 673, 675
uses of, 390
Japanese lady beetle. See Multicolored Asian
lady beetle
Japanese millet, biological control (plants)
purple loosestrife, 417
Japanese mitten crab, 87
Japanese mystery snail. See Chinese
Mystery Snail
Japanese oyster, 40
Japanese sedge. See Asiatic Sand Sedge
Japanese Silver-eye, 556
Japanese Stilt Grass, 466–69
impacts, 683
noxious designation, 665, 666,
668, 691
pathways of introduction, 676
Japanese White-Eye, xviii (v. 1), 246–48
and kahili ginger, 393
state-by-state occurrence, 298
and strawberry guava, 578
and velvet tree, 591
Japanese wisteria. See Wisteria
Jenkin’s spire shell. See New Zealand
Mud Snail
Jesup’s milkvetch, 651
and swallow-worts, 639
Johnson, Velma B. (“Wild Horse
Annie”), 273
Johnson grass. See Johnsongrass
Johnsongrass, 469–72
impacts, 678, 679, 680, 681, 683, 685
noxious designation, 667, 668, 670
pathways of introduction, 675
Jointed grass, 467, 654
Jubata Grass, 472–77
impacts, 679, 683, 685
noxious designation, 667, 691
pathways of introduction, 673
Kahila garland lily. See Kahili Ginger
Kahili. See Kahili Ginger
Kahili Ginger, 391–94
impacts, 681, 682
ISSG 100 worst invaders, 710
INDEX n I-21
noxious designation, 690
pathways of introduction, 673
Kariba weed. See Giant Salvinia
Karritree. See Princess Tree
Kelp destruction, Lacy Crust Bryozoan, 37
Kerr, Dr. Warwick, 106
Key West quail-dove, 285
Kiger mustang, 272
Kikuyugrass, 462, 478–81
impacts, 678, 679, 680
noxious designation, 665, 666, 668, 669,
670, 671, 690
pathways of introduction, 675
Killer bee. See Africanized Honey Bee
King, Helen Dean, 289
Klamath weed. See Common St. Johnswort
Knight anole, 223
Knobbed whelk, 80
Koa tree, 659
and fire tree, 556
and strawberry guava, 577
and velvet tree, 590
Kochia. See burning bush
Koi kandy. See Giant Salvinia
Koster’s Curse, 209, 515–18, 589
impacts, 683
ISSG 100 worst invaders, 710
noxious designation, 667, 692
pathways of introduction, 673, 674, 677
and velvet tree, 589
Kraunhia floribunda. See Japanese Wisteria
Kraunhia floribunda var. alba. See Japanese
Wisteria
Kraunhia floribunda var. sinensis. See Chinese
Wisteria
Kraunhia japonica. See Japanese Wisteria
Kraunhia sinensis. See Chinese Wisteria
Kudzu, 622–26
impacts, 684
ISSG 100 worst invaders, 710
and Japanese dodder, 611, 612
noxious designation, 666, 667, 668, 670,
671, 693
pathways of introduction, 674, 675
uses of, 624, 625
Kyack. See Alewife
Kyasuma grass, 459, 461, 654
noxious designation, 665, 666, 668, 669,
670, 671, 691
Lace bug, biological control (plants)
Canada thistle, 349
lantana, 521
Lacey Act, xvi (v. 1), 196, 200, 319, 699, 704
I-22 n INDEX
Lacy Crust Bryozoan, xv (v. 1), xxiv (v. 1),
36–38
state-by-state occurrences, 297, 300, 301,
303, 306
Lagurus cylindricus. See Cogongrass
Lake lamprey. See Sea Lamprey
Lamprey eel. See Sea Lamprey
Land invertebrate, ISSG 100 worst invaders, 711
Lantana, 234, 248, 518–22
impacts, 678, 679, 680, 681, 683, 685
ISSG 100 worst invaders, 710
noxious designation, 692
pathways of introduction, 674
uses of, 522
Lantana aculeate. See Lantana
Lantana camara. See Lantana
Lantana camara var. aculeata. See Lantana
Lantana camara var. nivea. See Lantana
Lantana wildtype. See Lantana
Lardizabala family, 594
Lardizabalaceae. See lardizabala family
Large St. Johnswort, 358, 651
Large-headed sedge, 432, 654
Largeleaf lantana. See Lantana
Larinus curtus, biological control (plants)
yellow starthistle, 431
Larinus minutus, biological control (plants)
spotted knapweed, 421
Larinus planus, biological control (plants)
Canada thistle control, 348
Latherleaf. See Asiatic Colubrina
Laycock, George, xxxi (v. 1)
Leaf-blotching bug, biological control (plants)
melaleuca, 561
Leaf rust, biological control (plants)
pepperweed, 409
Leaf worms. See European Earthworms
Leafy Spurge, 395–99
impacts, 678, 680, 681, 682
ISSG 100 invasive worst, 710
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 677
Least Bell’s Vireo, 662
and giant reeds, 464
and Japanese dodder, 612
Least Tern, 255, 662
and Asiatic sand sedge, 434
Legless red mealybug, biological control (plants)
common reed, 451
Legume family, 527. See also pea family
Leiopython albertisii, 220
Lens-pod hoary cress, 405, 651
noxious designation, 690
Lepidium draba. See Hoary Cress
Lepidium latifolium. See Perennial Pepperweed
Leptinotarsa texana, biological control (plants)
tropical soda apple, 534
Leptospirosis, Norway Rat as host, 290
Lespedeza cuneata. See Chinese Lespedeza
Lespedeza juncea var. sericea. See Chinese
Lespedeza
Lespedeza sericea. See Chinese Lespedeza
Lespedeza webworm, Chinese lespedeza
control, 353
Lesser celadine. See Fig Buttercup
Leucantha solstitialis. See Yellow Starthistle
Leucoptera spartifolilella, biological control
(plants)
brooms, 501
Linaria dalmatica ssp. dalmatica. See Dalmatian
Toadflax
Linaria genistifolia ssp. dalmatica. See Dalmatian
Toadflax
Linaria vulgaris. See Yellow Toadflax
Linepithema humile. See Argentine Ant
Linnet. See House Finch
Lionfish, xxiiit (v. 1), 175–78
state-by-state occurrences, 297,
298, 305, 307
Liothrips urichi, biological control (plants)
Koster’s curse, 518
Lipara species, biological control (plants)
common reed, 451
Lithobates catesbeianus. See American Bullfrog
Lithracus atronotatus, biological control (plants)
Brazilian peppertree, 548
Little, Clarence Cook, 282
Littleleaf sensitive briar, 574, 660
Littoraria irrorata. See marsh periwinkle
Littorina littorea. See Common Periwinkle
Littorina saxatilis. See Rough Periwinkle
Lius peisodon, biological control (plants)
Koster’s curse, 518
Lumbricus rubellus. See European Earthworms
Lumbricus terretris. See European Earthworms
Lonicera insularis. See Exotic Bush Honeysuckles
Lonicera japonica. See Japanese Honeysuckle
Lonicera maackii. See Amur Honeysuckle
Lonicera morrowiii. See Morrow’s Honeysuckle
Lonicera sibirica. See Exotic Bush Honeysuckles
Lonicera tatarica. See Tatarian honeysuckle
Lonicera x bella. See Bell’s Honeysuckle
Lonicera japonica. See Japanese Honeysuckle
Loosestrife family, 414
Lophyrotoma zonalis, biological control (plants)
melaleuca, 561
Louis’ swallow-wort. See Black Swallow-Wort

Love-apple. See cockroach berry
Lygodiaceae. See climbing fern family
Lygodium japonicum. See Japanese
Climbing Fern
Lygodium microphyllum. See Old World
Climbing Fern
Lygodium scandens. See Old World
Climbing Fern
Lymantria dispar. See Gypsy Moth
Lymantria xylina, biological control (plants)
Australian pine, 543
Lyme Disease Bacterium, xix (v. 1),
xxx (v. 1), 3–7
state-by-state occurrences, 296, 298, 299,
300. 301, 304, 306–10
Lythraceae. See loosestrife family
Lythrum salicaria. See Purple Loosestrife
MacArthur, Robert, xxxi (v. 1)
Maccartney rose, 523, 657
Machineel, 495, 659
Madwoman’s milk, 396, 651
Magainins, African Clawed Frog, 204
Magur. See Walking Catfish
Mahogany family, 551
Mahogany flat. See Common Bed Bug
Mallow bindweed, 607, 661
Malta starthistle, 427, 651
noxious designation, 669, 690
Mammals, 259–94
American species invasive abroad, 697
ISSG 100 worst invaders, 712
Mangosteen family, 358
Mangroves, 548, 550
Marine vomit. See Colonial Tunicate
Marlahan mustard. See Dyer’s Woad
Marsh marigold, 367, 651
Marsh periwinkle, biological control (plants)
cordgrass, 458
Mecinnus janthirus, biological control (plants)
toadflax, 426
Medusae, jellyfish generation, 46, 47
Medusahead, 481–84
and cheatgrass, 442,
impacts, 679, 680, 683, 691
noxious designation, 666, 670, 671
pathways of introduction, 677
Medusahead wildrye. See Medusahead
Megamelus species, biological control
(plants)
waterhyacinth, 342
Megastigmus aculeatus var. nigroflavus,
biological control (plants)
multiflora rose, 526
INDEX n I-23
Megastigmus transvaalensis, biological control
(plants)
Brazilian peppertree, 548
Me-jiro. See Japanese White-Eye
Melaleuca, 557–61
impacts, 680, 681, 684
ISSG 100 worst invaders, 710
noxious designation, 665, 666, 668, 669,
670, 671, 692
pathways of introduction, 674, 675, 666
Melaleuca quinquenervia. See Melaleuca
Melaleuca species. See Melaleuca
Melastoma elegans. See Koster’s Curse
Melastoma hirta. See Koster’s Curse
Melastoma hirtum. See Koster’s Curse
Melastomataceae. See melastome family
Melastome family, 515, 589
Meliaceae. See mahogany family
Melia species. See Chinaberry
Melia azedarach. See Chinaberry
Melopsitticus undulates. See Budgerigar
Membranipora membranacea. See Lacy Crust
Bryozoan
Mesembryanthemum crystallinum. See Crystalline
Ice Plant
Mesembryanthemum edule. See Ice Plant
Metriona elatior, biological control (plants)
tropical soda apple, 534
Metrosideros quinquenervia. See Melaleuca
Metsulfuron, chemical control (plants)
climbing ferns, 601
common St. Johnswort, 361
dyer’s woad, 365
halogeton, 382
hoary cress, 409
Japanese hops, 621
yellow Himalayan raspberry, 539
Metzneria paucipunctella, biological control (plants)
spotted knapweed, 421
Mexican bamboo. See Japanese knotweed
Mexican garter snake, 207
Mexican water-fern, 327, 649
Miconia. See Velvet Tree
Miconia calvescens. See Velvet Tree
Miconia magnifica. See Velvet Tree
Microorganisms, 1–10
ISSG 100 worst invaders, 710
Microstegium imberbe. See Japanese Stilt Grass
Microstegium vimineum. See Japanese Stilt Grass
Midges, biological control (plants)
cogongrass, 447
common reed, 451
Eurasian watermilfoil, 324, 325
leafy spurge, 399
I-24 n INDEX

Migratory Bird Treaty Act (2001), 255 Mosquitos

Mikania micrantha, 626, 710 Asian Tiger, 116–20
Mile-A-Minute, 626–29 as avian malaria vectors, 1, 3
impacts, 678, 684, 685 as tularemia vector, 290
noxious designation, 665, 666, 669, 670, as West Nile virus vector, 8, 9, 119
693. See also Mikania micrantha Yellow fever mosquito, 117
Mile-a-minute weed. See Mile-A-Minute Moths, biological control (plants)
Milfoil midge, biological control (plants) Australian pine, 543
Eurasian watermilfoil, 324 bindweed, 609
Military grass. See Cheatgrass Brazilian peppertree, 547–48
Milk thistle, 346, 651 brooms, 501
Milkweed family, 636 chocolate vine, 596
Millettia japonica. See Japanese Wisteria climbing ferns, 601
Mimosa. See Silk Tree common mullein, 357
Mimosa arborea. See Silk Tree common reed, 451
Mimosa julibrissin. See Silk Tree common St. Johnswort, 361
Mimosa wilt, biological control (plants) Eurasian watermilfoil, 324–25
silk tree, 575 field bindweed, 609
Missiongrass, 459–60, 655 fire tree, 557
Mites, biological control (plants) giant reed, 466
climbing ferns, 601 gorse, 511
field bindweed, 609 halogeton, 383
gorse, 511 hydrilla, 334
multiflora rose, 526 Japanese hops, 621
prickly Russian thistle, 413 Koster’s curse, 518
waterhyacinth, 342 leafy spurge, 398
Mollusks, 53–86 mile-a-minute, 629
American species invasive abroad, 696 prickly Russian thistle, 413
Mompha trithalama, biological control Russian knapweed, 421
(plants) sawtooth blackberry, 539
Koster’s curse, 518 spotted knapweed, 421
Monarch butterflies, 663 toadflax, 426
and Asiatic sand sedge, 434 velvet tree, 592
and swallow-worts, 639 waterhyacinth, 342
Money monitor. See Nile Monitor Moth mullein, 354, 651
Monk Parakeet, xxvt (v. 1), xxix (v. 1), Mowing. See cutting
248–52 Mud shad. See Gizzard Shad
state-by-state occurrences, 295, 297–300, Mulberry family, 562
304–8, 310 Mulching, tarping, or solarization, physical
Monogynella japonica. See Japanese Dodder control (plants)
Monopterus albus. See Asian Swamp Eel chocolate vine, 596
Montana Dyer’s Woad Cooperative Project, 365 common reed, 451
Mooney, H. A., xxxi (v. 1) cordgrasses, 457
Moraceae. See mulberry family English ivy, 605
Morella faya. See Fire Tree field bindweed, 609
Morning glory family, 606 goutweed, 379
Morrow’s honeysuckle, noxious designation, Japanese knotweed, 390
666, 668, 669, 671, 692 Johnsongrass, 472
and Japanese honeysuckle, 614. See also kikuyugrass, 481
Exotic Bush Honeysuckles West Indian marsh grass, 491
Mosquito fern, 327, 329, 650 winter creeper, 643
Mosquitofish, xxiiit (v. 1), 178–82 Mulhaden. See Alewife
ISSG 100 worst invaders, 181, 711 Mullein moth, biological control (plants)
state-by-state occurrences, 295–306, 307–10 mullein, 347

Multicolored Asian Lady Beetle, xxiiit (v. 1),
xxx (v. 1), 148–52
state-by-state occurrences, 307
Multiflora Rose, 522–26
impacts, 678, 680, 683, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 692
pathways of introduction, 674, 675, 676
Musk Thistle, 345, 348, 399–404
impacts, 680, 683
noxious designation, 666, 667, 668, 669,
670, 671, 672, 690
pathways of introduction, 677
Mus musculus. See House Mouse
Mustang. See Feral Horse
Mustard family, 362, 369, 404
Mute Swan, 252–56
state-by-state occurrences, 297, 300, 301,
304, 305, 309
Myakka bug, biological control (plants)
West Indian marsh grass, 492
Myiopsitta monachus. See Monk Parakeet
Myocastor coypus. See Nutria
Mycoplasmal conjunctivitis
House Finch decline, 241–42
spread, 243
Mylopharyngodon piceus. See Black Carp
Myrica faya. See Fire Tree
Myricaceae. See sweet gale family
Myriophyllum spicatum. See Eurasian
Watermilfoil
Myriothecium verrucaris, biological control
(plants)
kudzu, 625
Myrtaceae. See eucalyptus family; myrtle
family
Myrtle family, 557, 576
Mytella charruana. See Charru mussel
Myxobolus cerebralis. See Whirling Disease
Nanday Conure (parakeet), 250
Nandayus nenday. See Nanday Conure
Nanny shad. See Gizzard Shad
Nanophyte species, biological control (plants)
water chestnut, 338
Nasturtium draba. See Hoary Cress
National Gypsy Moth Slow the Spread (STS)
program, 142
National Invasive Species Act, 703
Native species, definitions, xiii (v. 1)
Natural selection, and brown anoles, 217
Naval Shipworm, 70–72
Neochetina species, biological control (plants)
waterhyacinth, 342
INDEX n I-25
Neodiplogrammus quadrivittatus. See sesbania
stem borer
Neogobius melanostomus. See Round Goby
Neomusotima conspurcatalis, biological control
(plants)
climbing ferns, 601
Neomusotima fuscolinealis, biological control
(plants)
climbing ferns, 601
Nepalese browntop. See Japanese Stilt Grass
New Zealand Mud Snail, xxvt (v. 1), 73–76
state-by-state occurrences, 295, 296, 298,
302, 303, 306, 308–10
Nicosulfuron, chemical control (plants)
quackgrass, 488
Nightcrawler, xxiiit (v. 1), xxxv (v. 1). See also
European Earthworms
Nigropalallidal encephalomalacia disease, in
horses, 430
Nigua. See Koster’s Curse
Nile Monitor, xxivt (v. 1), 225–28
state-by-state occurrences, 297
Nintooa japonica. See Japanese Honeysuckle
Niphograpta albiguttalis. See waterhyacinth moth
Nitrogen fixer
Australian pine, 541
brooms, 497, 500
fire tree, 556
gorse, 509–10
kudzu, 623, 627
Russian olive, 571–72
silk tree, 575
Nodding thistle. See Musk Thistle
Nonindigenous Aquatic Nuisance Prevention
and Control Act (1990), xv (v. 1),
85, 702
Nonnative species, xv (v. 1)
Norops sagrei. See Brown Anole
Norops sagrei sagrei, 214
Norops sagrei ordinates, 214
North Cascades National Park, and white pine
blister rust, 34
Northern African python, 220
Northern pearly eye butterfly, and Japanese stilt
grass, 468, 664
Northern Snakehead, xix (v. 1), xxiiit (v. 1),
182–85
state-by-state occurrences, 301, 306, 309
Northern watermilfoil, 321–22, 324, 325, 649
Northern wheatgrass, 487, 655
Norway Rat, xviii (v. 1), xxvit (v. 1), 259, 260,
262, 283, 287–90
state-by-state occurrences, 295–310
Noxious species, definition, xv (v. 1)
I-26 n INDEX
Noxious Weed Control and Eradication Act,
704–5
Nutria, xviii (v. 1), xxiiit (v. 1), xxvi (v. 1),
290–94
ISSG 100 worst invaders, 293, 712
state-by-state occurrences, 295– 298, 300–2,
304–6, 307, 308, 309
Oberea erythrocephala, biological control (plants)
leafy spurge, 398
Ohia tree, 556, 659
and strawberry guava, 577, 579
and velvet tree, 591
Old World Climbing Fern, 597–602
impacts, 684
noxious designation, 665, 667, 693
pathways of introduction, 674, 679
Octagonal-tail earthworm. See European
Earthworms
Oleaster. See Russian Olive
Oleaster family, 568
Oncideres cingulata. See twig girdler
Onopordum acanthium. See Scotch thistle
Oncorhynchus mykiss. See Rainbow Trout
Ophideres fullonica, biological control (plants)
chocolate vine, 596
Ophioglossum japonicum. See Japanese
Climbing Fern
Ophiomyia lantanae, biological control (plants)
lantana, 521
Ophiostoma novo-ulmi. See Dutch Elm Disease
Ophiostoma ulmi. See Dutch Elm Disease
Oppositeleaf Russian thistle, 412, 651
noxious designation, 666, 690
Orangeberry nightshade, 532, 661
Orange sheath tunicate. See Chain tunicate
Orchids, 495, 661
Orconectes rusticus. See Rusty Crayfish
Orconectes virilis. See Virile Crayfish
Organ Pipe Cactus National Monument
buffelgrass, 437, 438
Oriental Bittersweet, 629–33
impacts, 681, 684
noxious designation, 666, 668,
669, 671, 693
pathways of introduction, 674
and winter creeper, 641
Orseolia javanica, biological control (plants)
cogongrass, 447
Orthogalumna terebrantis, biological control
(plants)
waterhyacinth, 342
Osteopilus septentrionalis. See Cuban Treefrog
Ossabaw Island hog, 280
Oxyops vitiosa, biological control (plants)
melaleuca, 561
Oyster thief (green algae), 37
Pacific mosquito fern, 327, 649
Painted butterfly, biological control (plants)
Canada thistle, 348
Pale Swallow-Wort, 636–40
impacts, 678, 680, 684
noxious designation, 666, 668, 669,
671, 693
pathways of introduction, 674
Palila, and gorse, 510, 662
Pampas Grass, 460, 473–78
impacts, 679, 683, 685
noxious designation, 691
pathways of introduction, 674
Panicum amplexicaule. See West Indian Marsh
Grass
Paper Mulberry, 562–65
impacts, 681, 684
noxious designation, 693
pathways of introduction, 674
Paperbark tea tree. See Melaleuca
Paperbark tree. See Melaleuca
Papyrius papyriferus. See Paper Mulberry
Parapoynx diminutalis, biological control (plants)
hydrilla, 334
Paraquat, chemical control (plants)
cheatgrass, 443
medusahead, 484
Parrotfeather, 322, 649
noxious designation, 665, 666,
668, 671, 689
Partridge pea, 574, 651
Pasilla. See Chinaberry
Passer domesticus. See House Sparrow
Passer montanus, 244
Pasto buffel. See Buffelgrass
Pasture rose, 524, 657
Pathways of introduction, xxii–xxvi (v. 1)
intentional, xxiiit–xxivt (v. 1)
unintentional, xxiiit (v. 1)
Paulownia. See Princess Tree
Paulownia elongata. See elongate paulownia
Paulownia fortunei. See white-flowered
paulownia
Paulownia imperialis. See Princess Tree
Paulownia tomentosa. See Princess Tree
Peacock cichlid, 198
Pea family, 349, 496, 508, 527, 572, 622, 644
Peacock fly, biological control (plants)
yellow starthistle, 431
Pearl millet, 462, 655
 INDEX n I-27

Pempelia genistella, biological control (plants) Phragmites. See Common Reed

gorse, 511 Phragmites australis ssp. australis.
Pennisetum ciliare. See Buffelgrass See Common Reed
Pennisetum clandestinum. See Kikuyugrass Phragmites communis. See Common Reed
Pennisetum conchroides. See Buffelgrass Phragmites communis var. berlandieri.
Pennisetum inclusum. See Kikuyugrass See Common Reed
Pennisetum incomptum. See Buffelgrass Phragmites phragmites. See Common Reed
Pennisetum longstylum. See Kikuyugrass Phyllachora species, biological control (plants)
Pennisetum longstylum var. clandestinum. West Indian marsh grass, 492
See Kikuyugrass Phyllocoptes fructiphilus, biological control (plants)
Pennisetum macrostachyon. See Crimson multiflora rose, 526
Fountain Grass Phyllonorycter myricae, biological control
Pennisetum rueppelianum. See Crimson (plants)
Fountain Grass fire tree, 557
Pennisetum ruppelii. See Crimson Fountain Grass Phyllorhiza punctata. See Australian Spotted
Pennisetum setaceum. See Crimson Fountain Jellyfish
Grass Phyllotreta ochripes, biological control (plants)
Peppervine, 633, 634, 660, 661 garlic mustard, 372
noxious designation, 693. See also Phymatototrichum omnivorum, biological
Porcelainberry control (plants)
Pepperweed whitetop. See Hoary Cress common mullein, 357
Perennial peppergrass. See Hoary Cress winter creeper, 643
Perennial Pepperweed 404–9 Physical control (plants). See species entries
impacts, 680, 683 Phytophthora ramorum. See Sudden Oak Death
noxious designation, 665, 666, 667, 668, Piaropus crassipes. See Waterhyacinth
669, 670, 671, 672 Piaropus mesomelas. See Waterhyacinth
pathways of introduction, 677 Pierce’s disease, Glassy-Winged Sharpshooter
Perennial snapdragon. See Yellow Toadflax as vector, 137
Perna viridis. See Asian Green Mussel Pickerelweed, 342, 649
Persian ivy. See colchis ivy Pickerelweed family, 339
Persian lilac. See Chinaberry Picloram, chemical control (plants)
Persicaria arifolium var. perfoliatum. See Australian pine, 543
Mile-A-Minute chinaberry, 554
Persicaria perfoliata. See Mile-A-Minute common St. Johnswort, 361
Peruvian peppertree, 545, 659 gorse, 511
Pet trade release Japanese honeysuckle, 617
boa constrictors, 220 Japanese knotweed, 390
Burmese Python, 219 leafy spurge, 398
Common Myna, 233 multiflora rose, 525
Nile Monitor, 226 musk thistle, 403
Petromyzon marinus. See Sea Lamprey spotted knapweed, 420
Phakopsora apoda, biological control (plants) strawberry guava, 578
kikuyugrass, 481 tropical soda apple, 534
Phalaris setaceum. See Crimson Fountain Grass wisteria, 647
Phomopsis arnoldiae, biological control (plants) Pigface fig. See Ice Plant
Russian olive tree, 572 Piggyback plant, 370, 651
Phomopsis broussonetiae, biological control Pilewort. See Fig Buttercup
(plants) Pili grass, 438, 655
paper mulberry, 564 Pimienta de Brazil. See Brazilian Peppertree
Phothedes dulcis, biological control (plants) Pineapple guava. See Strawberry Guava
giant reed, 466 Pine bark adelgid, 31
Phragmataecia castaneae, biological control Pineland lantana, 519, 657
(plants) Pineland verbena. See pineland lantana
common reed, 451 Pine straw, and climbing ferns, 600, 601
I-28 n INDEX
Pineus strobi. See Pine bark adelgid
Pineywoods rooter. See Feral Pig
Pink-flowered tamarisk. See Tamarisk
Pink pampas grass. See Jubata Grass
Pink sand verbena, 386, 651
Pink snow mold, biological control (plants)
cheatgrass, 443
Pipevine, 594, 661
Piping Plover, 662
and Asiatic sand sedge, 434
and oriental bittersweet, 632
Plains bristlegrass, 437, 655
Planthopper, biological control (plants)
cordgrass, 458
waterhyacinth, 342
Plant Protection Act, 704
Plasmodium relictum capistranoae. See Avian
Malaria
Platycephala planifrons, biological control
(plants)
common reed, 451
Pleuropterus cuspidatus. See Japanese Knotweed
Pleuropterus zuccarinii. See Japanese Knotweed
Plowing, physical control (plants). See tilling
Plumeless thistle, 401, 652
noxious designation, 665, 666, 667,
668, 669, 670, 671, 672, 691.
See also Musk Thistle
Poaceae. See grass family
Poison hemlock, 374, 652
Poison ivy, 547, 661
and English ivy, 604
and kudzu, 623
Polygonaceae. See buckwheat family
Polygonum cuspidatum. See Japanese Knotweed
Polygonum perfoliatum. See Mile-A-Minute
Polygonum zuccarinii. See Japanese Knotweed
Pomacea canaliculata. See Golden Apple Snail
Pomacea insularum. See Island applesnail
Pondweed, 333–34, 649
Pontederiaceae. See pickerelweed family
Pontederia crassipes. See Waterhyacinth
Poor-man’s mustard. See Garlic Mustard
Popillia japonica. See Japanese Beetle
Porcelain ampelopsis. See Porcelainberry
Porcelainberry, 633–36
and English ivy, 603
impacts, 684
noxious designation, 666, 668, 693
pathways of introduction, 674
Porcelain vine. See Porcelainberry
Portuguese broom, noxious designation, 670,
692. See also Brooms
Postemergent herbicides, xx (v. 2)
Potamopyrgus antipodarum. See New Zealand
Mud Snail
Potato family, 530
Potato x disease, field bindweed as host, 609
Powderpuff tree. See Silk Tree
Proterorhinus marmoratus. See Tubenose goby
Preemergent herbicides, xx (v. 2)
Prickly glasswort. See Prickly Russian Thistle
Prickly lantana. See Lantana
Prickly rose, 524, 657
Prickly Russian Thistle, 409–14
and halogeton, 380
impacts, 678, 679, 683, 685
noxious designation, 665, 666, 667, 669, 691
pathways of introduction, 677
Prickly sage. See Lantana
Pride of India. See Chinaberry
Princess flower. See glorybush
Princess Tree, 121, 565–68
impacts, 684
noxious designation, 666, 693
pathways of introduction, 674, 676
uses of, 567
Prokeleisa marginata. See planthopper
Prostrate tickrefoil, 623, 652
Pseudocercospora humuli, biological control (plants)
Japanese hops, 621
Pseudodaleta unipuncta. See grass army worms
Pseudomonas bacteria, biological control (plants)
cheatgrass, 443
kudzu, 625
Pseudophilothrips ichini, biological control (plants)
Brazilian peppertree, 548
Pseudorabies, Wild Pig as carrier, 280
Psidium cattleianum. See Strawberry Guava
Psidium guajava. See common guava
Psidium littorale var. longipes. See Strawberry
Guava
Psylliodes chalcomera, biological control (plants)
musk thistle, 403
Pterois miles. See devil firefish. See also Lionfish
Pterois volitans. See Lionfish
Pt. Reyes bird’s beak, 457, 652
Public health and well-being impacts, xxx (v. 1)
Australian pine, 543
chinaberry, 553
English ivy, 605
giant hogweed, 375
Japanese hops, 620
Johnsongrass, 472
leafy spurge, 397
melaleuca, 560
paper mulberry, 563
Public Land Management Act of 2009, 706

Puccinia juncea var. solstitialis, biological control
(plants)
yellow starthistle, 431
Puccinia lygodii, biological control (plants)
climbing ferns, 601
Puccinia punctiformis, biological control (plants)
Canada thistle, 349
Pueraria montana. See Kudzu
Pueraria species. See Kudzu
Puerto Rican treefrog. See Coqui
Pulvinariella species. See ice plant scale insects
Punk tree. See Melaleuca
Purple African nightshade, 532, 657
Purpleflowering raspberry, 524, 657
Purple guava. See Strawberry Guava
Purple Loosestrife, 414–17
impacts, 681, 683
ISSG 100 invasive worst, 711
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 691
pathways of introduction, 674, 675, 677
Purple lythrum. See Purple Loosestrife
Purple pampas grass. See Jubata Grass
Purple plague. See Velvet Tree
Purple sesbane. See Rattlebox
Purple starthistle, 419, 428, 652
noxious designation, 665, 666, 669, 670,
671, 691
Purplestem angelica, 373, 652
Purple strawberry guava. See Strawberry Guava
Pyricularia grisea, biological control (plants)
buffelgrass, 439
kikuyugrass, 481
Python molurus bivittatus. See Burmese Python
Python sebae, 220
Quackgrass, 485–89
impacts, 678, 680, 683
noxious designation, 665, 666, 667, 670,
671, 672, 691
pathways of introduction, 677
Quagga Mussel, xxvt (v. 1), 76–79, 83
state-by-state occurrences, 295, 296,
298–303, 305, 306
Quaker Conure. See Monk Parakeet
Quaker Parrot. See Monk Parakeet
Quassia family, 585
Queen Anne’s lace, 373, 652
Rabbit flower. See Yellow Toadflax
Rainbow Trout, xviii (v. 1), xxivt (v. 1),
xxviii (v. 1), 166, 185–87
ISSG 100 worst invaders, 187, 711
state-by-state occurrences, 295–310
INDEX n I-29
Rainbow weed. See Purple Loosestrife
Rajania quinata. See Chocolate Vine
Ralstonia solanacearum, biological control (plants)
kahili ginger, 394
tropical soda, 534
Rambler rose. See Multiflora Rose
Ramorum blight. See Sudden Oak Death
Rana platernera. See Cuban Treefrog
Range expansion, xv (v. 1)
Range maps, xiv (v. 1)
Ranunculuaceae. See buttercup family
Ranunculus ficaria. See Fig Buttercup
Rapana venosa. See Veined Rapa Whelk
Rattlebox, 527–30
impacts, 679, 681, 682, 683, 685, 692
noxious designation, 692
Rattus norvegicus. See Norway Rat
Rattus rattus. See Black Rat
Razorback. See Feral Pig
Red baron. See Japanese Blood Grass
Redbelly tilapia, 196
Red-billed Leiothrix, and kahili ginger, 393, 662
and velvet tree, 591
Redcoat. See Common Bed Bug
Red honeysuckle, 505, 657
Red Imported Fire Ant, xxvit (v. 1), xxviii (v. 1),
152–56
ISSG 100 worst invaders, 711
state-by-state occurrences, 295–98, 305,
307–10
Red lionfish. See Lionfish
Red mulberry, 562, 659
Red sage. See Lantana
Red sesbania. See Rattlebox
Red sheath tunicate. See Chain Tunicate
Red starthistle. See purple starthistle
Red swamp crayfish, 89
Red wigglers. See European Earthworms
Red worms. See European Earthworms
Rehsonia floribunda. See Japanese Wisteria
Rehsonia sinensis. See Chinese Wisteria
Reptiles, 214–28
American species invasive abroad, 697
ISSG worst invaders, 712
Residuals, herbicides, xx (v. 2)
Reticulated python, 220
Reynoutria. See Japanese Knotweed
Reynoutria japonica, 387
Rhamnaceae. See buckthorn family
Rhamnus asiatica. See Asiatic Colubrina
Rhamnus splendens. See Asiatic Colubrina
Rhinocyllus conicus, biological control (plants)
Canada thistle, 348
musk thistle, 403
I-30 n INDEX

Rhinoncomimus latipes, biological control (plants) Running euonymus. See running

mile-a-minute, 628 strawberry bush
Rhinusa species, biological control (plants) Running strawberry bush, 641, 657
toadflax, 426 Rush wheatgrass, 487, 655
Rhizedra lutosa, biological control (plants) Russian knapweed, 418–19, 428, 652
common reed, 451 noxious designation, 665, 666, 667, 668,
Rhizobacteria, biological control (plants) 669, 670, 671, 672, 691
cheatgrass, 443 toxic, 430
Rhus terebinthifolia. See Brazilian Peppertree Russian Olive, 568–72, 705
Rhyssomatus marginatus. See sesbania impacts, 680, 682, 684, 686
seed weevil noxious designation, 666, 669, 693
Rice eel. See Asian Swamp Eel pathways of introduction, 674, 675, 676
Rice-paddy eel. See Asian Swamp Eel Russian thistle, 380. See Prickly Russian Thistle
Richard’s pampas grass. See toe toe Russian tumbleweed. See Prickly Russian Thistle
RIFA. See Red Imported Fire Ant Russian wheatgrass, 487, 655
Ringed Turtle-Dove, 235–36 Russian wild boar. See Feral Pig
Rio Grande ragweed, 438, 652 Rusty blackhaw, 641, 657
River herring. See Alewife Rusty Crayfish, xxiiit (v. 1), 93–95
Robust blackberry. See Yellow Himalayan state-by-state occurrences, 297, 299, 300,
Raspberry 301, 302, 304, 306, 309
Rock Dove. See Rock Pigeon Ryegrass, 487, 655
Rock Pigeon, xviii (v. 1), 256–59
state-by-state occurrences, 295–310 Sacramento Mountain thistle, and musk thistle,
Rocky Mountain National Park, chytrid frog 402, 652
fungus, 19 Safflower, 348, 652
quackgrass, 487 Salicaire. See Purple Loosestrife
Roof rat. See Black Rat Salmo trutta. See Brown Trout
Root rot Salsola species. See Prickly Russian Thistle
and Australian pine, 544 Salsola tragus. See Prickly Russian Thistle
and common mullein, 357 Saltcedar leaf beetle, biological control (plants)
and giant reed, 466 tamarisk, 585
and winter creeper, 644 Salt cedar. See Tamarisk
Rosa cathayensis. See Multiflora Rose Saltlover. See Halogeton
Rosaceae. See rose family Saltmarsh clubrush. See cosmopolitan bulrush
Rosa multiflora. See Multiflora Rose Salt marsh hay. See Cordgrasses and Their
Roseau. See Common Reed Hybrids
Roseau cane. See Common Reed Salt meadow cordgrass. See Cordgrasses and
Rose family, 522, 535 Their Hybrids
Rose stemgirdler, biological control (plants) Saltwort. See Prickly Russian Thistle
multiflora rose, 526 Salvinia. See Giant Salvinia
Round-leaved bittersweet. See Oriental Salvinia auriculata. See Giant Salvinia
Bittersweet Salvinia auriculata complex, 326
Rough Periwinkle, 61 Salviniaceae. See floating fern family
Round Goby, xxiv (v. 1), xxvt (v. 1), xxvii (v. 1), Salvinia molesta. See Giant Salvinia
187–90 Sameodes albiguttalis. See waterhyacinth moth
state-by-state occurrences, 299, San Clemente Island goats, 269, 270
301, 302, 305 Sand dune thistle, 402, 652
Royal paulownia. See Princess Tree Sapindaceae. See soapberry family
Rubus ellipticus. See Yellow Himalayan Sarcotheca bahiensis. See Brazilian Peppertree
Raspberry Sarthamnus scoparius. See Brooms
Rubus flavus. See Yellow Himalayan Raspberry Sassafras, 562, 659
Rubus gowreephul. See Yellow Himalayan Satintail. See Cogongrass. See also Brazilian
Raspberry satintail; California satintail
Rugosa rose, 524, 657 Sauce-alone. See Garlic Mustard
 INDEX n I-31

Saussurea species, 402 Sericea bush clover. See Chinese Lespedeza

Sawbelly. See Alewife Sericothrips staphylinus, biological control (plants)
Sawtooth blackberry, 537, 539, 657 gorse, 511
noxious designation, 667, 692 Serrate spurge. See toothed spurge
Scaldweed. See common dodder Sesbania flower beetle, biological control (plants)
Scaly bark oak. See Australian Pine rattlebox, 529
Scarlet wisteria. See Rattlebox Sesbania puniceae. See Rattlebox
Scheiffelin, Eugene, 238 Sesbania seed weevil, biological control (plants)
Schinus. See Brazilian Peppertree rattlebox, 529
Schinus species. See Brazilian Peppertree Sesbania tripetii. See Rattlebox
Schinus terebinthifolius. See Brazilian Peppertree Sesbania stem borer, biological control (plants)
Schistocerca americana, biological control (plants) rattlebox, 529
Chinese lespedeza, 353 Sethoxydim, chemical control (plants)
Schizaphiz graminum, biological control (plants) quackgrass, 488
giant reed, 466 Seven-spotted lady beetle, 149
Schoenobius giganatella, biological control (plants) Sewer rat. See Norway Rat
common reed, 451 Shattercane. See sorghum
Scientific Committee on Problems of Sheep, plants toxic to
Environment (SCOPE), xxxi (v. 1) buffelgrass, 438
Sclerotinia sclerotiorum, biological control (plants) common St. Johnswort, 360
Canada thistle, 349 halogeton, 382
Scolytus multistriatus. See smaller European elm Johnsongrass, 472
bark beetle Shenandoah National Park, and
SCOPE, xxxi (v. 1) gypsy moth, 143
Scotch broom, 509 Ship rat. See Black Rat
noxious designation, 666, 667, 670, 671, Shortspike watermilfoil. See northern
692. See also Brooms watermilfoil
Scotch thistle, 401, 403, 652 Short-tip gall midge, biological control (plants)
noxious designation, 665, 666, 667, 668, leafy spurge, 399
669, 670, 671, 672, 691 Shrubby nightshade, 532, 657
Scrambling nightshade. See Tampico soda apple Shrubby Russian thistle, 412, 657
Scrophulariaceae. See figwort family noxious designation, 665, 666, 667, 668,
Seabeach amaranth, 434, 652 669, 670, 671, 692
Seabeach evening primrose, 434, 652 Shrubs, 493–39
Sea-coast marsh elder, 434, 652 American species invasive abroad, 695
Sea fig, 384, 652. See also Ice Plant ISSG 100 worst invaders, 710–11
Sea Lamprey, xxiv (v. 1), xxvt (v. 1), 158, 183, noxious designation, 692
190–93 Sicilian starthistle, 427, 652
state-by-state occurrences, 299, 301, 302, noxious designation, 665, 666, 691
305, 310 Silk Tree, 572–75
Sea sandwort, 434, 652 impacts, 684
Sea-side arrowgrass, 455, 457, 655 noxious designation, 693,
Seaside knotweed, 434, 652 pathways of introduction, 674, 676
Sedge family, 336, 432 Silky acacia. See Silk Tree
Sedges, 455, 546, 654, 655. See also Asiatic Silky bush clover. See Chinese Lespedeza
Sand Sedge Silver Carp, 164, 194–96
Segmented worms, 48 state-by-state occurrences, 299, 300
Selloa. See Jubata Grass Silverberry, 570, 655
Senna obtusifolia, biological control (plants) Silver buffaloberry, 570, 656
kudzu, 625 Silverleaf nightshade, 532, 657
Septoria hodgesii sp. nov, biological control noxious designation, 665, 666, 667, 669,
(plants) 670, 671, 692
fire tree, 557 Silverleaf whitefly, and tropical soda apple as
Sericea lespedeza. See Chinese Lespedeza host, 533–34
I-32 n INDEX
Silver pampas grass. See Pampas Grass
Silversword, 356, 657
Silverthorn, 570, 658
Simaroubaceae. See quassia family
Simberloff, Daniel, xxxi (v. 1)
Sipha species. See sugarcane aphids
Sisymbrium alliaria. See Garlic Mustard
Sisymbrium officinalis. See Garlic Mustard
Skipjack. See Gizzard Shad
Slender-flowered thistle, 401, 652
Slender lespedeza, 350–51, 652
Slender perennial peppercress. See Perennial
Pepperweed
Slender Russian thistle, 411, 652
noxious designation, 665, 666, 691
Slender seapurslane, 434, 652
Slippery mullein. See moth mullein
Slow paralysis virus (SPV), bee virus, 105–6
Small Asian mongoose. See Indian Mongoose
Small cordgrass, 452, 655. See also Cordgrasses
and Their Hybrids
Small crowfoot. See Fig Buttercup
Smaller European elm bark beetle, 22, 23, 24
Small-flowered morning glory. See Field
Bindweed
Small-flowered tamarisk. See Tamarisk
Small Indian mongoose. See Indian Mongoose
Small-leaf climbing fern. See Old World
Climbing Fern
Smooth brome, 440, 655
noxious designation, 692
Smooth cordgrass. See Cordgrasses and Their
Hybrids
Smooth cordgrass hybrid. See Cordgrasses and
Their Hybrids
Smooth rose, 524, 658
Snow-on-the-mountain. See Goutweed
Snowpeaks raspberry, 537, 658
noxious designation, 667, 692
Soapberry family, 548
Soap bush. See Koster’s Curse
SOD. See Sudden Oak Death
Sodom apple. See Tropical Soda Apple
Sod-web worm, Canada thistle host, 347
Soft brome, 440, 655
noxious designation, 692
Soft chess. See Cheatgrass
Soil Conservation Service. See U.S. Department
of Agriculture Soil Conservation Service
Solanaceae. See potato family
Solanum chloranthum. See Tropical Soda Apple
Solanum khasianum var. chatterjeeanum.
See Tropical Soda Apple
Solanum viarum. See Tropical Soda Apple
Solanum viridiflorum. See Tropical Soda Apple
Solarization, physical control (plants).
See mulching or tarping
Soldierwood, 493, 658
Solenopsis invicta. See Red Imported Fire Ant
Solenopsis richteri. See Black imported fire ant
Sonoran Desert weedwackers, 438
Sophonia rufostachia. See two-spotted leafhopper
Sorghum, 470, 472, 655
Sorghum halepense. See Johnsongrass
Sorghum miliaceum. See Johnsongrass
Southern house mosquito, as disease vectors, 1,
3, 8, 9
South Texas. See Rio Grande ragweed
Soybean loper, 664,
and tropical soda apple as host, 533–34
Spanish broom, noxious designation, 667, 670,
671, 692. See also Brooms
Spanish gold. See Rattlebox
Spanish mustang, 271–72
Spanish reed. See Giant Reed
Spartina species. See Cordgrasses and Their
Hybrids
Spartium junceum. See Brooms
Spartium scoparium. See Brooms
Speargrass. See Cogongrass
Species Survival Commission (SSC), ISSG 100
worst invasive species, 710–12
Sphenophorus entus vestitus, biological control
(plants)
kikuyugrass, 481
Spiked loosestrife. See Purple Loosestrife
Spike watermilfoil. See Eurasian Watermilfoil
Spiny Water Flea, xxiv (v. 1), 95–99
state-by-state occurrences, 301, 302, 305,
306, 309
Spirochetes bacterium, 3
Spissistulus festinus. See three-cornered alfalfa leaf
hopper
Spotted jellyfish. See Australian Spotted Jellyfish
Spotted knapweed, 417–21
impacts, 679, 680, 683
noxious designatioins, 665, 666, 667, 668,
669, 670, 671, 672, 691
pathways of introduction, 677
Spotted Tilapia, 196–98
state-by-state occurrences, 296, 297, 303.
See also European Starling
Spurge family, 395
Spurgia esulae. See short-tip gall midge
Spurious mullein. See moth mullein
Squarrose knapweed, 419, 652
noxious designation, 665, 666, 670, 671, 691
Staff-tree family, 629

Staff-vine family, 640
Staghorn sumac, 586, 658
Star thistles, 402. See also Yellow Starthistle
St. Augustine grass, 479, 655
St. Barnaby’s thistle. See Yellow Starthistle
Steele, Dr. Allen, 4
Steelhead (coastal rainbow trout), 185, 186
St. Johnswort. See Common St. Johnswort
Stinking shumac. See Tree of Heaven
Stone plant family, 383
Strangleweed. See Japanese Dodder
Strawberry Guava, 279, 576–79
impacts, 678, 682, 684, 686
ISSG 100 worst invaders, 711
noxious designation, 693
pathways of introduction, 674, 676
Streptopelia decaocto. See Eurasian Collared-Dove
Streptopelia risoria. See Ringed Turtle-Dove
Strophocaulos arvensis. See Field Bindweed
Sturnus vulgaris. See European Starling
Sudangrass, 470, 655
noxious designation, 667
Sudden Oak Death, xix (v. 1), 25–29
state-by-state occurrences, 296, 306
Sugarcane aphids, biological control (plants)
kikuyugrass, 481
Sulfometuron, chemical control (plants)
Japanese honeysuckle, 617
Japanese hops, 621
Sumac family, 544
Summer cypress. See burning bush
Sun sensitivity in humans and livestock
common St. Johnswort, 360
giant hogweed, 375
Sun spurge. See madwoman’s milk
Sunflower family, 344, 399, 417, 427
Surfactant, herbicides, xx (v. 2)
Surinam cherry, 659
and strawberry guava, 578
Sus scrofa. See Feral Pig
Susumber. See turkey berry
Swallow-worts, 636–40
impacts, 678, 680, 684
noxious designation, 666, 668,
669, 671, 693
pathways of introduction, 674
Swamp beaver. See Nutria
Swamp dodder. See common dodder
Swamp fly-honeysuckle, 505, 658
Swamp rose, 524, 658
Swamp verbena, 414, 652
Sweetberry honeysuckles, 505, 658
Sweet breath of spring. See winter honeysuckle
Sweetbriar rose, 524, 658
INDEX n I-33
Sweet cicely, 370, 653
Sweet gale family, 554
Swine, plants toxic to, 360
Swine brucellosis, Wild Pig as carrier, 280
Synchytrium puerariae, biological control (plants)
kudzu, 625
Systemic herbicides, xx (v. 2)
Taeniatherum asperum. See Medusahead
Taeniatherum caput-medusae. See Medusahead
Taeniatherum crinitum. See Medusahead
Tahiti, 591, 592
Tall wheatgrass, 487, 655
Tall whitetop. See Perennial Pepperweed
Tamaricaceae. See tamarisk family
Tamarisk, 579–85
impacts, 682, 684, 686
ISSG 100 worst invaders, 711
noxious designation, 666, 668, 669, 670,
671, 672, 693
pathways of introduction, 674, 676
Tamarix chinensis. See Five-stamen Tamarisk and
Chinese Tamarisk
Tamarix gallica. See French Tamarisk
Tamarix parviflora. See Small-flowered Tamarisk
Tamarix ramossissima. See Pink-flowered
Tamarisk
Tamarix species. See Tamarisk
Tampico soda apple. See wetlands nightshade
Tanglehead. See pili grass
Taosa species, biological control (plants)
waterhyacinth, 342
Tatarian honeysuckle, noxious designation, 666,
668, 669, 671, 692
and Japanese honeysuckle, 614. See also
Exotic Bush Honeysuckles
Tebuthiuron, chemical control (plants)
buffelgrass, 439
Japanese honeysuckle, 617
Tectococcus ovatus, biological control (plants)
strawberry guava, 578
Teleonemia scrupulosa, biological control (plants)
lantana, 521
Teline monspessulanus. See Brooms
Tender fountain grass. See Crimson Fountain
Grass
Teredo navalis. See Naval Shipworm
Terellia ruficauda, biological control (plants)
Canada thistle, 348
Tetralopha scortealis. See lespedeza webworm
Tetranychus lintearis, biological control (plants)
gorse, 511
Tetranychus urticae, Japanese honeysuckle as
host, 617
I-34 n INDEX
Texas umbrella chinaberry, 552, 659
Texas umbrella tree. See Texas umbrella
chinaberry
Thai eggplant. See turkey berry
Thatch bromegrass. See Cheatgrass
Theodore Roosevelt National Park, and feral
horses, 271, 272, 273
Theory of Island Biogeography, The (MacArthur
and Wilson), xxxi (v. 1)
Thorny olive. See silverthorn
Three-cornered alfalfa leaf hopper, biological
control (plants)
Chinese lespedeza, 353
Three-leaf akebia, 594, 661
Thrypticus species, biological control (plants)
waterhyacinth, 342
Tick quackgrass, 487, 655
Tilapia, ISSG 100 worst invaders, 711
waterhyacinth control, 342. See also Spotted
Tilapia
Tilapia mariae. See Spotted Tilapia
Tilapia zilli. See Redbelly tilapia
Tilling, physical control (plants)
cheatgrass, 442
cogongrass, 446
dyer’s woad, 365
field bindweed, 609
medusahead, 484
musk thistle, 403
quackgrass, 488
Timandra convectaria, biological control (plants)
mile-a-minute, 629
Timandra griseata, biological control (plants)
mile-a-minute, 629
Tingis ampliata, biological control (plants)
Canada thistle, 349
Tinker’s penny, 359, 653
Tipton weed. See Common St. Johnswort
Tithymalus esula. See Leafy Spurge
Toadflax, 421–26
impacts, 678, 679, 680, 683, 686
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 674, 675
Tobacco budworm, 664
Japanese honeysuckle as host, 617
tropical soda apple as host, 533–34
Tobacco hornworm, 664
tropical soda apple as host, 533–34
Tobacco leaf curl, biological control (plants)
Japanese honeysuckle, 617
Tobacco mild green mosaic tobamovirus,
biological control (plants)
tropical soda apple, 534
Toe toe, 475, 655
Tomato hornworm, 664
tropical soda apple as host, 533–34
Tomato pinworm, 664
tropical soda apple as host, 533–34
Tomato spotted wilt, field bindweed as host, 609
Tomato weed. See silverleaf nightshade
Toothed spurge, 396, 653
Toothworts, 370, 371, 650, 653
Tordon, chemical control (plants)
gorse, 511
Torrey’s nightshade. See western horsenettle
Toxicodendron altissima. See Tree of Heaven
Toxic plants
brooms, 500
buffelgrass, 438
chinaberry, 553
common St. Johnswort, 360
English ivy, 605
field bindweed, 609
giant hogweed, 375
halogeton, 382
Johnsongrass, 472
kikuyugrass, 480
Koster’s curse, 517
lantana, 521
leafy spurge, 397–98
Oriental bittersweet, 632
rattlebox, 529
toadflax, 425
tropical soda apple, 531
yellow starthistle, 430
Trabutina mannipara, biological control (plants)
tamarisk, 585
Trapa bicornis. See devil’s pod
Trapa bispinosa. See Water Chestnut
Trapaceae. See water caltrop family
Trapa natans. See Water Chestnut
Trapa natans var. bispinosa. See Water Chestnut
Trapa natans var. natans. See Water Chestnut
Trapdoor snail. See Chinese Mystery Snail
Trap-neuter-release (TNR) programs,
feral cats, 268
Tree of Heaven, 248, 585–89
impacts, 684
noxious designation, 666, 668, 669, 671, 693
pathways of introduction, 674
uses of, 588
Trees, 540–93
American species invasive abroad, 695
ISSG 100 worst invaders, 710–11
noxious designation, 692–93
Trichapion latrivetre. See sesbania
flower beetle
 INDEX n I-35

Trichomonas gallinae parasite, Eurasian Trumpet creeper, 645, 661

Collared-Dove as host, 236 Trumpet grass. See West Indian Marsh Grass
Trichosirocalus horridus, biological control (plants) Tubenose goby, 188
musk thistle, 403 Tuckeroo tree. See Carrotwood
Trichosporium visiculosum, biological control Tucson Mountain Park, 438
(plants) Tumbleweed. See Prickly Russian Thistle
Australian pine, 543 Tunicates, 39–45
Triclopyr, chemical control (plants) Turkey berry, 531, 658
Asiatic colubrina, 495 noxious designation, 665, 666, 667, 668,
Australian pine, 543 669, 670, 671, 692
Brazilian peppertree, 547 Turkeyfish, 175
brooms, 501 Turtles, and Australian pine, 543
carrotwood, 550 Tu Si Zi. See Japanese Dodder
chinaberry, 554 Twig girdler, biological control (plants)
Chinese lespedeza, 352 Australian pine, 543
chocolate vine, 596 Twinberry honeysuckle. See bearberry
common mullein, 36 honeysuckle
English ivy, 605 Twitchgrass. See Quackgrass
Eurasian watermilfoil, 324 2,4-D, chemical control (plants)
exotic bush honeysuckles, 506 common St. Johnswort, 361
giant hogweed, 375 dyer’s woad, 365
gorse, 511 Eurasian watermilfoil, 324
Japanese barberry, 514 field bindweed, 609
Japanese knotweed, 390 gorse, 511
Koster’s curse, 518 Japanese honeysuckle, 617
kudzu, 625 leafy spurge, 398
multiflora rose, 525 multiflora rose, 525
musk thistle, 403 musk thistle, 403
Oriental bittersweet, 632 prickly Russian thistle, 413
paper mulberry, 564 spotted knapweed, 420
porcelainberry, 635 strawberry guava, 578
prickly Russian thistle, 413 tamarisk, 584
princess tree, 568 velvet tree, 592
purple loosestrife, 417 water chestnut, 338
rattlebox, 529 waterhyacinth, 342
strawberry guava, 578 Twoleaf toothwort. See crinkleroot
swallow-worts, 640 Two-spotted leafhopper, biological control
tamarisk, 585 (plants)
tree of heaven, 588 fire tree, 557
tropical soda apple, 534 Tyta luctuosa, biological control (plants)
velvet tree, 592 field bindweed, 609
winter creeper, 643
wisteria, 647 Ugena microphylla. See Old World Climbing Fern
yellow Himalayan raspberry, 538, 539 Ulex europaeus. See Gorse
Triploid grass carp, 174 Umbrellatree. See Chinaberry
Triticum repens. See Quackgrass Unaspis euonumi, biological control (plants)
Triticum vaillantianum. See Quackgrass winter creeper, 643
Trompetilla. See West Indian Marsh Grass Unintentional pathways of introduction,
Tropical curlygrass fern, 600, 653 xxvt (v. 1), 676–77
Tropical Soda Apple, 530–35 Upland frog. See African Clawed Frog
impacts, 678, 680, 681, 684, 685 Urophora species, biological control (plants)
noxious designation, 665, 666, 668, 669, spotted knapweed, 421
670, 671, 692 yellow starthistle, 431
pathways of introduction, 677 Uruguayan pampas grass. See Pampas Grass
I-36 n INDEX

U.S. Department of Agriculture Soil Vitis brevipedunculata. See Porcelainberry

Conservation Service, xvi (v. 2), Viviparus georgianus, 59
xviii (v. 2), 437, 499, 524, 623 VOC. See volatile organic compound
Ustilagao bulleta, biological control (plants) Volatile organic compound, and kudzu, 624
cheatgrass, 443
Wahiawa Botanical Garden, and velvet tree, 590
Vaccinium flase bottom, field bindweed Waiawi. See Strawberry Guava
as host, 609 Walking Catfish, xviii (v. 1), 198–201
Vanessa cardui. See painted butterfly ISSG 100 worst invaders, 200, 711
Varanus niloticus. See Nile Monitor state-by-state occurrences, 297
Varanus salvator. See Asian water monitor Wall louse. See Common Bed Bug
Variable leaf waterhyacinth, 339–40, 649 Wasps, biological control (plants)
Variable watermilfoil, 322, 649 Australian pine, 543
noxious designation, 666, 667, 668, Brazilian peppertree, 548
671, 689 multiflora rose, 526
Varnish tree. See Tree of Heaven velvet tree, 592
Varroa destuctor. See Varroa Mite Water caltrop. See Water Chestnut
Varroa Mite, xix (v. 1), 102–5 Water caltrop family, 335
Africanized Honey Bee as host, 103, 108 Water Chestnut, 335–39
state-by-state occurrences, 295–310 impacts, 682, 685
Veined Rapa Whelk, 79–82 noxious designation, 665, 666, 667, 668,
state-by-state occurrences, 308 669, 670, 671, 689
Velvet dock. See Common Mullein pathways of introduction, 674, 677
Velvet Tree, 515, 589–93 Water clover, 327, 330, 650
impacts, 682, 684 Water fern. See Giant Salvinia
and Koster’s curse, 515 Waterhyacinth, xviii (v. 1), 173, 293, 336,
noxious designation, 667, 693 339–43, 522
pathways of introduction, 674 impacts, 680, 681, 682, 685
Verbascum thapsus. See Common Mullein ISSG 100 worst invasives, 710
Verbenaceae. See verbena family noxious designation, 665, 666, 668, 669,
Verbena family, 518 670, 671, 689
Vertebrates, 157–294 pathways of introduction, 674, 677
American species invasive abroad, 697 Waterhyacinth moth, biological control (plants)
amphibians, 201–14 waterhyacinth, 342
birds, 228–59 Water milfoil family, 322
fish, 157–201 Water spangles. See common salvinia
ISSG 100 worst invaders, 711–12 Water straw grass. See West Indian Marsh Grass
mammals, 259–94 Water thyme. See Hydrilla
reptiles, 214–28 Water velvet. See Giant Salvinia
Verticillium dahliae, biological control (plants) Wauke. See Paper Mulberry
tree of heaven, 589 Wavyleaf thistle, 345, 653
Vincetoxicum medium. See Pale Swallow-Wort Weevils, biological control (plants)
Vincetoxicum nigrum. See Black Swallow-Wort Australian pine, 543
Vincetoxicum rossicum. See Pale Swallow-Wort brooms, 501
Vines, 594–644 Canada thistle, 348, 349
ISSG 100 worst invaders, 710–11 common mullein, 357
noxious designation, 693 Eurasian watermilfoil, 324, 325
Violets, 370, 653 garlic mustard, 372
Violet tunicate, 39 giant salvinia, 330
Virginia creeper, 619, 661 gorse, 511
Virginia cutgrass, 467, 656 kudzu, 625
Virginia rose, 524, 658 mile-a-minute, 628–29
Virile crayfish, 93, 94 musk thistle, 403
Vitaceae. See grape family purple loosestrife, 417

rattlebox, 529
tropical soda apple, 534
spotted knapweed, 421
toadflax, 426
tropical soda apple, 534
velvet tree, 592
water chestnut, 338
waterhyacinth, 342
yellow starthistle, 430–31
West African pennisetum. See Kikuyugrass
Western horsenettle, 533, 653
noxious designation, 666, 691
Western morning glory, 607, 661
Western mosquito fish. See Mosquitofish
Western wheatgrass, 442, 487, 656
West Indian mahogany, 495, 659
West Indian Marsh Grass, 489–92
impacts 682, 683
noxious designation, 692
pathways of introduction, 675, 677
West Indian raspberry, 537, 658
noxious designation, 692
West Nile Virus, xix (v. 1), 7–10
mosquito as vector, 119
state-by-state occurrences, 295–310
West Virginia white butterfly, and garlic
mustard, 371
Wetlands nightshade, 531–32, 661
noxious designation, 665, 666, 667, 668,
669, 670, 671, 693
Wetlands soda apple. See wetlands
nightshade
Wharf rat. See Norway Rat
Whin. See Gorse
Whirling disease parasite, salmonids, 187
Whistling pine. See Australian Pine
White amur. See Grass Carp
White avens, 370, 653
White basswood, 562, 659
White bottlebrush tree. See Melaleuca
White cedar. See Chinaberry
White-flowered paulownia, 565–66, 659
White ginger, 391, 392, 393, 394, 653
White grass. See Virginia cutgrass
White herring. See Alewife
White horsenettle. See silverleaf nightshade
White lace bryozoan. See Lacy Crust Bryozoan
White leaf rust, biological control (plants)
perennial pepperweed, 409
White-lipped python, 220
White mulberry, 562, 659
White Pine Blister Rust, 29–35
state-by-state occurrences, 296–310
White ricefield eel. See Asian Swamp Eel
INDEX n I-37
White-spotted jellyfish. See Australian
Spotted Jellyfish
White swallow-wort, 637, 661
noxious designation, 671, 693
White-tailed deer, 663
and earthworms, 51
White top. See Hoary Cress
White-winged parakeet, 250
Wild blackberry. See Yellow Himalayan
Raspberry
Wild cane. See Giant Reed
Wild cinnamon, 495, 659
Wild coffee. See Asiatic Colubrina
Wild cucumber, 619, 661
Wild donkey. See Feral Burro
Wild Free-Roaming Horse and Burro Act
(1971), 273, 274
Wild ginger. See Kahili Ginger
Wild grape, 603, 661. See also Porcelainberry
Wild horse. See Feral Horse
Wild oats. See Cheatgrass
Wild parsnip, 373, 653
Wild pigs. See Feral Pig
Wild raspberry. See Yellow Himalayan
Raspberry
Wild rye, 483, 656. See also Quackgrass
Wild snapdragon. See Toadflax
Wilelaiki. See Brazilian Peppertree
Willow Flycatcher, 662
and giant reeds, 464
Wilson, E. O., xxxi (v. 1)
Windwitch. See Prickly Russian Thistle
Winter creeper, 640–44
impacts, 684
noxious designation, 693
pathways of introduction, 674
Winter honeysuckle, 504–5, 658
Winter shad. See Gizzard Shad
Wiregrass. See Quackgrass
Wisteria, 644–47
impacts, 681, 684
noxious designation, 693
pathways of introduction, 674
Wistar Institute, Philadelphia, 289
Wisteria sinensis. See Chinese Wisteria
Wisteria floribunda. See Japanese Wisteria
Witchgrass. See Quackgrass
Witchweed. See Prickly Russian Thistle
WNV. See West Nile Virus
Wolf’s milk. See Leafy Spurge
Wolf’s primrose, 386, 653
Woman’s tongue, 572–73, 660
Woodbine. See Japanese Honeysuckle
Woolly. See Hemlock Woolly Adelgid
I-38 n INDEX
Wooly Dutchman’s pipe, 594, 661
and climbing ferns, 600
Wooly mullein. See Common Mullein
World Conservation Union (IUCN), ISSG 100
worst invasive alien species, 710–12
Wormwood, 434, 658
Wrinkle wrinkle. See Common Periwinkle
Xenopus laevis. See African Clawed Frog
Xylella fastidiosa, glassy-winged sharpshooter as
vector, 137. See also bacterial leaf scorch
Yellow anaconda, 220
Yellow-brown stink bug. See Brown Marmorated
Stink Bug
Yellow cane. See Common Reed
Yellow-chevroned parakeet, 250
Yellow cockspur. See also Yellow Starthistle
Yellow eel. See Asian Swamp Eel
Yellow fever mosquito, 117
Yellow ginger, 391–92, 393, 394, 653
Yellow Himalayan Raspberry, 535–39
impacts, 684
ISSG 100 worst invaders, 711
noxious designation, 667, 692
pathways of introduction, 676
Yellow honeysuckle, 505, 658
Yellow raspberry. See Yellow Himalayan
Raspberry
Yellow sage. See Lantana
Yellowspine thistle, 345, 653
Yellow Starthistle, 402, 427–31
impacts, 679, 680, 683, 685, 686
noxious designation, 665, 666, 667, 668,
669, 670, 671, 691
pathways of introduction, 677
Yellow toadflax. See Toadflax
Yersinia pestis, carried by black
rats’ fleas, 262, 290
Zacate buffel. See Buffelgrass
Zapata bladderpod, 438, 653
Zasloff, Dr. Michael, 204
Zauclophora pelodes, biological control (plants)
Australian pine, 543
Zebrafish. See Lionfish
Zebra Mussel, xix (v. 1), xxiv (v. 1), xxvt (v. 1),
xxvii (v. 1), xxviii (v. 1), xxix (v. 1), 58,
76, 77, 78, 79, 80, 82–86, 189, 190
ISSG 100 worst invaders, 711
state-by-state occurrences, 295–10
Zingiberaceae. See ginger family
Zonate leafspot, biological control (plants)
tree of heaven, 589
Zooids
bryozoan young, 36, 37
tunicate young, 39–0, 42
Zosterops japonicas. See Japanese White-Eye
Zyginidia guyumi, biological control (plants)
giant reed, 466
n About the Authors
SUSAN L. WOODWARD received her PhD in Geography—with a specialization in biogeog-
raphy—from the University of California, Los Angeles in 1976. Her doctoral research
included three years along the Lower Colorado River studying feral burros, considered by
some then and now to be an invasive species. When her work began, the burro (along with
the feral horse) had just been placed under the jurisdiction of the U.S. Bureau of Land
Management (BLM), which had a federal mandate to manage this living symbol of the Old
West. The results of her field work provided the BLM with some of its earliest baseline data
on burro population biology and ecology.
Dr. Woodward taught biogeography, physical geography, and human ecology for 22 years
at Radford University in Virginia, before retiring in 2006. She is the author of Biomes of Earth
(2003) and served as general editor and author of three volumes for Greenwood Guides to
Biomes of the World (2009).

JOYCE A. QUINN retired from California State University, Fresno as professor emerita after
21 years of teaching a variety of courses in physical geography and mapping techniques. She
earned an MA from the University of Colorado and a PhD from Arizona State University,
both in Geography, specializing in the effect of climate and soils on the distribution of
plants. She has traveled extensively throughout North America, Latin America, Europe,
northern and southern Africa, Uzbekistan, Nepal, China, Southeast Asia, Micronesia, and
elsewhere. She is a member of the Cactus and Succulent Society of America and the
California Invasive Plant Council and is the author of two volumes of Greenwood Guides to
Biomes of the World (2009).
n Encyclopedia of Invasive
Species
This page intentionally left blank
n Encyclopedia
of Invasive Species
From Africanized Honey Bees
to Zebra Mussels
Volume 2: Plants

Susan L. Woodward and Joyce A. Quinn

Copyright 2011 by ABC-CLIO, LLC
All rights reserved. No part of this publication may be reproduced, stored in a retrieval
system, or transmitted, in any form or by any means, electronic, mechanical,
photocopying, recording, or otherwise, except for the inclusion of brief quotations in a
review, without prior permission in writing from the publisher.
Library of Congress Cataloging-in-Publication Data
Woodward, Susan L., 1944 Jan. 20–
Encyclopedia of invasive species : from africanized honey bees to zebra mussels / Susan L. Woodward and
Joyce A. Quinn.
p. cm.
Includes bibliographical references and index.
ISBN 978–0–313–38220–8 (cloth : alk. paper) — ISBN 978–0–313–38221–5 (ebook)
1. Introduced organisms—Encyclopedias. I. Quinn, Joyce Ann. II. Title.
QH353.W66 2011
578.60 2—dc23 2011026543

ISBN: 978–0–313–38220–8
EISBN: 978–0–313–38221–5
15 14 13 12 11 1 2 3 4 5
This book is also available on the World Wide Web as an eBook.
Visit www.abc-clio.com for details.
Greenwood
An Imprint of ABC-CLIO, LLC
ABC-CLIO, LLC
130 Cremona Drive, P.O. Box 1911
Santa Barbara, California 93116-1911
This book is printed on acid-free paper
Manufactured in the United States of America
n Contents
General Introduction: Invasive Species—Concepts and Issues n xiii

VOLUME 1: INVASIVE MICROORGANISMS, FUNGI,

AND ANIMALS
Preface n xxxv
Alphabetical List of Invasive Microorganisms, Fungi, and Animal
Entries n xxxix

Microorganisms
Avian Malaria (Plasmodium relictum capistranoae) n 1
Lyme Disease Bacterium (Borrelia burgdorferi) n 3
West Nile Virus (West Nile Virus) n 7

Fungi
Bat White-Nose Syndrome Fungus (Geomyces destructans) n 11
Chestnut Blight Fungus (Cryphonectria parasitica) n 14
Chytrid Frog Fungus (Batrachochytrium
dendrobatidis) n 18
Dutch Elm Disease Fungi (Ophiostoma novo-ulmi
and O. ulmi) n 21
Sudden Oak Death (Phytophthora ramorum) n 25
White Pine Blister Rust (Cronartium ribicola) n 29
Invertebrates
Bryozoan
Lacy Crust Bryozoan (Membranipora membranacea) n 36
Tunicates
Chain Tunicate (Botrylloides violaceus) n 39
Colonial Tunicate (Didemnum vexillum) n 42
Cnidarian
Australian Spotted Jellyfish (Phyllorhiza punctata) n 45
vi n CONTENTS

Annelid worms
European Earthworms (Lumbricus terrestris, L. rubellus,
Aporrectodea caliginosa, Dendrobaena octaedra, and
others) n 48
Mollusks
Asian Clam (Corbicula fluminea) n 53
Asian Green Mussel (Perna viridis) n 56
Chinese Mystery Snail (Cipangopaludina
chinensis malleata) n 58
Common Periwinkle (Littorina littorea) n 61
Giant African Snail (Achatina fulica) n 64
Golden Apple Snail (Pomacea canaliculata) n 67
Naval Shipworm (Teredo navalis) n 70
New Zealand Mud Snail (Potamopyrgus
antipodarum) n 73
Quagga Mussel (Dreissena rostriformis bugensis) n 76
Veined Rapa Whelk (Rapana venosa) n 79
Zebra Mussel (Dreissena polymorpha) n 82
Crustaceans
Chinese Mitten Crab (Eriocheir sinensis) n 86
Green Crab (Carcinus maenas) n 90
Rusty Crayfish (Orconectes rusticus) n 93
Spiny Water Flea (Bythotrephes longimanus) n 95
Arachnids
Honeybee Tracheal Mite (Acarapis woodi) n 99
Varroa Mite (Varroa destructor) n 102

Insects
Africanized Honey Bee (Apis mellifera scutellata) n 106
Argentine Ant (Linepithema humile) n 110
Asian Longhorned Beetle (Anoplophora
glabripennis) n 113
Asian Tiger Mosquito (Aedes albopictus) n 116
Brown Marmorated Stink Bug (Halyomorpha halys) n 120
Common Bed Bug (Cimex lectularius) n 123
Emerald Ash Borer (Agrilus planipennis) n 127
 CONTENTS n vii

Formosan Subterranean Termite (Coptotermes

formosanus) n 131
Glassy-Winged Sharpshooter (Homalodisca vitripennis) n 134
Gypsy Moth (Lymantria dispar) n 138
Hemlock Woolly Adelgid (Adelges tsugae) n 142
Japanese Beetle (Popillia japonica) n 146
Multicolored Asian Lady Beetle (Harmonia axyridis) n 148
Red Imported Fire Ant (Solenopsis invicta) n 152

Vertebrates
Fish
Alewife (Alosa pseudoharengus) n 157
Asian Swamp Eel (Monopterus albus) n 160
Bighead Carp (Hypophthalmichthys nobilis) n 163
Brown Trout (Salmo trutta) n 166
Gizzard Shad (Dorosoma cepedianum) n 168
Grass Carp (Ctenopharyngodon idella) n 172
Lionfish (Pterois volitans / P. miles) n 175
Mosquitofish (Gambusia affinis and G. holbrooki) n 178
Northern Snakehead (Channa argus) n 182
Rainbow Trout (Oncorhynchus mykiss) n 185
Round Goby (Neogobius melanostomus) n 187
Sea Lamprey (Petromyzon marinus) n 190
Silver Carp (Hypophthalmichthys molitrix) n 194
Spotted Tilapia (Tilapia mariae) n 196
Walking Catfish (Clarias batrachus) n 198
Amphibians
African Clawed Frog (Xenopus laevis) n 201
American Bullfrog (Lithobates catesbeianus) n 205
Coqui (Eleutherodactylus coqui) n 208
Cuban Treefrog (Osteopilus septentrionalis) n 211
Reptiles
Brown Anole (Norops [=Anolis] sagrei) n 214
Burmese Python (Python molurus bivittatus) n 217
Green Iguana (Iguana iguana) n 221
viii n CONTENTS

Nile Monitor (Varanus niloticus) n 225

Birds
Cattle Egret (Bubulcus ibis) n 228
Common Myna (Acridotheres tristis) n 232
Eurasian Collared-Dove (Streptopelia decaocto) n 234
European Starling (Sturnus vulgaris) n 237
House Finch (Carpodacus mexicanus) n 240
House Sparrow (Passer domesticus) n 243
Japanese White-Eye (Zosterops japonicus) n 246
Monk Parakeet (Myiopsitta monachus) n 248
Mute Swan (Cygnus olor) n 252
Rock Pigeon (Columba livia) n 256
Mammals
Black Rat (Rattus rattus) n 259
Feral Burro (Equus asinus) n 262
Feral Cat (Felis silvestris catus) n 265
Feral Goat (Capra hircus) n 268
Feral Horse (Equus caballus) n 271
Feral Pig (Sus scrofa) n 275
House Mouse (Mus musculus) n 281
Indian Mongoose (Herpestes javanicus) n 284
Norway Rat (Rattus norvegicus) n 287
Nutria (Myocastor coypus) n 290
State-by-State Occurrences of Invasive Microorganisms, Fungi, and
Animals n 295

Glossary n 311
Index n I-1

VOLUME 2: INVASIVE PLANT SPECIES

Preface n xiii

Invasive Plants in the United States: A Brief Overview n xvii

Alphabetical List of Invasive Plant Entries n xxiii
 CONTENTS n ix

Aquatic Plants
Eurasian Watermilfoil (Myriophyllum spicatum) n 321
Giant Salvinia (Salvinia molesta) n 326
Hydrilla (Hydrilla verticillata) n 331
Water Chestnut (Trapa natans) n 335
Waterhyacinth (Eichhornia crassipes) n 339

Forbs
Canada Thistle (Cirsium arvense) n 344
Chinese Lespedeza (Lespedeza cuneata) n 349
Common Mullein (Verbascum thapsus) n 353
Common St. Johnswort (Hypericum perforatum) n 358
Dyer’s Woad (Isatis tinctoria) n 362
Fig Buttercup (Ficaria verna) n 366
Garlic Mustard (Alliaria petiolata) n 369
Giant Hogweed (Heracleum mantegazzianum) n 372
Goutweed (Aegopodium podagraria) n 376
Halogeton (Halogeton glomeratus) n 379
Ice Plant and Crystalline Ice Plant (Carpobrotus edulis
and Mesembryanthemum crystallinum) n 383
Japanese Knotweed (Fallopia japonica) n 387
Kahili Ginger (Hedychium gardnerianum) n 391
Leafy Spurge (Euphorbia esula) n 395
Musk Thistle (Carduus nutans) n 399
Perennial Pepperweed and Hoary Cress (Lepidium
latifolium and Cardaria draba) n 404
Prickly Russian Thistle (Salsola tragus) n 409
Purple Loosestrife (Lythrum salicaria) n 414
Spotted Knapweed (Centaurea stoebe) n 417
Toadflax (Linaria dalmatica ssp. dalmatica and
Linaria vulgaris) n 421
Yellow Starthistle (Centaurea solstitialis) n 427
Graminoids
Asiatic Sand Sedge (Carex kobomugi) n 432
Buffelgrass (Pennisetum ciliare) n 435
x n CONTENTS

Cheatgrass (Bromus tectorum) n 439

Cogongrass (Imperata cylindrica) n 443
Common Reed (Phragmites australis ssp. australis) n 447
Cordgrasses and Their Hybrids (Spartina alterniflora, Spartina
densiflora, Spartina patens, Spartina anglica, and Spartina
alterniflora x foliosa n 452
Crimson Fountain Grass (Pennisetum setaceum) n 458
Giant Reed (Arundo donax) n 462
Japanese Stilt Grass (Microstegium vimineum) n 466
Johnsongrass (Sorghum halepense) n 469
Jubata Grass and Pampas Grass (Cortaderia jubata and Cortaderia
selloana) n 472
Kikuyugrass (Pennisetum clandestinum) n 478
Medusahead (Taeniatherum caput-medusae) n 481
Quackgrass (Elymus repens) n 485
West Indian Marsh Grass (Hymenachne
amplexicaulis) n 489

Shrubs
Asiatic Colubrina (Colubrina asiatica) n 493
Brooms (Cytisus scoparius, Spartium junceum, Genista
monspessulana, and Cytisus striatus) n 496
Exotic Bush Honeysuckles (Lonicera maackii, L. morrowii,
L. tatarica, and L. x bella) n 502
Gorse (Ulex europaeus) n 508
Japanese Barberry (Berberis thunbergii) n 512
Koster’s Curse (Clidemia hirta) n 515
Lantana (Lantana camara) n 518
Multiflora Rose (Rosa multiflora) n 522
Rattlebox (Sesbania punicea) n 527
Tropical Soda Apple (Solanum viarum) n 530
Yellow Himalayan Raspberry (Rubus ellipticus) n 535
Trees
Australian Pine (Casuarina equisetifolia) n 540
Brazilian Peppertree (Schinus terebinthifolius) n 544
 CONTENTS n xi

Carrotwood (Cupaniopsis anacardioides) n 548

Chinaberry (Melia azedarach) n 551
Fire Tree (Morella faya) n 554
Melaleuca (Melaleuca quinquenervia) n 557
Paper Mulberry (Broussonetia papyrifera) n 562
Princess Tree (Paulownia tomentosa) n 565
Russian Olive (Elaeagnus angustifolia) n 568
Silk Tree (Albizia julibrissin) n 572
Strawberry Guava (Psidium cattleianum) n 576
Tamarisk (Tamarix chinensis T. ramosissima,
T. parviflora, and T. gallica) n 579
Tree of Heaven (Ailanthus altissima) n 585
Velvet Tree (Miconia calvescens) n 589

Vines
Chocolate Vine (Akebia quinata) n 594
Climbing Ferns (Lygodium japonicum and Lygodium micro-
phyllum) n 597
English Ivy (Hedera helix) n 602
Field Bindweed (Convolvulus arvensis) n 606
Japanese Dodder (Cuscuta japonica) n 610
Japanese Honeysuckle (Lonicera japonica) n 614
Japanese Hops (Humulus japonicus) n 618
Kudzu (Pueraria montana) n 622
Mile-A-Minute (Persicaria perfoliata) n 626
Oriental Bittersweet (Celastrus orbiculatus) n 629
Porcelainberry (Ampelopsis glandulosa var. brevipedunculata) n
633
Swallow-Worts (Cynanchum rossicum and Cynanchum louiseae) n
636
Winter Creeper (Euonymus fortunei) n 640
Wisteria (Wisteria sinensis and Wisteria floribunda) n 644

Tables and Lists about Invasive Plants

Common Names and Scientific Names n 649
State-by-State Designations of Invasive or Noxious Weeds n 665
xii n CONTENTS

Pathways of Introduction for Plants n 673

Impacts of Invasive Plants n 678
Major Organizations and Publications Concerned about Invasive
Plants n 687
Plant Species Listed as Invasive or Noxious by Organizations and
State and Federal Governments n 689
Set Appendices

Appendix A: American Species That Are Invasive Abroad n 695

Appendix B: Major Federal Legislation and Agreements Pertaining to

Invasive Species n 699
Appendix C: Selected International Agreements and Conventions
Pertaining to Invasive Species n 707

Appendix D: ISSG’s 100 of the World’s Worst Invasive Alien

Species n 710
Glossary n 713
General Bibliography: Selected Classic and Contemporary Works and
Major Internet Data Sources n 723
Index n 727
n Preface
Invasive species have gained our attention in different ways. Susan Woodward, who wrote
about invasive microorganisms, fungi, and animals in Volume 1, had her interest in invasive
animals first sparked as a student of biogeography in the 1970s. Birds such as the European
Starling and House Sparrow were featured in textbooks to demonstrate how animals spread
in an environment that was new to them or how quickly they evolved adaptations to varying
local conditions across a whole continent. Graeme Caughley’s work on irruptions of red deer
in New Zealand was new, and the modeling of invasions and management of exotics in their
infancy. As a doctoral student at UCLA, she studied feral burros along the lower Colorado
River, viewing them as an example of humans “changing the face of the earth” (the buzz-
words of those days) by transporting domesticated and wild animals around the world.
Under contract to the Bureau of Land Management (BLM) at that time, she collected baseline
data on population dynamics, diet, home range size, and other aspects of burro behavior
and ecology that would help that agency devise policies and practices for the animals’ man-
agement. With an applied aspect to her work, she straddled what has become two main per-
spectives on invasive species in general: an academic interest in the science of invasions and
a management interest in preventing arrivals, eradicating or controlling the spread of those
species that were able to establish populations, and managing those whose numbers and dis-
tribution were for all practical purposes already beyond eradication.
Joyce Quinn, a biogeographer whose research dealt with distribution of plants and their
relationships with the natural environment, such as climate and soils, wrote the invasive
plants section in Volume 2. In spite of her experience, she found that researching and writ-
ing this book led her to learn more. Some plants that she had thought were an integral part
of the “natural” landscape, such as common mullein, are actually alien plants that had
become naturalized and are now widespread in the United States. She notes:

Several years ago, an uninvited plant sprouted unexpectedly in my yard. I tried for years
to get rid of it, dutifully pulling off the little sprouts as they emerged. After four or five
years, I gave up and decided to let the plant grow. In a couple of years, it became an
attractive tree about 10 ft. (3 m) tall. It had smooth speckled bark, long lacy compound
leaves, and clusters of small purple star-shaped flowers. As I was doing research for this
Encyclopedia, I discovered that my new plant was a chinaberry tree. While attractive, it
had few redeeming qualities so I decided to eliminate it. A friend helped me saw the
trunk, about 8 in. (20 cm) in diameter, slightly below ground level. I immediately poured
undiluted glyphosate on the freshly cut stump, thinking that was the end of it. I paid no
attention until four months later when I saw a 6 in. (15 cm) sprout! I sprayed it with her-
bicides, but another sprout soon emerged. I sprayed again, but at the time of this writing,
I still do not know if I have managed to kill the invader. I fear that it will be an on-going
process. If I have had such trouble with just one alien invasive plant, the challenges that
land managers, conservationists, and agriculturalists have in battling invasive species
seem insurmountable.
xiv n PREFACE

Scope
The purpose of the Encyclopedia of Invasive Species is to provide an introduction to the species,
issues, and management options involved with invasive animals, fungi, microorganisms, and
plants. The number of plants and animals introduced into the United States is staggering.
Only a relatively few establish self-sustaining populations, and very few of these actually become
invasive (in the scientific sense of greatly and rapidly expanding their range in the United States).
Still, there are hundreds of invasive species—too many to be included in a reference book of this
sort. For many species, much remains to be learned, and it is premature to develop full entries
for them, but this still leaves many to choose from. In selecting the 168 species for inclusion
in the Encyclopedia, we have tried to offer a wide spectrum of invasive species that includes some
present in the United States from colonial times, and some that have just been detected; some
that completed their spread across the country long ago, and others that are in the midst of rapid
population growth and range expansion. We also wanted to include some species that are found
throughout the country, and some that are limited to a region or single state.
For animals, we aimed to include representatives from all major classes of vertebrates and
a good variety of invertebrates. The reader will find common, well-known invaders and
others that may be a surprise. We also wanted to showcase a few fungi, especially those that
have been major transformers of urban, suburban, and natural forests, and at least acknowl-
edge the presence of invading microorganisms with a tiny sample of those threatening the
health of native animals and, in some cases, humans as well. Finally, we wished to have a
geographically broad selection of invasive animals, with all 50 states and Puerto Rico having
some members of their nonindigenous fauna represented. Florida, Hawai’i, and California
have the largest numbers of officially recognized invasive species. Residents of these states
will undoubtedly find nonnative organisms causing significant impact in natural and artifi-
cial ecosystems missing from our accounts. This was necessary in order to include some
organisms limited to other states.
For plants, we also tried to include a little bit of everything. Volume 2 addresses a variety
of growth forms, ranging from aquatic plants to trees and vines, and all regions of the United
States. Some plants are widespread throughout the country, while others are localized. Many
plants were deliberately brought to the United States as ornamentals or for some useful char-
acteristic, while others were accidentally introduced. The length of entries dedicated to each
species is variable. The taxonomic relationships of some plants and similar species are not
always clearly defined. Some plants, for example, hybridize so freely that it becomes difficult
to distinguish different species. A few accounts of invasive plants treat two or more related
species in the same entry because their effects and management are similar. As with animals,
we could always find “one more” species that should be included, but it was not possible to
include all. A wealth of information from various sources can be accessed by the reader
wanting to know more. The General Bibliography at the end of Volume 2 has a list of recom-
mended resources, including websites.
The Encyclopedia is specifically meant for high school and college students, but addresses
many of the informational needs of the curious naturalist, horticulturalist, or any home-
owner or environmentally concerned citizen who is interested in the origins and consequen-
ces of invasive plants and animals.
Although some invasive species have been part of the landscape of the United States for lit-
erally hundreds of years, the wide-reaching effects of most are only beginning to become real-
ized. Some invasive species are detrimental to native ecosystems and threaten biodiversity,
others are more economically damaging to crops and livestock, and a few pose a danger to
 PREFACE n xv

human health. Invasive species are a major part of current global environmental change.
Experts consider them the second-greatest threat to native species after habitat destruction
and fragmentation. The control and interdiction of invasive species coupled with the damage
some incur on crops, pastures, livestock, native ecosystems, and human health and well-
being costs billions of dollars each year. The invasive species problem is dynamic—as those
in the Mid-Atlantic states weathering their first onslaught of the brown marmorated stink
bug know well—and endlessly fascinating.

How to Use the Encyclopedia

Volume 1 begins with an introduction to inform the reader of the nature and scope of issues
related to invasive species in the United States. Separate sections deal with the terminology
related to invasive species, the invasion process from an ecological point of view, the pathways
by which nonnative species have been and continue to be introduced to the United States,
some of the ecological and economic impacts of invasives, and a brief outline of the history of
modern invasion science. A final section of the introduction describes the human factors that
determine what species come in, where they succeed, and if and how they are managed.
The introduction is followed by 88 entries describing microorganisms, fungi, inverte-
brates, and vertebrates. Entries are arranged alphabetically within major taxonomic groups.
The species described represent the large number introduced and invasive in the continental
United States, Hawai’i, and Puerto Rico.
Each entry in both volumes includes the following elements, unless noted otherwise:

Native Range
Distribution in the United States
Description
Related or Similar Species
Introduction History
Habitat
Diet (animals only)
Life History (animals, fungi, and microorganisms only)
Reproduction and Dispersal (plants only)
Impacts
Management
Selected References

Additionally, each entry in both volumes is accompanied by at least one photograph and
maps that show the original and invasive range of the species in question according to the
best information available. Often range maps are, by necessity, approximate. This is espe-
cially true for organisms not native to the United States or Europe, where biological surveys
are more complete than on other continents.
In Volume 1, the entries are followed by a list of state-by-state occurrences of invasive
animals, fungi, and microorganisms; a glossary; and an index to both volumes.
Volume 2 begins with a brief overview of invasive plants in the United States, which
loosely follows the organization within species accounts, describing in general the scope of
xvi n PREFACE

the invasive plant problem, including the ways, both intentional and accidental, that plants
were brought into the country; and some of the effects invasive plants have on native plant
and animal species, natural ecosystems, agricultural or fishing industries, recreational activ-
ities, or human health. The ways in which invasive plants reproduce and expand their range
is summarized, as is information on management and prevention of invasive plants species.
A sidebar on herbicides accompanies the overview.
The 80 entries on invasive plants are arranged by growth form categories: aquatics, forbs,
graminoids, shrubs, trees, and vines. Photographs of each species show different parts of the
plant. Interesting facets of a plant’s use or history or of strategies attempted for its control are
related in sidebars.
Several supplementary lists follow the invasive plant entries to provide background infor-
mation, and various tables summarize plant data in different, easily accessible ways, includ-
ing a table of common and scientific names of both plants and animals briefly mentioned in
the text of Volume 2, and a list of organizations concerned with invasive plants in the United
States. Two tables of noxious or invasive plants, one organized by state and the other by spe-
cies, as well as a table of species listed by type of impact are also available.
Volume 2 concludes the set with these appendices to the Encyclopedia: a list of American
species that are invasive in other parts of the world; a list of federal laws related to the pre-
vention and management of invasive species; international agreements and conventions
dealing with invasive species; and the IUCN/SCC Invasive Species Specialist Group’s list of
100 of the “World’s Worst Invasive Alien Species,” with an indication of those covered in
the Encyclopedia. The glossary, a selected bibliography of classic and contemporary writings
and online information sources, and the index to the set complete Volume 2.
It is our hope that our efforts will stimulate thought and make the natural world more
accessible to the general public. Informed readers can help make the decisions that will cur-
tail the spread of species that have only recently arrived, prevent the arrival of yet others, and
manage those that are currently invasive

Acknowledgments
Both authors thank the photographers who graciously allowed their photos to be used,
often donating them, or sometimes providing them at a reduced fee. They deserve our spe-
cial thanks for giving life to the species descriptions. Bugwood.org and its associated person-
nel at the Center for Invasive Species and Ecosystem Health, University of Georgia, deserves
special mention as a clearinghouse for providing informational sources and photographs.
Joyce Quinn prepared the excellent maps for the species accounts.
Each author is most appreciative of the other’s contributions to the development of the
project, the overall organization of the volumes, and constructive critiques of text and illus-
trations throughout the manuscript preparation process. We complement each other and
work well as a team. We acknowledge Kevin Downing, originally of Greenwood Press and
now serving the broader ABC-CLIO community as editorial operations manager, who initi-
ated the proposal for the Encyclopedia set, and David Paige of ABC-CLIO, who guided us
through subsequent discussions and organizational details. Anne Thompson, development
editor, later offered guidance in the specifics of the manuscript, and Erin Ryan helped with
the specifications for the illustrations. We thank all four for creating a positive and flexible
working environment and offering valuable suggestions all along the way.
n Invasive Plants in the United States:
A Brief Overview

Although alien invasive plants in the United States come from many regions of the world,
many of them originated in Eurasia, probably because of the long-term connection the
United States has had with European immigrants. The United States has had trade and
immigration from China, Japan, and other eastern Asian countries for more than 100 years,
and some of the alien species native to Asia may have first been introduced to Europe. Few,
if any, places in the United States have been left unaffected by alien invasive plants. Because
of its relatively remote location and extreme climate, Alaska probably has the fewest. In con-
trast, the tropical island environment of Hawai’i is most vulnerable to introduced plants.
Island ecosystems developed independently, without controlling factors such as mammalian
herbivores and predators, so invaders can be especially destructive.
Different growth forms are likely to be invasive in different climates or environments.
Grass and drought-tolerant shrub species present the biggest problems in the arid west,
while moisture-loving vines, trees, and shrubs gain a foothold in the humid eastern states.
Although some species are noxious weeds or invasive throughout the entire country, other
plants can be safely grown as a garden ornamental in climates less suited to them.

Arrival of Alien Plants in the United States

Plant introductions may be either intentional or unintentional. Many of the invasive
plants plaguing the United States were first brought to the country as ornamentals, some
as early as the 1600s, some as late as the 1990s. It is understandable that early immigrants
from Europe and other world regions would want a remembrance of their homeland in their
gardens. We might offer them forgiveness, because they were unaware of how their favorite
plants would behave in a new setting without natural enemies or competition to keep them
in check. Other plants were brought to this country because they were important in house-
hold use, such as for natural dyes or as medicine. Although modern society may sneer at the
supposed medicinal properties of “weeds,” we should remember that early settlers could not
run down to the local pharmacy to purchase relief or a cure for a malady. Some plants
actually are effective, some not.
Some plants, such as kudzu, were encouraged for use by federal agencies, such as the Soil
Conservation Service, as beneficial for erosion control, or as fast-growing forage, such as
Johnsongrass, for livestock. Widespread erosion during the Depression in the 1930s saw
widespread use of exotic plants to reclaim overused or damaged land. U.S. government
agencies were unaware at the time of the potentially devastating effects of many of the spe-
cies they were recommending.
Accidental introductions may be caused by ignorance, carelessness, or just poor luck.
Seeds may adhere to clothing or shoes and then be transported long distances across
expanses that would otherwise be barriers to plant migration. Seeds or plant pieces are small
enough to be overlooked when they arrive mixed into an otherwise-nondescript shipment of
xviii n INVASIVE PLANTS IN THE UNITED STATES

grain or crop seeds. Sand, soil, or water used as ship’s ballast, which is then dumped at U.S.
ports, may carry seeds or pieces of unwanted plants.

Realization of the Problem

The science of ecology is relatively new, and many plant relationships were largely
unknown in the United States until the years leading up to the first Earth Day in 1970,
one of the events that triggered the rise of the environmental movement. It is now realized
by many individuals and many federal and state agencies that alien species without their
natural controls can have disastrous consequences. The proliferation of organizations and
state agencies that classify plants as “noxious” or “invasive” or “prohibited” attest to the
increased awareness of problems associated with introduced species. Biological controls,
meaning insects or pathogens that attack alien invasive plants, are investigated thoroughly
to assure that they do not detrimentally affect related native species or economically impor-
tant crops. Declarations and agricultural screenings at airports and other ports of entry into
the United States attempt to eliminate or at least limit the accidental movement of alien plant
species into U.S. territory.

Impacts of Alien Invasive Plants

Invasive plants not only damage the unique natural environments of the United States,
they also have economic impacts. Alien plants often outcompete native plants for resources,
such as light, water, soil nutrients, or even space. By reducing or eliminating native plants,
they indirectly affect animals that depend on those plants. The result is a reduction in biodi-
versity and the creation of simpler ecosystems that are more susceptible to disturbances. The
United States then loses the natural heritage of a varied landscape and, often, its unique
plants or animals.
Invasive plants have a detrimental economic impact. By encroaching on or taking over
croplands, they reduce crop yields. As alternate hosts, they may introduce diseases or insects
into agricultural areas. Many invasive plants are prickly or otherwise undesirable and hinder
recreational activities, such as hiking, boating, swimming, or fishing. Farmers, ranchers, and
park management agencies spend millions of dollars in attempts to eradicate or merely con-
trol or limit the spread of invasive plants.

Reproduction and Dispersal of Alien Invasive Plants

Like native plants, invasives reproduce and spread either by seed or by sprouting from
vegetative parts. Seeds may be distributed in several ways, including wind, water, animals,
and human activities. Plant pieces are also either deliberately spread, such as when garden-
ers share their extras, or accidentally, such as when carried in a boat trailer to another water
system.
Invasive plants usually do not restrict themselves to the type of habitat they occupied in
their native range. They are usually not invasive in the region where they originated because
competition from other plants, or some other factor such as an insect predator, keeps them
under control. In the absence of competition or a natural enemy in its introduced territory in
the United States, invasive plants may extend to a wider range of environments.
 INVASIVE PLANTS IN THE UNITED STATES n xix

Management Options
No simple solution exists. The problem of eradicating or controlling alien invasive species
is as complex as the array of invaders themselves. The ideal management is to avoid the prob-
lem altogether—to not deliberately plant an invasive species in one’s garden, and to eliminate
what species are already there. This is not always practical or even possible. Cutting down the
tree in your yard will do little to control the invasive species if 20 or more trees grow in the
adjacent woods. The second-best approach is to reduce or eliminate the plant’s potential
spread by preventing seed production or transportation of plant pieces that may sprout.
Some plants respond to physical controls, such as pulling or digging, cutting, mowing, or
grazing. Others require applications of chemical substances, herbicides, which either kill
existing plants or prevent seeds from germinating. A more “natural” way to subdue invasive
plant species is to unleash insect predators or pathogens that harm or control the plants in
some way. Each method has drawbacks. Physical control is usually labor intensive and time
consuming. Chemicals may cause harm to desirable plants or animals and threaten the natural
environment. Unless thoroughly researched, imported biological controls can themselves
become invasive pests. Whatever method is used, it all costs money and takes time.
Management goals—containment, reduction, or eradication—should be realistic for the
situation. While it may not be possible, either physically or monetarily, to totally eradicate
an infestation of an invasive plant, it may be feasible to contain it or to reduce its dominance.
The most basic goal is containment, which means preventing an invasive species from
spreading to new areas, perhaps by physically erecting a barrier or preventing seed produc-
tion. Reduction refers to reducing the amount of area infested by the weed or reducing its
dominance. Total eradication, or completely eliminating the invader, may only be possible
with small stands or new infestations.

What Can We Do?

Everyone seems to have a different definition of what constitutes “alien and invasive,” and
the concept of invasive species just has not reached the public. Some state and federal agen-
cies still recommend and distribute invasive plant species for various “beneficial” purposes.
Several blogs on the Internet tout the benefits of growing attractive alien plants in private
gardens, stating that they do not care if the plant is invasive elsewhere. One blogger even
professed liking the way the plant looked along roadsides.
Although some individuals will demand to retain their “right” to grow the plants they
want, many more citizens will be willing to do the right thing if they only know about the
consequences posed by invasive species. The issue here is ignorance, not as a pejorative
term, but as meaning unaware. Many states and municipalities now have educational pro-
grams to inform citizens of problems associated with alien species. Billboards urge boaters
to clean their boats and trailers so that they do not carry invasive aquatic plants to pristine
water bodies. Signs are posted at trailheads in state parks urging hikers to clean their boots
before leaving or entering a trail, so as to not unwittingly disperse seeds. Native plant soci-
eties urge gardeners to avoid alien invasive plants and to plant native species compatible
with their climate. Although many invasive species have a hold in the United States that is
too strong or entrenched to ever be eradicated, newly invasive species can be controlled or
eliminated if the public is aware of the consequences of some imported plants. Those port-
of-entry screenings are much less of an inconvenience than the potential devastation caused
by an invasive species slipping through.
xx n INVASIVE PLANTS IN THE UNITED STATES

Herbicides

H erbicides are a complicated lot, with several active ingredients sold under
many different brand names. Major categories include preemergent or poste-
mergent, contact or systemic, and nonselective or selective. Each active ingredient,
meaning the substance that kills the plants, has a specific chemical structure that
affects plants differently. Some may damage or kill the plant by interfering in some
way with plant growth, including but not limited to restricting root growth, inhib-
iting cell division, or blocking photosynthesis. Depending on the herbicide and
how it works, it may be applied as a foliar spray, to the basal bark, to the cut stump,
or into slashes cut through the bark. Some herbicides may be toxic to mammals,
fish, or birds, and skin contact may be harmful to some humans. Others may con-
taminate either groundwater or surface water and harm amphibians. Depending
on the plant targeted, herbicides are mixed to different strengths. Many factors in-
fluence the proper selection and use of herbicides.

Types of Herbicides
Preemergent herbicides are also called residuals because they remain in the soil.
They prevent seeds of both annuals and perennials from germinating, and also
sometimes affect shoots sprouting from perennial roots. Although some may tar-
get specific weeds, other preemergents are nonselective, preventing the germina-
tion of seeds of all types of plants. Postemergent herbicides are used on plants
that have grown past the seedling stage.
Contact herbicides are fast-acting and kill the parts of the plant that the chemi-
cals touch. They are usually ineffective for perennials because they do not affect
root growth, and are best used on annuals. Systemic herbicides are absorbed into
the tissue of the plant and moved to other locations, such as the roots. Results
are slower, but the entire plant is killed, making systemics a good choice for
perennials.
Nonselective herbicides affect any type of plant. The most common two catego-
ries of selective herbicides are those that target only broadleaf plants, and those
that target only graminoids. Herbicides selective to broadleaf plants, therefore,
can be used without harm in grasslands, and those selective to graminoids may
be used to eradicate grasses without harm to broadleaf vegetation.

Application Methods
Herbicides may be applied to the foliage, usually by spraying. A surfactant, a sub-
stance added to the spray, is sometimes needed to make the herbicide better
adhere to the leaves, especially if the leaf cuticle is waxy or otherwise imper-
meable. Foliar sprays can be either contact or systemic. In some species, herbicides
may be absorbed into the plant through the bark rather than through the foliage.
This method may be applicable if spraying the canopy may harm nontarget vegeta-
tion. Different herbicides have different effectiveness in basal bark application.
Cutting the trunks or stems close to ground level, then painting the cut stump with
herbicide, is another method that may avoid damaging other plants. It also
requires less product. Hack-and-squirt involves cutting slashes through the bark
directly into the xylem of the plant, usually on a trunk or thick stem. Herbicides
are then injected directly into the wounds.
 INVASIVE PLANTS IN THE UNITED STATES n xxi

Regardless of what method or what type of herbicide is used, the timing of

application is important. Although each species reacts somewhat differently, some
generalizations can be made. Herbicides are frequently most effective when seeds
are being produced, to prevent seed development. Biennial plants are generally
easier to kill in the first season, when they are in an early stage of growth.
Systemic herbicides must be applied when the plants are healthy and actively
growing and translocating sugars to the roots. They are less effective during times
of cold or drought when plants are stressed.
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n Alphabetical List of Invasive Plants
Entries

Entries in the encyclopedia are arranged by growth form categories. Following are the inva-
sive plant entries in alphabetic order.

Asiatic Colubrina (Colubrina asiatica)

Asiatic Sand Sedge (Carex kobomugi)
Australian Pine (Casuarina equisetifolia)
Brazilian Peppertree (Schinus terebinthifolius)
Brooms (Cytisus scoparius, Spartium junceum, Genista monspessulana, and Cytisus
striatus)
Buffelgrass (Pennisetum ciliare)
Canada Thistle (Cirsium arvense)
Carrotwood (Cupaniopsis anacardioides)
Cheatgrass (Bromus tectorum)
Chinaberry (Melia azedarach)
Chinese Lespedeza (Lespedeza cuneata)
Chocolate Vine (Akebia quinata)
Climbing Ferns (Lygodium japonicum and Lygodium microphyllum)
Cogongrass (Imperata cylindrica)
Common Mullein (Verbascum thapsus)
Common Reed (Phragmites australis ssp. australis)
Common St. Johnswort (Hypericum perforatum)
Cordgrasses and Their Hybrids (Spartina alterniflora, Spartina densiflora, Spartina patens,
Spartina anglica, and Spartina alterniflora x foliosa)
Crimson Fountain Grass (Pennisetum setaceum)
Dyer’s Woad (Isatis tinctoria)
English Ivy (Hedera helix)
Eurasian Watermilfoil (Myriophyllum spicatum)
Exotic Bush Honeysuckles (Lonicera maackii, Lonicera morrowii, Lonicera tatarica, and
Lonicera x bella)
Field Bindweed (Convolvulus arvensis)
Fig Buttercup (Ficaria verna)
Fire Tree (Morella faya)
Garlic Mustard (Alliaria petiolata)
Giant Hogweed (Heracleum mantegazzianum)
xxiv n ALPHABETICAL LIST OF INVASIVE PLANTS ENTRIES

Giant Reed (Arundo donax)

Giant Salvinia (Salvinia molesta)
Gorse (Ulex europaeus)
Goutweed (Aegopodium podagraria)
Halogeton (Halogeton glomeratus)
Hydrilla (Hydrilla verticillata)
Ice Plant and Crystalline Ice Plant (Carpobrotus edulis and Mesembryanthemum
crystallinum)
Japanese Barberry (Berberis thunbergii)
Japanese Dodder (Cuscuta japonica)
Japanese Honeysuckle (Lonicera japonica)
Japanese Hops (Humulus japonicus)
Japanese Knotweed (Fallopia japonica)
Japanese Stilt Grass (Microstegium vimineum)
Johnsongrass (Sorghum halepense)
Jubata Grass and Pampas Grass (Cortaderia jubata and Cortaderia selloana)
Kahili Ginger (Hedychium gardnerianum)
Kikuyugrass (Pennisetum clandestinum)
Koster’s Curse (Clidemia hirta)
Kudzu (Pueraria montana)
Lantana (Lantana camara)
Leafy Spurge (Euphorbia esula)
Medusahead (Taeniatherum caput-medusae)
Melaleuca (Melaleuca quinquenervia)
Mile-A-Minute (Persicaria perfoliata)
Multiflora Rose (Rosa multiflora)
Musk Thistle (Carduus nutans)
Oriental Bittersweet (Celastrus orbiculatus)
Paper Mulberry (Broussonetia papyrifera)
Perennial Pepperweed and Hoary Cress (Lepidium latifolium and Cardaria draba)
Porcelainberry (Ampelopsis glandulosa var. brevipedunculata)
Prickly Russian Thistle (Salsola tragus)
Princess Tree (Paulownia tomentosa)
Purple Loosestrife (Lythrum salicaria)
Quackgrass (Elymus repens)
Rattlebox (Sesbania punicea)
Russian Olive (Elaeagnus angustifolia)
Silk Tree (Albizia julibrissin)
Spotted Knapweed (Centaurea stoebe)
Strawberry Guava (Psidium cattleianum)
 ALPHABETICAL LIST OF INVASIVE PLANTS ENTRIES n xxv

Swallow-Worts (Cynanchum rossicum and Cynanchum louiseae)

Tamarisk (Tamarix chinensis, Tamarix ramosissima, Tamarix parviflora, and Tamarix
gallica)
Toadflax (Linaria dalmatica ssp. dalmatica and Linaria vulgaris)
Tree of Heaven (Ailanthus altissima)
Tropical Soda Apple (Solanum viarum)
Velvet Tree (Miconia calvescens)
Water Chestnut (Trapa natans)
Waterhyacinth (Eichhornia crassipes)
West Indian Marsh Grass (Hymenachne amplexicaulis)
Winter Creeper (Euonymus fortunei)
Wisteria (Wisteria sinensis and Wisteria floribunda)
Yellow Himalayan Raspberry (Rubus ellipticus)
Yellow Starthistle (Centaurea solstitialis)
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n Aquatic Plants
n Eurasian Watermilfoil
Also known as: Spike watermilfoil
Scientific name: Myriophyllum spicatum
Synonyms: None
Family: Watermilfoil (Haloragaceae)
Native Range. Widely distributed in Europe, Asia, and North Africa.
Distribution in the United States. Found in every state except Wyoming and Hawai’i. A
major problem in the Northeast, Upper Midwest, Gulf Coast drainages, Southwest river
systems, and the Pacific Northwest, it is still expanding its range.
Description. Eurasian watermilfoil is an herbaceous aquatic perennial that is submersed,
meaning it grows primarily under the water surface. It usually grows in water depths of
3–13 ft. (1–4 m), but can be rooted on bottom sediments as deep as 33 ft. (10 m) beneath
the surface. The species is variable in appearance. The slender, hairless stems have no leaves
toward the base due to lack of light. Growing stem tips are often reddish brown. Upon
reaching the water surface, plant stems branch and form a dense canopy. Plants also branch
when they are cut, browsed or otherwise damaged. In shallow water, stem densities can
exceed 30 per sq. ft. (300/m2). Olive-green leaves, 4–5 per node, are whorled around the
stem. Each leaf is finely divided into narrow segments or leaflets, usually 12–16 pairs, but
sometimes as few as 5 or as many as 24. The soft, feathery leaflets are 0.5 in. (1.3 cm) long.
When removed from the water, leaflets become limp and cling to the stem above the node.
With cooler fall temperatures, stems die back to the propagating root crowns and plants
overwinter rooted in bottom sediment. In warmer areas, plants can continue growing
throughout the winter months. Carbohydrate storage is in the shoots and roots, and plants
do not grow turions. Growth begins rapidly when spring temperatures near 59ºF (15ºC).
Roots are fibrous and often become fragmented.
Beginning in early summer and continuing for several months, flowers develop on red-
dish spikes that extend 2–4 in (5–10 cm) out of the water. Stems beneath the spikes become
thickened and curved to lie parallel with the surface. Small yellow or pinkish, four-part
flowers occur in whorls of four (rarely five) around the inflorescence, with female flowers
at the base and male flowers at the top. After pollination, flower spikes sink down into the
water. Four small hard capsules, 0.05–0.1in (1–3 mm), develop from each flower. When
flowering is complete, stems fragment, growing roots at the nodes before breaking away. If
conditions are not suitable, or stems and leaves are eaten by herbivores, plants may not
reach the surface and flower.
Related or Similar Species. Ten species of Myriophyllum are native to the eastern United
States, but taxonomy is unclear, causing confusion in identification. The number of leaf divi-
sions, 5–24 pairs, is usually sufficient to distinguish Eurasian watermilfoil from other species.
The most closely related species native to North America is northern watermilfoil, also known
as shortspike watermilfoil, with 5–10 leaflet pairs that remain rigid when removed from water.
The stem beneath the flower spikes of the northern watermilfoil is straight and not thickened.
322 n AQUATIC PLANTS

Northern watermilfoil forms

blackish-green turions, or scaly
shoots from underwater buds,
whereas the Eurasian species
does not.
Parrotfeather, native to the
Amazon River in South America,
is commonly used in aquariums
and garden ponds. Although it
is less pervasive than Eurasian
watermilfoil, it forms denser
stands which provide more
breeding habitat for mosquitos.
Leaves, in whorls of 4–6, occur
in two types, submersed and
emergent. Submersed leaves,
with 20–30 leaf divisions, are
most commonly confused with
Eurasian watermilfoil. The emer-
gent leaves have only 6–18 divi-
sions, and are bright green. The
most distinctive difference is that
the emergent leaves and stems
grow as much as 1 ft. (0.3 m)
up out of the water. This plant
causes problems similar to those
of Eurasian watermilfoil, and is a
noxious weed in several states.
Although native to the sou-
thern United States and endan-
gered in Kentucky, Ohio, and
After its initial introduction to the Eastern Seaboard, Eurasian Pennsylvania, variable water-
watermilfoil spread rapidly throughout the Northeast, into the milfoil is an invasive species in
Tennessee River valley, and into the Pacific Northwest. (Native range some New England states. It
adapted from USDA GRIN and selected references. Introduced range has two types of leaves. The
adapted from USGS Nonindigenous Aquatic Species Database and whorled leaves below the water
selected references.)
surface are reddish-green to
brown, very finely dissected or
threadlike, and 0.5–2.5 in. (1.4–6.4 cm) long. Emergent leaves, which grow on stalks rising
6–8 in. (15–20 cm) out of the water, are small, 0.25 in. (0.6 cm) and oval. The native
coontail, also called common hornwort, differs from Myriophyllum species in that it is a float-
ing plant that does not root. Its toothed leaves, making it feel rough to touch, are bright
green, stiff, and brittle. The whorled leaves, which curve upward to resemble a tail, remain
rigid when removed from water. It is not invasive and is an important part of natural
ecosystems.
Introduction History. The introduction of Eurasian watermilfoil to North America is not
clear. It may have been accidentally introduced into the northeastern United States some-
time between the late 1800s and 1940, perhaps in ship ballast in the late 1800s. Reports
 EURASIAN WATERMILFOIL n 323

A. Finely divided leaves are whorled around the stem. (Chris Evans, River to River CWMA, Bugwood.org.)
B. Leaflets cling to the stem when the plant is out of the water. (Richard Old, XID Services, Inc., Bugwood.org.)
C. Tiny flowers are whorled around the flower spikes. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology
University of Connecticut, Bugwood.org.)

prior to 1940, however, may be due to misidentified plants. The first recorded collection
was in Washington, D.C., in fall 1942. By 1985, it had spread to 33 states and adjacent parts
of Canada. Plants may have been intentionally introduced to the Tennessee Valley Authority
(TVA) system in 1953 by a resort owner, and some anglers have deliberately stocked lakes or
ponds. Some Oklahoma populations developed when earthworm farmers used Eurasian
watermilfoil for packing material. Long-distance dispersal can be attributed to the aquarium
and water garden trade, through which it is still sold.
Habitat. Eurasian watermilfoil tolerates a wide range of environmental conditions, from fro-
zen ponds in Minnesota to shallow, warm bays in Florida. It primarily invades disturbed areas
where native plants are unable to adapt to changes, and is less likely to spread into healthy eco-
systems. Although it prefers quiet, stagnant water commonly found in ponds, lakes, reservoirs,
pools, ditches, and irrigation canals, either deep or shallow, it also grows on muddy shores
and in flowing water. Because it tolerates fresh to slightly brackish water, with salinity as high
as 15 ppt, it is found in estuaries. It is also tolerant of many types of water pollution, as well as
pH levels 5.4–11. Because it is herbaceous and protected by water, it survives in ice-covered
lakes. It requires high light levels compared with other submersed plants, but grows rapidly over
a broad temperature range.
Reproduction and Dispersal. Although plants reproduce by seed, vegetative growth is
responsible for its spread in North America. New plants, which grow from each stem node,
will root in the muddy bottom. Stems become naturally broken apart by wind and waves,
and by self-fragmentation after flowering. Boat traffic or weed cutting machines will also
cut and spread fragments. Fragments are inadvertently carried on boats and trailers, and
by currents downstream in river systems.
Seedlings are rare. Seeds need temperatures higher than 57ºF (14ºC) to germinate, but
amount of light is not important. Too much sediment, however, impedes germination.
Impacts. Eurasian watermilfoil is considered the worst waterweed in the continental
United States. Millions of dollars are spent nationwide in control. Several states, including
Washington, Minnesota, Wisconsin, Vermont, and New York, spend approximately $1 million
each year. Although no direct financial assessment exists, infestations of Eurasian watermilfoil
negatively affect recreational use of water bodies, making boating, swimming, and fishing
324 n AQUATIC PLANTS

unpleasant, if not impossible. Plant canopies in stagnant water provide ideal mosquito habitat.
Dense stands of plants clog irrigation systems and intakes for hydropower.
By growing rapidly and creating dense canopies on the water surface, Eurasian watermil-
foil reduces light penetration early in the growing season, thereby shading out native plants,
such as the native northern watermilfoil. Although the northern watermilfoil is able to suc-
cessfully compete in aquarium tank studies, it fails to do so in the field. With decreased
plant biodiversity, fewer animals, including benthic invertebrates and native fish, can sur-
vive. Fish have fewer spawning areas and do not grow as large. Decomposition of so much
biomass at the end of summer changes water chemistry, increasing phosphorus and nitrogen
levels. Eurasian watermilfoil also decreases the amount of oxygen beneath the mats, and
raises both the temperature and pH of the water.
Management. Thorough inspection and cleaning of boats and trailers may prevent frag-
ments from being carried to noninfested water bodies. Several methods of control also have
negative impacts on native plants, fish, and other wildlife. Mechanical harvesting and herbi-
cides kill beneficial insects and may do more long-term harm. Mechanical harvesters, rototillers,
and dredges are also expensive, labor intensive, and disruptive to the environment. Physical
removal provides short-term control, while insect predation represents long-term control.
Physical means of control are ineffective in the long term because broken pieces of Eurasian
watermilfoil will grow into new plants. Small plants may be hand-pulled, but all pieces must
be removed from the water. Mechanical harvesting can control growth in larger areas, but
should be done more than once in a summer season. Harvesting implies that cut plants are
removed and disposed of elsewhere. Rototilling the bottom pulls up the roots, which can then
be removed. Water level manipulation may be practical in reservoirs, canals, or ditches, but
not in natural areas. A higher water level may limit light to the plants, effectively shading them
out. Lower water levels expose plants to dehydration in summer and freezing in winter.
Drawdown of 6.5 ft. (2 m) in the Tennessee Valley Authority system in the Southeast effec-
tively reduces populations. Light may also be reduced by planting vegetation on banks or
erecting shade structures. Vertical screen barriers inserted to a water depth of 13 ft. (4 m) will
confine the infestations and prevent plant fragments from moving into other parts of the water
body, perhaps a way to keep docks or swimming areas free. Those types of areas may also be
kept clean by covering the sediment with a light-blocking substance.
Care must be taken with any attempt at chemical control because herbicides may also kill
fish and nontarget aquatic plants, cause more algae growth, and contaminate the water sup-
ply. Not all herbicides are approved for use in all types of water bodies, and selection of
product must take into account whether control is needed in a natural lake or a roadside
ditch. Some herbicide applications will suppress growth for six weeks up to one year.
Fluridone is selective for watermilfoil and other nonnative weeds and is not harmful to
recreation activities or drinking-water supplies. One application controls weeds for the sea-
son. Also selective, triclopyr and 2,4-D afford excellent control and do not affect native
aquatics. Other herbicides have also been found to be effective.
Research on insects and pathogens for potential biological control has been conducted for
30 years. Over 20 insect species have been observed feeding on Eurasian watermilfoil in its
native range, but few have been investigated seriously because they do not appear to be host
specific. Several species, some native to North America and some accidentally introduced,
are possibilities. Three small herbivores, a pyralid moth (Acentria ephemerella), the milfoil
midge (Cricotopus myriophylli), and a weevil (Euhrychiopsis lecontei) have been associated
with declines in populations of Eurasian watermilfoil in some lakes. The moth, native to
Europe but now widespread from the East Coast west to Minnesota and Iowa, prefers
 EURASIAN WATERMILFOIL n 325

Eurasian watermilfoil but does not feed on it exclusively. Each female lays 100–300 eggs,
often with two generations per year. The larvae feed on the apical meristems, reducing plant
growth, height, and biomass, and inhibiting canopy formation. The milfoil midge may have
been accidentally introduced to North America because it prefers Eurasian watermilfoil over
the native northern watermilfoil. It is now widely distributed in the northern states from the
West Coast to New York, but only in areas that support Eurasian watermilfoil. Its larvae may
reduce the plant’s biomass by boring into the stems. Although it may be present in many
lakes, it is difficult to identify. The weevil, native to and widely distributed in North
America, is a watermilfoil specialist. All life stages eat tips, stems, and leaves. Although it ini-
tially fed on native northern watermilfoil, the weevil now prefers the introduced Eurasian
watermilfoil. This weevil is now available commercially, but needs shoreline habitat where
adults overwinter. Development of shorelines with houses and other structures inhibits its
establishment.
Inconsistent results with insects may be caused by slight chemical differences, such as
nitrogen content, in different populations of Eurasian watermilfoil, which in turn provide
different nutritive value to feeding insects, altering their vigor. Similarly, waterfowl in
Alabama eat a lot of Eurasian watermilfoil, but the same species avoid it in the Great Lakes
region.
Grass carp (see Volume 1, Vertebrates, Fish, Grass Carp) (Ctenopharyngodon idella), a fish,
is used for control of several aquatic pest species. Grass carp, however, prefer other food and
only eat Eurasian watermilfoil if nothing else is available. Their presence can also deplete
desired plant biomass, limiting food and shelter for native fish, invertebrates, and waterfowl.
Most states do not allow importation of grass carp. Two fungi, Mycoleptodiscus terrestris and
Colletotrichum gloeosporioides, are possibilities.

Selected References
Agricultural Research Service. “Foiling Watermilfoil.” Agricultural Research Magazine, 1999. Accessed at
United States Department of Agriculture. http://www.ars.usda.gov/is/AR/archive/mar99/
foil0399.htm.
Johnson, R. L., and B. Blossey. “Eurasian Watermilfoil.” In Biological Control of Invasive Plants in the
Eastern United States, by R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-04,
Morgantown, WV, 2002. http://www.invasive.org/eastern/biocontrol/6EurasianMilfoil.html.

Successful Eradication of Eurasian Watermilfoil

in Cayuga Lake

E urasian watermilfoil dominated the waters of Cayuga Lake, one of the Finger
Lakes of upstate New York, to the exclusion of most other water plants until
1991 when large populations of the pyralid moth were discovered there. Ten years
later, the biomass of Eurasian watermilfoil was only 10 percent of what it had
been, with no canopy, resulting in long-term control and reversion to the natural
ecosystem. Native plants have repopulated the lake, economic viability for
recreation-based communities has resumed, and herbicide use has been reduced.
326 n AQUATIC PLANTS

“Parrotfeather (Myriophyllum aquaticum).” Non-native Invasive Freshwater Plants, Department of

Ecology, State of Washington, n.d. http://www.ecy.wa.gov/Programs/wq/plants/weeds/
aqua003.html.
Ramey, Victor. “Eurasian water-milfoil, Myriophyllum spicatum.” Center for Aquatic and Invasive Plants,
University of Florida, 2001. http://plants.ifas.ufl.edu/node/278.
Rook, Earl J. S. “Ceratophyllum demersum, Common Hornwort.” 2002, 2004. http://www.rook.org/earl/
bwca/nature/aquatics/ceratophyllum.html.
Rook, Earl J. S. “Myriophyllum heterophyllum, Two Leaf Milfoil.” 2002, 2004. http://www.rook.org/earl/
bwca/nature/aquatics/myriophyllumhet.html.

n Giant Salvinia
Also known as: water velvet, salvinia, African pyle, aquarium watermoss, kariba weed,
water fern, koi kandy, floating fern
Scientific name: Salvinia molesta
Synonyms: Salvinia auriculata
Family: Floating Fern (Salviniaceae)
Native Range. Coastal regions, as far as 125 mi. (200 km) inland, of southeastern Brazil
in the states of Sao Paulo, Parana, Santa Catarina, and Rio Grande do Sul, and in
northeastern Argentina. From 24º S to 32º S latitude.
Distribution in the United States. Over 40 drainage basins, primarily along the Gulf
Coast and Atlantic Coast, from Texas east to Florida, north to Virginia, and in New York
State. Also in Arizona, California, Hawai’i, and Puerto Rico. It is still expanding its range.
Description. Giant salvinia is a free-floating fern. Horizontal stems, or rhizomes, float just
below the water surface. The stems produce three leaves at each node, two floating or emer-
gent and one submergent. The oval or round floating leaves, green or yellow-green, are
opposite each other on the surface. Young floating leaves are thin, 0.3–0.6 in. (8–15 mm)
wide, and lie flat on the water surface. Leaf size varies with space and nutrient conditions,
but leaves of mature plants are usually heart-shaped or oblong, 0.5–1.5 in. (1.3–3.8 cm)
wide and sometimes as wide as 2.4 in. (6 cm) when unfolded. They may become brown at
the edges as they age. The crowding of plants forces the leaves of older plants to fold in
the center, creating chains of upright leaves extending out of the water. The upper surface
of leaves is covered with rows of bristly hairs. Each hair stalk divides into four extensions
that rejoin at the tips to form a cage that resembles an eggbeater. These structures can be
seen with minor magnification, such as a hand lens. The hairs, giving plants a velvety
appearance, are an air trap that provides buoyancy. Hairs on the upper leaf surface repel
water, while the short, straight hairs on the lower leaf surface attract water, a combination
that keeps the leaf properly oriented. Although possibly damaged in older leaves, the hairy
structures are clearly visible on young leaves.
Giant salvinia has no roots. The underwater structures that resemble roots are filamen-
tous fronds or leaves. The third leaf growing from the surface node is highly divided into
brown or whitish filaments, which dangle in the water. The filaments are covered with very
short hairs. The length of the filaments, several inches, stabilizes the floating plants.
Descending stalks among the root-like leaves hold chains of egg-shaped spore cases.
Related or Similar Species. Several plants closely related to giant salvinia, called the
Salvinia auriculata complex, include Salvinia auriculata, S. biloba, and S. herzogii. All have
the same type of eggbeater hairs, but are difficult to identify. Although all are federal noxious
weeds, none occur in the United States.
 GIANT SALVINIA n 327

Common salvinia, also

known as water spangles, is an
invasive aquatic species from
Central and South America that
closely resembles giant salvinia.
It produces two round or oval
floating leaves, 0.2–0.75 (0.4–
2.0 cm) long, from each rhi-
zome node. Older leaves fold
at the midrib and stand more
erect. In shady locations, the
leaves remain round and emer-
ald green; leaves in full sun
become larger, longer, and
rusty brown. The third leaf
from the node is the submerged
filaments. Both the underside of
the floating leaves and the fila-
ments are covered with long
hairs. The most distinguishing
feature is that the hairs on the
upper leaf surface are not joined
at the tip, and therefore lack the
eggbeater structures.
Mat-forming waterfern plants
native to the United States
include three Azolla species,
all of which have very small
leaves. Mosquito fern, native
to midwestern and eastern
North America, is less than
0.5 in. (1.3 cm) wide, with First found growing in a natural water body in 1995, giant salvinia is a
lacy-looking leaves. The gray- recent invader dispersed by the aquarium and water garden trades.
green or rusty-red leaves fre- (Native range adapted from USDA GRIN and selected references.
quently overlap, giving the plant Introduced range adapted from USGS Nonindigenous Aquatic Species
a quilted look. It can form large Database and selected references.)
mats, 1.5 in. (4 cm) thick, in
stagnant water. Mexican water-fern, native to the midwestern and western United States, has
tiny (1 mm), overlapping leaves. Each leaf has two lobes, one on the surface and one sub-
merged. Pacific mosquitofern plants, native to the West Coast, are 0.4–0.8 in (1–2 cm) across,
with green overlapping leaves, sometimes tinged with pink, orange, or red. Species of water
clover, so called because their leaves are divided into four parts resembling clover, have leaves
which are borne on long stalks either projecting out of the water or submerged.
Introduction History. Giant salvinia was first recorded growing in a natural area in 1995,
in a private pond in South Carolina, from which it has since been eradicated. It is believed to
have been introduced via the aquarium and water garden trade, subsequently discarded into
a waterway when the plants outgrew the aquarium or pond. Flooding may also have acci-
dentally released the species from pond cultivation. The plant is still offered for sale in
328 n AQUATIC PLANTS

A. Floating plants can completely cover a lake. (Ted D. Center, USDA Agricultural Research Service,
Bugwood.org.) B. Leaves are usually heart-shaped. (© Barry Rice, Sarracenia.com.) C. The upper surface of
leaves is covered with bristly hairs. (© Barry Rice, Sarracenia.com.) D. Under crowded conditions, the leaves fold
in the middle. (Troy Evans, Great Smoky Mountains National Park, Bugwood.org.) E. “Egg beater” cages on the
leaf hairs. (© Barry Rice, Sarracenia.com.)

nurseries and mail-order suppliers in several states, and is frequently a contaminant in

aquatic nursery plant stock.
Common salvinia, which has been cultivated in gardens and greenhouses since the late
1880s, is still widely available for sale. It is also a common contaminant in other water plant
stock.
Habitat. Giant salvinia is found in quiet, nutrient-rich water, such as lakes or ponds,
oxbows and backwater swamps, marshes, rice fields, ditches, and any slowly flowing water
courses. Plants grow only in freshwater and are intolerant of as little as 11 ppt salt. Growing
in tropical, subtropical, or warm temperate climates, the species can tolerate freezing air
temperature but not ice formation in the water. Giant salvinia is capable of overwintering
in mild, southern climates.
Plants do best at water temperatures of 68–86ºF (20–30ºC), and dense mats do not
develop at temperatures lower than 50ºF (10ºC). Exposure to 26.5ºF (−3ºC) or above
109ºF (43ºC) will kill buds in two hours’ time.
Reproduction and Dispersal. Although giant salvinia has many spore cases, it is not
known to be fertile, and its means of reproduction is strictly vegetative. Each rhizome node
has as many as five buds that produce new rhizome branches, which radiate outward. Stress
from cool temperatures or desiccation stimulates latent buds to sprout. Rhizome pieces are
transported to other water bodies by boats, propellers, boat trailers, jet ski intakes, and other
water gear. Young plants or pieces are carried downstream by currents or are distributed
when ponds or lakes flood and overflow. Wind may also push floating plants to new loca-
tions within water bodies. The plant’s hairiness minimizes desiccation when it is transported
out of the water.
Impacts. The vegetative reproductive method of giant salvinia allows it to rapidly create a
network of plants that grow into dense mats covering the water surface. Under ideal green-
house conditions, it can double its numbers every 2–4 days. In one month, one plant can
multiply to 8,000 plants, and to more than 67 million plants in two months. Under good
conditions and warm weather, the size of a mat in a natural environment may double in
7–10 days. Vegetation mats may cover entire lakes or other water bodies. In other countries,
 GIANT SALVINIA n 329

Recuperation of Invasive Species?

A fter the British Petroleum oil spill in the Gulf of Mexico in April, 2010, the
governor of Louisiana ordered the Davis Pond Freshwater Diversion Project
to be fully opened, an attempt to stem the inflow of oil by flooding the area with
fresh water. The influx of so much fresh water lowered the salinity of the water-
ways and allowed freshwater invasive species to gain a foothold. Giant salvinia
was one of the plants identified in November to be taking advantage of the lower
salinity levels, and it is questionable whether the species can be eliminated in this
new environment.
Source: Bob Marshall, The Times-Picayune, “Freshwater areas’ next big battle has arrived.”
November 21, 2010. www.nola.com/outdoors/index.ssf/2010/11/freshwater_areas)_next_big_
batt.html.

mats covering 96 sq. mi. (250 km2) and measuring 2–3 ft. (0.6–0.9 m) thick have been
recorded, indicating its potential. One individual plant was documented to have increased
to cover 40 sq. mi. (104 km2) in three months. Plants grow faster when water is saturated
with nutrients, perhaps resulting from fertilizer runoff.
Mats alter ecosystems by blocking light, which kills algae and other organisms that serve
as food for fish and other species higher on the food chain. Plants alter the water quality,
lower the pH level, and increase temperature and carbon dioxide content. Decaying vegeta-
tion uses oxygen, depleting the supply for fish, insects, and other aquatic organisms.
At Enchanted Lake on O’ahu, Hawai’i, giant salvinia threatens nesting habitats of endan-
gered waterbirds, including the Hawaiian Coot, Hawaiian Gallinule, and the Hawaiian Stilt.
Although populations of giant salvinia have been reduced in that location, the plant contin-
ues to expand its range to other sites, including the big island of Hawai’i.
Its rapid spread may threaten cultivated aquatic crops, such as rice. It clogs water lines
carrying drinking water, irrigation canals, and intakes for hydroelectric power plants. The
dense mats interfere with boating, fishing, and recreational activities or water sports.
Common salvinia has escaped into natural areas by flooding or by intentional release,
growing into dense mats in Texas and Louisiana, where it is a noxious weed. In Louisiana,
it is documented to have covered a waterway 12 mi. (19.3 km) long and 360 ft. (110 m)
wide as thickly as 8–10 in. (20–25 cm). It may displace native duckweed, which is an
important food for waterfowl. It may also dominate native mosquito fern in summer.
Management. New infestations of giant salvinia can be prevented by thorough washing of
all boating and other equipment after leaving the water. Even one piece can start a new pop-
ulation. Although small infestations can be removed by hand, the species must be identified
and removed quickly because the plant spreads rapidly. Other alien floating plants, such as
waterhyacinth, however, may invade after giant salvinia is eradicated.
Physical methods of removal, either by hand or by mechanical harvesters, are only fea-
sible for small infestations that have not yet become established. All pieces should be bagged
and disposed of in an upland landfill, not near water. Because giant salvinia does not tolerate
salinity, infestations can be controlled by flooding with seawater.
Although chemical applications may be effective, they require long-term contact.
Repeated applications of fluridone, hexazinone, glyphosate, or diquat may control the plant.
330 n AQUATIC PLANTS

Chemical control is further complicated because masses of decaying plants severely deplete
the oxygen in the water.
Because several insects feed on giant salvinia in its native Brazil, where it does not form
dense mats, biological control is possible. Large populations, however, may also need her-
bicide applications. A host-specific weevil (Cyrtobagous salviniae) from giant salvinia’s
native range in southeastern Brazil and adjacent countries has succeeded in suppressing
populations in Australia, Asia, and Africa. This tiny insect, 0.1 in. (2.5 mm) long, lives
in or on the leaves and submerged rhizomes. After the eggs hatch in a cavity chewed into
the leaf bud by the female, the larvae eat the leaf bud base and tunnel into the rhizomes
and petioles of the plant. Until temperatures drop below approximately 75ºF (24ºC), this
weevil can generate a new generation every month. Populations can reach 30 adults and
90 larvae per sq. ft (300 adults and 900 larvae per m2) of water surface. Cyrtobagous salvi-
niae is host-specific and will not eat North American native water plants, such as mosquito
fern or water clover.
In preliminary trials in Texas, using Cyrtobagous salviniae weevils collected from common
salvinia plants in Florida, results were inconclusive because of other variables such as flood
and drought, which affected the weevils. Confusion exists regarding the identity of those
weevils, however, and it is believed that the weevils in the United States may be genetically
distinct from the weevils that are very effective in other parts of the world. Release of
Cyrtobagous salviniae imported from Australia is pending positive identification of the weevil
species collected in the United States.

Selected References
Jacono, C. C. “Common—Salvinia minima Identification.” U.S. Geological Survey, 2003. http://
salvinia.er.usgs.gov/html/identification1.html.
Jacono, C. C. “Giant Salvinia—Salvinia molesta Identification.” U.S. Geological Survey, 2003. http://
salvinia.er.usgs.gov/html/identification.html.
Julien, M. H. “Floating Fern (Salvinia).” In Biological Control of Invasive Plants in the Eastern United
States, by R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-04, Morgantown,
WV, 2002. http://wiki.bugwood.org/Archive:BCIPEUS/Floating_Fern_(Salvinia).
National Biological Information Infrastructure (NBII), Comité français de l’UICN (IUCN French
Committee), and IUCN SSC Invasive Species Specialist Group (ISSG) “Salvinia molesta (Aquatic
Plant, Herb).” ISSG Global Invasive Species Database. 2005. http://www.issg.org/database/species/
ecology.asp?fr=1&si=569
“Salvinia Complex.” Plant Pest and Health Prevention Services (PHHPS). California Department of
Food and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/salvinia.htm.

Successful Biological Control

C yrtobagous salviniae weevils, from Brazil, were released in Australia in 1980. In

only 14 months, they succeeded in eliminating 90 percent of giant salvinia from
Lake Moondarra, which was previously almost completely covered with plants. This
weevil destroyed more than 8,000 tons of the plant in less than a year.
Source: Julien, M. H. 2002.
 HYDRILLA n 331

n Hydrilla
Also known as: water thyme, waterthyme, Florida elodea, Indian star vine
Scientific name: Hydrilla verticillata
Synonyms: None
Family: Frog’s-bit (Hydrocharitaceae)
Native Range. Probably from the warmer areas of southern Asia and India and Korea.
Hydrilla is so widely spread throughout the world, on all continents except Antarc-
tica, that its native range is difficult to determine. The dioecious form in the United States
is native to India, while the monoecious form is native to Korea.
Distribution in the United States. Most of the eastern half of the country. The Atlantic
Coast and Gulf Coast states,
from Maine south and west to
Texas; north through Okla-
homa, Arkansas, Missouri, Iowa,
Wisconsin, and Upper Michi-
gan, and east to Maine. Western
states, including Colorado,
Arizona, Nevada, California,
Idaho, and Washington. Also in
Puerto Rico. Most invasive in
the southern and coastal states.
The dioecious form is found in
the southern states, while the
monoecious form is found north
of North Carolina. A population
in Connecticut derived from the
dioecious strain is expanding its
range, with new populations
being discovered in the northern
and midwestern states.
Description. Hydrilla is an
herbaceous perennial that grows
submersed in freshwater. Plants
are most often rooted in bottom
sediments, but broken pieces
may be free-floating. Its appear-
ance, such as number of leaves,
length of stem, and leaf shape,
is variable and dependent on
age and environmental condi-
tions. Slender stems growing up
from the bottom may be 6.5 ft.
(2 m) or more long, depending Two forms of hydrilla are in the United States, a dioecious form in the
on the water depth. Some have warmer South and a monoecious form in cooler, northern climates.
been measured at 30 ft. (9 m). (Native range approximated from selected references. Introduced range
Stems of dioecious plants in adapted from USGS. Aquatic Species Database and selected references.)
332 n AQUATIC PLANTS

A. Plants may form dense floating mats. (David J. Moorhead, University of Georgia, Bugwood.org.) B. Narrow,
pointed leaves are whorled around the stem. (Robert Vidéki, Doronicum Kft., Bugwood.org.) C. Plant pieces
entangled in boat motors may be dispersed to other water bodies. (Wilfredo Robles, Mississippi State University,
Bugwood.org.) D. Small tubers can produce many new plants. (Robert Vidéki, Doronicum Kft., Bugwood.org.)

deep water have few branches, but they branch profusely upon reaching the surface, forming
dense vegetation mats. Monoecious plants, however, branch at the sediment rather than at
the surface, sending a denser array of stems growing up through the water column. Small,
0.25–0.8 in (0.6–2 cm) long, narrow, pointed leaves, with finely toothed margins, grow in
whorls of 3–10 along the stems. The lower surfaces of leaves on mature hydrilla sometimes
have small sharp spines on the reddish midrib, making the plant feel scratchy. The green
leaves are translucent when clean, but the top leaves, sun bleached or damaged by bacteria,
may range from yellowish to brownish green. Leaf axils have pairs of tiny scales, only
0.5 mm long, with an orange-brown hairy fringe. Turions, which are condensed shoots 0.25
in. (0.6 cm) long with whorls of fleshy leaves, grow in leaf axils. Turions are dark green and
look spiny. In cold climates, the stems and leaves die back seasonally, but the tubers and
turions send up new stems in spring.
Small, pale tubers, 0.2–0.5 in (0.5–1.3 cm), that resemble tiny potatoes develop at the
ends of stolons or rhizomes, both on and below the surface of the bottom sediment where
the plant is rooted. They may be as deep as 12 in. (30 cm) into the sediment.
Tiny female flowers grow singly on long, 4 in. (10 cm), thread-like stalks from leaf axils
near the top of the stems. Three whitish sepals enclose three transparent petals. The
funnel-shaped flowers float upright on the surface, the petals keeping the flower centers
dry. Tiny male flowers, also with three sepals and three petals, range in color from whitish,
reddish, brownish, to greenish. Male flowers break loose from the leaf axils and float freely
on the water surface, looking like tiny inverted bells. Pollination is by wind.
Related or Similar Species. Hydrilla may be difficult to distinguish from the nonnative
Brazilian waterweed, also called Brazilian elodea, or the native Canadian waterweed with
which it often grows. Brazilian waterweed has 4–5 leaves on each whorl, and the fine serra-
tions on the leaf margins are only detectible under magnification. Canadian waterweed has
whorls of 3–6 leaves. Neither plant has tubers.
 HYDRILLA n 333

Introduction History. An aquarium dealer shipped the dioecious variety of hydrilla plants
from Sri Lanka to Florida in the 1950s. After being deemed unsatisfactory, the plants were
dumped into a canal near Tampa Bay. In 1955, some of the plant population was taken to
Miami for the pet trade, where careless or accidental release allowed it to infest many water
bodies. By 1988, it had spread to 49,500 ac. (20,000 ha) of water bodies in Florida. The
monoecious strain, from Korea, was first reported in Delaware in 1976 and in the Potomac
River around 1980.
Habitat. Other than a need to be submerged in freshwater, hydrilla has few specific require-
ments. It grows in still or slow-flowing water of various depths in tropical to temperate climates
where water temperatures can be 50–95ºF (10–35ºC). Temperatures of 68–81ºF (20–27ºC) are
optimal for photosynthesis, but hydrilla will continue to grow slowly at lower temperatures. It
can grow in clear water with low nutrients, or in murky, nutrient-rich water. Although opti-
mum pH is 7, hydrilla grows within a range of acidity. Plants are found in many types of water
bodies, including lakes, ponds, reservoirs, springs, rivers, canals, ditches, tidal zones, and rice
fields. It tolerates polluted water and has moderate tolerance for salinity, up to 7 percent,
allowing it to infest the upper reaches of estuaries such as Chesapeake Bay. Because it can grow
with little sunlight, it is able to colonize deeper water than most native aquatic plants.
Reproduction and Dispersal. Hydrilla is one species with two forms, a monoecious and a
dioecious biotype. The southern biotype is dioecious, with only female plants. Although
they flower, they cannot set seed. The northern biotype is monoecious, with the male and
female flowers on the same plant, but seeds are rare. The only known seed production in
the United States is in monoecious stands in North Carolina. Seeds mature in September
and sink to the bottom sediment, where they germinate the following spring. Although,
the germination rate is less than 50 percent, new plants can flower after two months.
Hydrilla reproduction is almost exclusively vegetative, from stem fragments, stolons, or
rhizomes and turions. Even stem fragments that contain only 1–3 whorls of leaves can pro-
duce new plants. Tubers develop at the ends of rhizomes or stolons and then grow new
plants. One tuber can produce up to 500 new tubers per sq. ft. (5,000 per m2). Tubers
remain viable for several days out of water and for more than four years in sediment.
Turions, produced in the leaf axils, are specialized buds. After detaching from the parent
plant, they are dispersed by water currents to form new colonies. Floating segments produce
more turions than do rooted stems. Both tubers and turions are overwintering organs in cold
climates. Turions begin to grow when temperatures reach 64.5ºF (18ºC). Tubers and
turions can also remain viable under ice cover, dry conditions, and herbicide application.
The only method of dispersal to new water bodies is through human activities, such as
boating and fishing or through the aquarium trade. Some California and Washington popu-
lations may have been transported as hitchhikers on water lily plants.
Impacts. Masses of hydrilla stems just beneath the surface impede navigation and com-
mercial activities and interfere with recreation. Plants clog irrigation canals, flood-control
canals, and intake pumps for irrigation and for hydroelectric plants. Recreational activities,
such as swimming, boating, and sport fishing, are curtailed by hydrilla mats. Stagnant water
in dense mats is prime mosquito-breeding territory. Because plants initially grow from the
bottom, infestations may go unseen until it is too late for efficient control.
Because plants can grow in low light, only 1 percent of what is available, and in silty
water, hydrilla begins growing during the short days of early spring before native plants
begin to grow. Plants grow rapidly, up to 1 in. (2.5 cm) a day. Dense, branching mats block
334 n AQUATIC PLANTS

light, displacing native aquatic plants such as pondweeds, tapegrass, and coon’s tail. They alter
the community structure by changing the water chemistry, raising the pH, and increasing tem-
perature. As the decay of the large biomass depletes oxygen in the water, zooplankton die and
the fish population changes. Hydrilla even outcompetes two other nonnative aquatic pests,
Eurasian watermilfoil (see Aquatic Plants, Eurasian Watermilfoil) and Brazilian waterweed.
Fish, especially chain pickerel, congregate under or near mats, where they use the cover
for hunting.
The economic impact is tremendous. The most abundant aquatic plant in Florida,
hydrilla covered approximately 95,000 ac. (38,500 ha) in 1994. In 1994–1995, Florida
spent about $14.5 million, not including loss of income from curtailed recreational activ-
ities, on controlling hydrilla. From 1980 to 2005, Florida spent $174 million. Millions are
spent each year on herbicides and harvesters just trying to control the plants.
Management. Hydrilla has been successfully eradicated from several sites in California
and Washington. Because hydrilla grows new plants from pieces, physical means of control
such as harvesting or chopping only serve to spread the infestation. Small populations can
be harvested if all parts are removed from the water. Sites should be closely monitored for
regrowth. Mechanical harvesting of large areas can cost $1,200 or more per ac. ($2,950
per ha) and is usually prohibitive because as many as six harvests a year are required due
to the plant’s rapid growth. Mechanical harvesting is used to open boat lanes in waterways
that are totally infested. In regulated areas such as reservoirs, water drawdown may be some-
what effective in reducing stem densities, but it will not kill the plants because of regrowth
from tubers. Covering bottom sediment with opaque fabric that totally blocks light will sup-
press growth around boat docks or swimming areas.
Chemical control is possible with herbicides approved for water use. Fluridone, slow to
take effect, will reduce infestations but will not eliminate hydrilla. Endothall is used when
results are desired more quickly.
The most effective biological control is grass carp (see Volume 1, Vertebrates, Fish, Grass
Carp) (Ctenopharyngodon idella), also called white amur, a nonnative invasive fish that con-
sumes hydrilla. Care must be taken that no fish escape the target water body, because they will
eat all aquatic plants, including natives. Grass carp are prohibited in most states unless the fish
are sterile. Since 1981, over 40 insects that feed on hydrilla have been identified in various
parts of the world. Possibilities include a beetle (Bagous affinis) from India and Pakistan, whose
larvae eat hydrilla tubers. The beetle requires alternating wet and dry periods that may be
simulated by artificial drawdowns. No established populations of the beetle exist in the
United States. Of two leaf-mining flies (Hydrellia pakistanae and H. bulciunasi) released in
Florida, one had no impact, and the other had limited success. An aquatic moth (Parapoynx
diminutalis), accidentally released, defoliates plants, but stems remain viable. Tropical insects
may not do well in northern climates.

Selected References
Evans, Chris, and Joseph LaForest. “Hydrilla verticillata.” Center for Invasive Species and Ecosystem
Health, University of Georgia, 2008. http://wiki.bugwood.org/Hydrilla_verticillata.
“Hydrilla (Hydrilla verticillata).” Non-native Invasive Freshwater Plants, Department of Ecology, State
of Washington, n.d. http://www.ecy.wa.gov/programs/wq/plants/weeds/aqua001.html.
Masterson, J. “Hydrilla verticillata.” Smithsonian Marine Station at Fort Pierce, 2007. http://www.sms
.si.edu/irlSpec/Hydrilla_verticillata.htm.
Ramey, Victor. “Hydrilla verticillata.” Center for Aquatic and Invasive Plants, University of Florida,
2001. http://plants.ifas.ufl.edu/node/183.
 WATER CHESTNUT n 335

Successful Hydrilla Eradication in Washington State

P robably introduced with nonnative water lilies, hydrilla was discovered on

June 1, 1995, in two interconnected lakes, Pipe Lake and Lucerne Lake, in
Washington. These two lakes north of Seattle comprise 73 ac (30 ha). From 1995
through 2000, the entire lake system was treated yearly with an aquatic herbicide.
Although the population of hydrilla thinned, new tubers germinated. No sprouts
developed, however, from seeds or turions. In 2001 and 2002, selectively treating
sections of the lake where hydrilla still existed was not effective. In 2003, herbicide
applications combined with divers and snorkelers hand-pulling plants succeeded in
ridding the lakes of hydrilla. Since 2004, no plants were reported in Lucerne Lake,
and as of 2007, none were found in either lake. Monitoring continues.
Source: Department of Ecology, State of Washington, n.d.

n Water Chestnut
Also known as: Ball nut, European water chestnut, water caltrop, horned water chestnut,
water-chestnut
Scientific name: Trapa natans
Synonyms: Trapa bispinosa, T. natans var. bispinosa, T. natans var. natans
Family: Water Caltrop (Trapaceae)
Native Range. Western Europe, northern Africa, and Asia, including the Indus River
basin, southeastern China, and Vietnam. Possibly southern Africa.
Distribution in the United States. From the northeastern states west to Wisconsin and
south to Virginia. The worst infestations are in Chesapeake Bay and the Potomac River and
in the Connecticut River Valley, Hudson River, and Lake Champlain region. It has the
potential to spread to warm temperate and subtropical regions.
Description. Water chestnut is an aquatic annual with stems that anchor in the muddy
substrate. A system of long, fine roots holds the long stems in the mud. Feathery leaves grow
in whorls around the submerged stems, which can grow 13–16.5 ft. (4–5 m) tall, depending
on water depth. These sessile stem leaves are similar to adventitious root hairs. At the surface,
the stem terminates in a rosette of floating leaves. The glossy rosette leaves, 0.75–1.5 in.
(2–4 cm) long, are triangular and saw-toothed. An inflated spongy petiole, 6 in. (15 cm)
long, gives the rosette buoyancy.
In late June to mid-July, inconspicuous white flowers develop in the center of the leafy
rosettes. Flowers, which have four 0.3 in. (8 mm) long petals, are insect pollinated. Each
flower produces one black fruit that contains one seed. Also called a nut, the fruit is a
woody ball, about 1 in. (2.5 cm) wide and weighing 0.2 oz. (6 g). The four triangular sepals

The Water Chestnut

T he water chestnut used in Chinese cooking is Eleocharis dulcis, in the sedge

family (Cyperaceae). The corms are the edible part.
336 n AQUATIC PLANTS

become hard horn-like prongs

on the fruit. The prongs or
horns, as long as 0.5 in.
(1.25 cm), are covered with
backward pointing barbs. Fruit
develops in the water, just
below the surface leaf rosette.
As annuals, plants die in the
fall and do not overwinter in
the water. Regrowth is from
the nuts.
Related or Similar Species.
No related species of Trapa, or
any other genus of the eastern
hemisphere family Trapaceae,
occur naturally in the United
States. Native to China, Trapa
bicornis, called devil pod, bat
nut, or horn nut, is used for
food in Asia. The glossy black
nut, 2.5–3 in (6.3–7.6 cm) has
two distinct horns and resem-
bles the profile of a bat in flight.
Although not commonly known
and not found in natural settings
in the United States, Trapa bicor-
nis is a noxious weed in the state
of Washington. It is a problem in
the Caspian Sea and in irrigation
systems in India.
Other water plants have
Although present in the United States since the mid-1800s, water some of the same characteris-
chestnut remains localized in the northeastern states. (Native range tics. The introduced waterhya-
adapted from USDA GRIN and selected references. Introduced range cinth, also popular in the
adapted from USGS Nonindigenous Aquatic Species Database and aquarium and water garden
selected references.) trade, can be distinguished by
its tall spike of showy lavender
flowers. Neither Eurasian watermilfoil (see Aquatic Plants, Eurasian Watermilfoil) nor
hydrilla (see Aquatic Plants, Hydrilla) have a floating rosette of leaves.
Introduction History. The specific origin of the genotype in the United States, first seen in
1859 in Massachusetts, is unknown. In 1877, the well-known botanist Asa Gray cultivated it
in his botanical garden in Cambridge, Massachusetts, and by 1879, it had escaped to local water-
ways. By 1884, a population was noticed near Scotia, New York, followed by populations in
Vermont and Massachusetts. By 1923, plants were growing in Maryland, and the species had
spread 40 mi. (65 km) up the Potomac River outside Washington, D.C., covering about
10,000 ac. (4,000 ha). Although the Army Corps of Engineers attempted to remove the infesta-
tion, water chestnut had migrated to the Bird River in Baltimore County by 1955, and to the
Sassafras River, a tributary to Chesapeake Bay, by 1964. Sometime before the late 1950s, water
 WATER CHESTNUT n 337

A. Populations can rapidly increase to completely cover water bodies. B. Small white flowers have four petals and
four sepals. C. Woody fruit have four distinctive “horns.” D. Glossy triangular leaves float in rosettes on the water
surface. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology University of Connecticut, Bugwood.org.)

chestnut was introduced into one of the New York Finger Lakes, part of the larger Great Lakes
Basin, possibly as a result of the connection with the Mohawk River via the Erie Canal. The plant
is now established in parts of Lake Ontario. In 1998, it was found in the South River in Quebec,
which has connections with Lake Champlain. Introductions may have been intentional, as food
for waterfowl, or accidental, from water gardens.
Habitat. Water chestnut is found in shallow, nutrient-rich, fresh-water lakes and rivers,
generally with a pH of 6.7–8.2. Water is usually stagnant or slow moving. Plants can grow
in water as deep as 16.5 ft (5 m), but are usually found where depth is 1–6.6 ft (0.3–2 m).
Reproduction and Dispersal. Water chestnut reproduces both by seed and asexually.
Flowers develop from the center of the rosette on the water surface, but as new leaves are
produced on the meristem, the older leaves and developing fruit sink below the surface.
Fruit ripens in about one month, and the mature fruit falls to the bottom of the water body
where it remains during winter. The prongs or horns on the fruit may serve to help anchor
the seed. Seeds need a dormancy period of about four months and germinate in late spring
on bottom sediments. Although most seeds germinate in the first two years, they are viable
for as long as 12 years. Leaves, which enable the plant to photosynthesize, develop on the
stem while it is growing toward the surface. The stem and leaves reach the surface by mid-
May. After the rosette develops at the surface, the stem branches, growing secondary stems
which also produce rosettes. One rooted stem can create 15–20 rosettes, and each rosette
can produce as many as 20 flowers and 20 seeds. One acre (0.4 ha) of water chestnut can
produce enough seeds to cover 100 ac. (40 ha) the following year.
Asexual dispersal occurs when plants are uprooted or rosettes break free to float with cur-
rents and waves. Nuts are also dispersed by water and may become trapped in feathers of
waterfowl.
Human activity is a major means of dispersal of water chestnut. It is an ornamental in
aquariums and water gardens in the United States and may be improperly disposed of into
natural water bodies.
Impacts. Infestations of water chestnut have both ecological and economic impacts.
Plants grow rapidly, and infestations can increase dramatically. A population of 50 plants
in 1997 on the Bird River in Maryland increased to cover more than three ac. (1.2 ha) in
1998. By 1999, it covered 20 ac. (8 ha).
Floating rosettes of water chestnut can create mats that completely cover the water sur-
face, blocking light to the water below. Density of rosettes on the water surface can be as
many as 5 per sq. ft. (50 per m2), which can cover the surface with leaves three layers deep.
338 n AQUATIC PLANTS

The shade suppresses the growth of other submersed or floating water plants. Because plants
are annuals, much dead material is discarded each year, which changes the water quality.
Reduction of oxygen levels kills fish.
Mats of vegetation interfere with navigation and recreational activities, such as boating,
fishing, and swimming, particularly along shorelines. Bays of southern Lake Champlain,
which were previously used for fishing, are now inaccessible. The spiky seeds are hard
enough to puncture shoe leather and can severely damage bare skin.
Removal is expensive and labor intensive. In 2002, Vermont allotted $500,000 to harvest
and remove water chestnut.
Management. Although control and eradication are difficult, efforts in the 1950s and
1960s by the Army Corps of Engineers succeeded in controlling water chestnut in the
United States. When the program was suspended, however, due to its success and budget
problems, populations rebounded. By 1994, water chestnut infested more territory than it
had before control efforts were begun.
Physical control includes hand removal, pulling and raking plants from boats. Plants are
easily pulled out of the mud, allowing small populations to be removed. Because seeds are
viable for several years, however, eradication is not ensured. Large populations, thick mats
that cover miles of river, require both mechanical harvesting and herbicides. Mechanical
removal and harvesting by volunteers removed approximately 400,000 lb. (181,500 kg) of
water chestnut from the Bird River and Sassafras River in Maryland in 1999. Machinery,
however, cannot operate in the shallow water occupied by much of the water chestnut.
Chemical applications of 2,4-D are somewhat helpful, and the herbicide seems to have no
adverse effect on wildlife.
The search for natural enemies of water chestnut that could be used for biological control
has been conducted in Western Europe and northeastern Asia, where the extremes of the
temperate zone climates are similar to those of regions infested in the United States. A leaf
beetle (Galerucella bimanica), which is widespread in northeastern Asia, can cause complete
defoliation of whole populations. Different stages in its life cycle eat different parts of the
leaves, leaving only a vein skeleton behind. However, the beetle also attacks many other
plants and is a pest of cultivated water chestnut in Asia. A related species, Galerucella nym-
phaeae, which is most common in Europe, does less damage but also feeds on water lilies.
Two weevils, Nanophyte japonica from central Japan and China and an unidentified
Nanophyte species from China and Russia, eat leaves. Although both appear to feed only
on Trapa species, neither produces much damage. Biological agents from India cannot sur-
vive the cooler climate in the northeastern United States, but searches can be conducted in
tropical and subtropical areas of its native range if water chestnut expands its distribution
to the southern United States.

Not Invasive Everywhere

P opulations of water chestnut are dwindling so much in Europe that it has become rare.
The Council of Europe declared it a protected species. In Asia, it has both nutritional
and medicinal value. The nutlike fruit is cultivated and used as food for both humans and
livestock. Parts of the plant are also used to treat elephantiasis, fevers, rheumatism, and skin
disorders.
 WATERHYACINTH n 339

Selected References
Cao, Ling. “Trapa natans.” USGS Nonindigenous Aquatic Species Database, Gainesville, Florida, 2010.
http://nas.er.usgs.gov/queries/FactSheet.aspx?SpeciesID=263.
Pemberton, R. W. “Water Chestnut.” In Biological Control of Invasive Plants in the Eastern United States,
by R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-04, Morgantown, WV,
2002. http://wiki.bugwood.org/Archive:BCIPEUS/Water_Chestnut
“Water Chestnut, Trapa natans.” Invasive Species in the Chesapeake Bay Watershed Workshop, Sea
Grant, MD, 2007. http://www.mdsg.umd.edu/issues/restoration/non-natives/workshop/water
_chestnut.html.

n Waterhyacinth
Also known as: Water-hyacinth, floating water hyacinth
Scientific name: Eichhornia crassipes
Synonyms: Eichhornia speciosa, Piaropus crassipes, P. Mesomelas, Pontederia crassipes,
Heteranthera formosa
Family: Pickerelweed family (Pontederiaceae)
Native Range. Amazon Basin of northern South America, from Venezuela to Brazil.
Distribution in the United States. Widespread from New York state west to Wisconsin,
Texas, and Colorado, and in the Pacific Coast states. A problem primarily in the
southeastern states and in California, Hawai’i, Puerto Rico, and the Virgin Islands.
Description. Waterhyacinth is a floating perennial plant that forms a rosette of leaves
above the surface of the water. While the rosette of leaves extends from a few inches to over
3 ft. (1 m) above the water surface, 16 in. (40 cm) is most common. Leaves, borne upright
on stalks, occur in whorls of 6–10. The round or oval leaves, up to 10 in. (25 cm) wide,
but more likely 6 in. (15 cm) or less, are thick and leathery, waxy or glossy. They are slightly
curved inward toward the stem and sometimes have wavy edges. The stems, which are
actually leaf petioles, are spongy air-filled tissue, bulbous and inflated, which enables the
plant to float. Petioles, usually up to 12 in. (30 cm) long, are short in uncrowded conditions,
but elongate where plants become crowded. Each plant sends out floating stolons in all
directions. The combination of many plants and associated intertwined stolons forms dense,
impenetrable mats that can be 6.5 ft. (2 m) thick.
Approximately 50 percent of the plant biomass is fibrous roots, extending deep into the
water. Many fine lateral roots give the roots a distinctly feathery appearance.
Showy flowers, usually 8–15 on each inflorescence, grow in clusters at the top of a
12–19.5 in. (30–50 cm) stalk which extends above the leaves. The six-petaled flowers,
1.5–2.8 in. (4–7 cm) in diameter, are lavender, purplish blue, or pinkish, with bright yellow
centers. Thin, delicate petals are oval or oblong. The flowers have 6 stamens and 3 stigmas.
The fruit is a thin-walled capsule with three cells. Each capsule usually contains about
50 seeds, but can have as many as 450. Seeds are small, 0.2 by 0.04 in. (5 by 1 mm), with
longitudinal ridges. As the fruit matures, the flower stalk bends to release the seeds under
water.
Related or Similar Species. Other Eichhornia species from South or Central America are
less invasive. Anchored waterhyacinth is listed as a Federal Noxious Weed. Because it must
be rooted, it occurs only in shallow ponds or along the shorelines of lakes and rivers.
Variable leaf waterhyacinth, native to Puerto Rico, is considered difficult to maintain in an
aquarium and is unlikely to be found naturalized. Leaves beneath the water surface are
340 n AQUATIC PLANTS

linear, while those floating on

the surface are round. Brazilian
waterhyacinth, native to Brazil,
is found only locally in Florida.
Introduction History. Water-
hyacinth was introduced as an
ornamental at the Cotton States
Exposition in New Orleans in
1884. Within 70 years, these
fast-growing plants covered
126,000 ac. (5,100 ha) in
Florida. Plants are widely avail-
able in nurseries for fish ponds
and aquariums.
Habitat. Although most
often found floating in water,
waterhyacinth plants can sur-
vive for months on moist sedi-
ment. They have a wide tolerance
to environmental conditions,
including nutrient levels, tem-
perature, and toxic substances.
Optimum water temperatures
are 77–86°F (25–30°C). Tem-
peratures below 54°F (12°C)
and above 92°F (33°C) inhibit
growth. Although its range is
limited by cold, it is able to grow
new leaves after mild freezes. In
northern states, plants will grow
only during the summer and
After introduction to New Orleans as an ornamental plant, waterhyacinth cannot survive cold winters. A
spread throughout most of the warmer states, primarily due to the water hard frost will kill the plant,
garden industry. (Native range adapted from USDA GRIN and selected but reinfestation may occur
references. Introduced range adapted from USGS Nonindigenous from seeds. Waterhyacinth will
Aquatic Species Database and selected references.) grow in water with a pH of
4–10, but it prefers neutral,
nutrient-rich water. It can also survive in water with salinity as high as 0.24 percent, and is tol-
erant of water level changes and variations in water flow. Plants need high light levels to thrive.
Reproduction and Dispersal. Waterhyacinth plants produce seeds, but reproduction and
spread is primarily vegetative. New plants form at nodes along the floating stolons that radi-
ate from the parent plant. Plants grow very fast, and populations can double in six days.
Results of one study showed that only two plants produced 1,200 daughter plants in only
four months. When they break free, these new plants are spread by water currents or pushed
by the wind and redistributed to form new colonies. Plant pieces are frequently carried to
new locations on boats and trailers. Improper disposal of excess or unwanted plants, such
as dumping into a canal or lake, can create additional invasive populations.
 WATERHYACINTH n 341

A. Rapidly growing plants can quickly cover acres or entire lakes. (Graves Lovell, Alabama Department of
Conservation and Natural Resources, Bugwood.org.) B. Spongy, inflated petioles enable plants to float.
(Richard Old, XID Services, Inc., Bugwood.org.) C. Glossy leaves encircle a showy inflorescence. (Wilfredo
Robles, Mississippi State University, Bugwood.org.) D. Roots are thick and feathery. (Rebekah D. Wallace,
Bugwood.org.)

Although plants can flower several times a year, and one plant can produce over 3,000
seeds, expansion by sexual reproduction is limited. Fruit and seeds are rare except in
warmer climates, and few seedlings are seen in the field. Plants are self-fertile but are also
pollinated by bees. The number of fruit produced per plant is highly variable, and under
high temperatures and low relative humidity, few fruit develop. Seeds remain viable in
mud for 15–20 years and are able to survive on both muddy banks and on waterhyacinth
mats. Germination requirements are not clearly determined, but high light intensities and
alternating temperatures favor the process. Buried seeds fail to germinate. Birds may also
transport seeds to new areas, and seeds may also be carried on boats and trailers.
Impacts. Waterhyacinth infestations alter hydrologic systems, impede water traffic,
degrade recreational sites, change water chemistry and quality, decrease biological diversity,
damage wildlife habitat, and increase habitat for undesirable species. Because of rapid
growth, infestations can quickly cover many acres. Mats of waterhyacinth can completely
cover lakes, rivers, canals, and ditches and can be dense enough for a person to walk on.
One acre of healthy plants can weigh 200 or more tons. Plants can block drainage and cause
flooding, impede water flow for irrigation, and clog pumps for hydroelectric power. Mats
may also increase the sediment load by interfering with drainage.
Floating mats can envelop water ways, making them impenetrable by watercraft, both
commercial and recreational. Swimming sites are overrun with plants, and waterfront prop-
erties have suffered a decrease in value because of waterhyacinth. Mats restrict fishing
access, and equipment can become entangled and lost in the root systems. Alterations in
the water environment, such as warmer water temperatures beneath dense mats, change
the species composition and reduce the number of sport fish.
The mats reduce oxygen supply in the water by covering the surface and preventing gas
exchange with the atmosphere. Waterhyacinth reduces biological diversity because it shades
out native water plants, crushes them with the weight of the mat, or literally pushes them
away. With no light, phytoplankton cannot survive, which alters the invertebrate composi-
tion and also the fish diversity. When native aquatic plants die, they further deplete oxygen
supplies as they decay and add organic matter to the water. As a result, fish and other water
342 n AQUATIC PLANTS

fauna suffocate. As well as blocking access to water for land animals, dense mats also elimi-
nate native plants those animals depend on for food, nesting, or shelter. In the Florida
Everglades, for example, the snail kite, a raptor that feeds on apple snails, is negatively
impacted because waterhyacinth has smothered the aquatic food plants for the snails.
Fish-spawning areas and waterfowl habitat are diminished.
The moist mats increase habitat for undesirable mosquitoes and parasitic flatworms and
can provide microhabitat for insects that spread diseases such as schistosomiasis and malaria
in favorable climates. Because evapotranspiration from plants can be 1.8 times that of an
open water surface, the growth of the plants can substantially decrease the water supply.
Control of waterhyacinth, often merely to a maintenance level, costs several millions of
dollars each year.
Management. Preventing the spread of waterhyacinth is a key to its control, because even
one plant can quickly produce hundreds. Garden pond plants must not be disposed of in
native waterways. Boats and equipment used in water must be thoroughly cleaned of all
plant debris before being transferred to other water bodies.
Because of the vast scale of infestation once plants become established, physical methods
of eradication, such as dumping plants onshore to die or chopping them to small bits, are
expensive and have limited effectiveness. Not only is removal a short-term solution, but
the requirements for transportation of plants, both on the water and on land, is enormous
because of the weight of the plants, which are 95 percent water.
Chemical control is also limited, not only because of expense but because some herbi-
cides cannot be used around water because of detrimental effects on water ecosystems.
Some formulations of glyphosate can kill waterhyacinth in eight weeks. Although it is non-
toxic to fish, glyphosate is slightly toxic to aquatic invertebrates. Applications of 2,4-D are
most effective, especially in hot weather. It is selective to broadleaf plants and some mono-
cots, including waterhyacinth, but is somewhat toxic to birds, fish, and aquatic inverte-
brates. Copper sulfate or copper chelate inhibit growth but are toxic to fish, especially
trout, and to some mammals, aquatic invertebrates, and soil organisms.
Biological control may be somewhat accomplished with plant-eating fish or insects.
Although fish eat the roots and do not destroy plants, they may be used to reduce densities
in sport-fishing areas. Possible species include Chinese grass carp (see Volume 1, Vertebrates,
Fish, Grass Carp) (Ctenopharyngodon idella) and two tilapia species (Tilapia melanopleura, T.
mossambica). Chinese grass carp prefers other foods but will feed on waterhyacinth where it is
dense or where little else is available. Because of their diet preferences, they can also reduce
the abundance of native plants. Several states require that any individuals imported be sterile
so they do not outcompete stocked fish or damage native aquatic vegetation.
Other biological control, however, is promising, and over 100 species of insects have
been considered. Two weevils (Neochetina bruchi and N. eichhorniae), native to Argentina,
control the plant in some locales, but success has been inconsistent. The weevils place eggs
inside plant tissue, where larvae then feed and weaken the plant. Because they also feed on
native pickerelweed, however, they are not suitable for use in the United States. Similarly,
the waterhyacinth moth (Sameodes = Niphograpta albiguttalis) and a mite (Orthogalumna tere-
brantis) may also damage native plants. Biological control is slow relative to the rapid growth
and spread of the plants. New research involves species of Thrypticus flies and plant-hopper
species of Taosa and Megamelus, all native to the source regions of waterhyacinth. The fly lar-
vae feed on leaf stalk tissue, while the plant-hoppers are sap-sucking insects that insert
pathogens into the plants, weakening them. Two possible fungus species, Acremonium
 WATERHYACINTH n 343

Is Waterhyacinth Good for Anything?

S everal projects involving practical uses of waterhyacinth, some which are more
applicable to developing countries, are being conducted in various parts of
the world. The fibrous tissue, when blended with other materials, makes a suitable
paper or fiber board. Dried plants are woven into baskets and matting, and rope
made from the fibers is used to wrap around canes to make a type of durable rat-
tan furniture. Because the roots harbor aerobic bacteria that feed on waste, plants
may be used to filter sewage. Plants or their roots provide nutritious food for pigs,
ducks, and pond fish such as carp, tilapia, or catfish. Other ideas include molding
charcoal briquettes from burned remains, development of biogas, or use as fertiliz-
ers, mulch, or compost.

zonatum, associated with the waterhyacinth mite, and Cercospora rodmanii, native to the
southeastern United States, currently provide low levels of control.

Selected References
Batcher, Michael S. “Element Stewardship Abstract, Eichhornia crassipes.” Global Invasive Species
Team. The Nature Conservancy, 2000; updated 2009. http://wiki.bugwood.org/Eichhornia
_crassipes.
Center, T. D., H. Cordo, M. P. Hill, and M. H. Julien. “Waterhyacinth.” In Biological Control of Invasive
Plants in the Eastern United States, by R. Van Driesche et al. USDA Forest Service Publication
FHTET-2002-04, Morgantown, WV, 2002. http://www.invasive.org/eastern/biocontrol/4
WaterHyacinth.html.
“Water Hyacinth.” Practical Answers, Technical Information Online, n.d. http://practicalaction.org/
practicalanswers/product_info.php?products_id=189.
“Water Hyacinth, Eichhornia crassipes.” Center for Aquatic and Invasive Plants. University of Florida,
IFAS, n.d. http://plants.ifas.ufl.edu/node/141.
n Forbs
n Canada Thistle
Also known as: Californian thistle, creeping thistle, field thistle, corn thistle
Scientific name: Cirsium arvense
Synonyms: Cirsium setosum, Breea arvensis, B. incana, Carduus arvensis, and others.
Family: Sunflower (Asteraceae)
Native Range. Southwestern Asia, including Afghanistan, Iran, Turkey, and the Caucasus
region. Widespread, and possibly native to Europe and more extensive areas in Asia.
Distribution in the United States. Most of the country except the southern states and
Hawai’i. Especially invasive in the northern states, from northern California east to Maine
and south to Virginia. The southern limit may be influenced by high summer temperatures
and shorter summer days.
Description. Canada thistle is an herbaceous perennial with erect stems growing 1.5–4 ft.
(0.5–1.2 m) tall, branching toward the top. It has four interfertile varieties, the most
common being Cirsium arvense var. horridum and numerous genotypes, which account for
its variable appearance. New shoots and seedlings initially form a rosette, although it may
be poorly developed with few leaves. Mature plants may have several slender stems arising
from the root system. Stem leaves are sessile and alternate. All leaves are oblong to lance-
shaped, 2–8 in. (5–20 cm) long, with either entire or shallowly lobed and toothed, spiny
margins. The undersides of leaves are often covered with soft, wooly hairs, but the upper
surface is green and glabrous. Unlike many other thistles, leaf bases do not extend down
the stem to form wings, and the stem is almost glabrous.
The plant has a deep, extensive network of both horizontal and vertical roots. Seedlings
first grow a fibrous taproot that serves as a storage organ, before developing creeping roots
in 2–4 months. Although primarily in the top 18–24 in. (45–60 cm) of soil, the vertical roots
may extend down 6.5–10 ft. (2-3 m), and sometimes as deep as 23 ft. (7 m). The horizontal
roots can spread outward several feet, and although brittle and easily fragmented, can sur-
vive in frozen soil.
Canada thistle is dioecious, meaning that male and female flowers are on different plants.
Although a patch may be all one sex, both male and female plants can develop from the same
clone. The flowers, which bloom from June to October, are rose-purple, lavender, or some-
times white. Cylindrical, oval, or bell-shaped flower heads grow in umbrella-shaped clus-
ters. Female flowers may have a distinct vanilla-like smell. Flower heads are 0.4–0.75 in.
(1–2 cm) in diameter, and the spines are short and slender. The involucre, or whorl of bracts
beneath the flower head, is purplish and either glabrous or with white wooly hairs. Old, dry
flower stems remain standing erect.
Fruits are small and dry brownish or tan achenes, with a tan, feathery appendage called a
pappus. The pappus in Cirsium species has branched hairs, giving the genus the general name
of “plumed thistles.” Male flowers have a slightly shorter pappus without a seed, 0.4–0.6 in.
(1–1.4 cm), compared with the pappus on female flowers, 0.6–0.8 in. (1.5–2 cm). The pappus
on both types of flowers is deciduous, leaving most seeds in the flower head.
 CANADA THISTLE n 345

Related or Similar Species.

Several Cirsium species are
native to the United States.
Yellowspine thistle, native to
the southwestern United States
is a clump-forming perennial
similar in size to Canada this-
tle but with narrower lance-
shaped leaves, 4–10 in. (10–
25 cm) long and less than 1.2
in. (3 cm) wide. Its stems are
not winged and are densely
wooly. Leaf margins are deeply
and coarsely lobed and toothed.
Lower leaf surfaces are densely
wooly, while the upper surfaces
are grayish and only lightly
covered with white wooly
hairs. White, pink, or pale pur-
ple flower heads, 0.8–1.6 in.
(2–4 cm) long, have a sparse
cover of white wooly hairs. The
stout, yellow spines, 0.1–0.5
in. (5–12 mm) long, are spread-
ing or curved slightly down-
ward. Seeds are pale orangish
brown with tan pappus bristles,
1–1.2 in. (2.5–3 cm) long.
Wavyleaf thistle, native to
the western third of the United
States and spotty in eastern
states, is also known as gray Canada thistle grows best in northern climates with long summer days.
thistle and hybridizes with yel- (Native range approximated from USDA GRIN and selected references.
lowspine thistle. Although it Introduced range adapted from USDA PLANTS Database, Invasive Plant
grows in compact clumps, it is Atlas of the United States, and selected references.)
not a widely spreading species
because its lateral roots are short and only weakly creeping. The shallow and coarsely lobed
leaves resemble those of yellowspine thistle, but rosette leaves are longer, 6–12 in. (15–
30 cm), and more than 1.2 in. (3 cm) wide. Stems are densely wooly and not winged.
Upper leaf surfaces are lightly wooly. Flower heads resemble those of yellowspine thistle,
but have short, slender spines. Its seeds are tan.
Several thistles native to Europe and North Africa are pests that resemble Canada thistle.
The pappus of Carduus species is not branched or feathery. The taxonomy of musk thistle is
not clear (see Forbs, Musk Thistle). It may be several species or several subspecies. This
biennial species is common in southern California, the Midwest, and Appalachia.
Bull thistle is a biennial found throughout the United States, growing best in heavy, fertile
soils. Reaching 6.5 ft. (2 m) tall, it has foliage with stiff hairs and prickly, winged stems
caused by the leaf bases extending down the stem. Stems and undersides of leaves are
346 n FORBS

A. Stems may branch near the top. (Steve Dewey, Utah State University, Bugwood.org.) B. Lobed and toothed
rosette leaves have spiny margins. (Ohio State Weed Lab Archive, The Ohio State University, Bugwood.org.)
C. Flowers are usually purple or lavender, and white flowers (right) are rare. (Steve Dewey, Utah State University,
Bugwood.org.) D. The involucre is the whorl of bracts beneath the flowerhead. (Chris Evans, River to River CWMA,
Bugwood.org.) E. Seeds of the female flower heads have a long pappus. (Richard Old, XID Services, Bugwood.org.)

covered with a network of white hairs that resemble cobwebs. The upper leaf surface is
bumpy and covered with long, stiff hairs. Bull thistle has a thick, fleshy taproot, sometimes
branched, extending 28 in. (70 cm) deep.
Common in coastal California, Italian thistle is rare elsewhere in the United States. It is a
summer annual with seeds that generally germinate the following spring. Milk thistle is a
pest in the dry coastal regions of California. As a winter annual, it germinates in late summer
or fall, producing flowers and seeds the following spring or summer.
Introduction History. Canada thistle was inadvertently brought to the United States in
the early 1600s. It was declared a noxious weed in Vermont by 1795 and in an additional
43 states by 1954.
Habitat. Canada thistle commonly invades disturbed sites, even the modest soil erosion
caused by gopher mounds, and is rare in healthy pastures. It is common on altered sites,
such as road and railway right-of-ways, pastures, agricultural fields, and abandoned fields.
It is frequently found in moist upland environments, such as prairies and grasslands in the
Midwest and Great Plains, and in riparian habitats in the Intermountain West. In the eastern
states, it grows in drier habitats, such as sand dunes and sandy fields. Because plants do not
tolerate shade, Canada thistle grows only in non-forested sites. It grows in a variety of soil
textures, including clay, loam, silt, gravel, and chalk, but prefers deep, well-drained mesic
soils and will not tolerate waterlogged soils. The plant can also tolerate 2 percent salt.
Reproduction and Dispersal. Canada thistle reproduces both sexually and vegetatively.
Because it is a long-day plant, meaning that at least 14 hours of daylight are required for
flowering, plants have more flowers and a longer bloom period at higher latitudes. In order
to produce seed, female plants and male plants must be within 0.25 mi. (0.4 km) of each
other. Some predominantly male plants, however, are self-fertile, with both male and female
flowers. Seed develops 8–10 days after insect pollination. One plant may produce 5,000
seeds each year, and a dense thicket covering 10 sq. ft. (1 m2) can produce over 70,000 seeds.
 CANADA THISTLE n 347

Although the bristly plumed seeds may be wind-dispersed more than 0.6 mi. (1 km), the pap-
pus breaks off easily, and most seeds fall near the parent plant. Seeds may also be dispersed by
water, in animal droppings, by birds and small mammals, and by human activity, such as on
vehicles and equipment or as a contaminant in packing material, soil, crop seed, or hay.
Most seeds germinate within one year, but unless deeply buried, some remain viable in the soil
for 20 or more years. Germination is best in the top 0.4 in. (1 cm) of soil, at 68–86ºF (20–
30ºC) with abundant moisture, bright light, and little competition. Canada thistle does not
easily establish from seed where soil is undisturbed, and seeds have low viability after passing
through a digestive tract. Germination and rosette growth take place in mid to late spring, and
seedlings rapidly develop a root system. Roots may initially grow 0.4 in. (1 cm) per day, and
the plant grows a vertical root 5 ft. (1.5 m) deep in the first year. Plants flower the following
summer. Most germinating seeds become female plants.
Local expansion is by roots and shoots. Canada thistle forms clumps or patches because
of its creeping root system. Although most horizontal roots grow 3.3–6.6 ft. (1–2 m) a year,
they can grow as much as 19.5 ft. (6 m) in one season, radiating away from the parent plant.
The shoots develop in winter and emerge in spring, from as much as 3 ft. (1 m) beneath the
surface. Wherever a vertical shoot grows, approximately every 2–3 ft. (0.6–0.9 m), another
horizontal root and a vertical root develop in the same location. Because sprouts develop
quickly as the horizontal root grows, the lateral roots do not extend very far beyond the
perimeter of the thicket. Although growth and density of shoots varies with environmental
conditions, 3.3 ft. (1 m) of horizontal root may have 8–24 buds. Root buds, however, are
not necessary. New shoots may sprout from root fragments less than 1 in. (2.5 cm) long
and create a clone 9 ft. (2.8 m) in diameter in one year. Most roots live only two years, but
the plants grow new sections every year. Stem pieces also grow new plants.
Whether from seed or from roots or stems, plants grow quickly after a rosette stage lasting
2–4 weeks. When plants bolt, the flowering stems may grow 1.2 in. (3 cm) per day, and
flower after 10 weeks. Because root buds produce new shoots all summer, a stand may be
in several stages of growth.
Impacts. Because of its creeping root system, Canada thistle readily establishes and
crowds out native plant species. Clones can reach 115 ft. (35 m) in diameter. It changes
the structure of the community, displaces native species, and reduces plant and animal
diversity. Plants can produce as many as 29 flowering shoots in 10 sq. ft. (1 m2), shading
out other plants and directly competing for water and soil nutrients. Canada thistle may also
have alleleopathic qualities. Insect pollinators may prefer Canada thistle flowers, to the detri-
ment of native plant species.
Although plants may accumulate nitrates that are toxic to animals, the substance causes
few problems because the plants are not grazed by livestock. Spiny leaves scratch or punc-
ture livestock hides, resulting in infections.
Canada thistle costs tens of millions of dollars in crop loss annually, and millions for control.
Shoot densities of fewer than 2 per 10 sq. ft. (1 m2) may cause a 15 percent loss in forage. A
density of 20 shoots per 10 sq. ft. (1 m2) can cause a yield loss of 35–50 percent in crops, such
as barley, winter wheat, or alfalfa. It is a host for the bean aphid and bean stalk borer, which
affect corn and tomatoes, and is a host for the sod-web worm, which also affects corn crops.
Density of standing stalks and litter can increase both fire frequency and intensity.
Management. Eradication is difficult once Canada thistle is established, and plants
should be removed when first spotted. A combination of methods is most effective.
Different ecotypes, however, respond differently to management methods, and results are
highly variable. Regardless of the method used, priority should be given to killing the entire
348 n FORBS

clone, including the root system and any root pieces. Although spread by seed is relatively
rare, an additional goal should also be decreased seed production and destruction of the
seed bank. The best control is to prevent infestation by maintaining good pastures, but
treated areas may be re-infested from areas not cared for, such as roadsides.
Removal of Canada thistle by physical means is difficult. Occasional cultivation or mow-
ing are poor practices because they stimulate growth of horizontal roots, and cultivation
breaks roots, which sprout new plants from pieces. Repeated cultivation or mowing, how-
ever, especially when done in early spring before much growth has taken place, can deplete
root reserves. Cultivating 3–4 in. (7.5–10 cm) deep every 20 days can eliminate up to 90 per-
cent of plants. Mowing every 7–21 days for four years may eliminate the plant. Cultivation
and mowing, however, are not appropriate in natural areas. Prescribed burns done during
the dormant season may stimulate native species to grow, which both decreases the amount
of bare soil and shades out Canada thistle. Fire during the growing season, however, dam-
ages native plants. Grazing is ineffective because livestock do not eat Canada thistle and
avoid areas where it grows, giving it a competitive advantage. Because Canada thistle is not
competitive in shade, planting crops such as alfalfa may prohibit its early establishment.
Chemical applications are most successful when physical means are used to stress the
plant early in the season. They may not be appropriate, however, for natural areas where
herbicides can also kill native plants, alter succession, and open the site for more invasion.
Depending on the site and growth stage of Canada thistle, herbicide possibilities include
glyphosate, clopyralid, and chlorsulfuron. Picloram, dicamba, metsulfuron, and 2,4-D are
not recommended, either because they are ineffective or are too damaging to native species.
Different clones have varying tolerances to herbicides, and Canada thistle’s deep root system
makes plants resistant. Applications are most successful on young plants or plants weakened
by mowing or tilling and should be done before flowering to prevent seed production. Care
should be taken to treat all areas because plants do not always remain connected as roots die
or are severed. Chemical applications are most effective when combined with physical or
biological controls, such as the European seed weevil.
Biological control is not promising. Canada thistle is a major crop pest in its native
European range where no major enemies keep it under control. Potential insects must be
host-specific to avoid adverse impacts on native Cirsium species or crop plants, such as saf-
flower and globe artichoke, which are closely related to Canada thistle. Most insects weaken
and kill individual plants, but do not offer widespread eradication, and no one agent is
totally effective. Larvae of the native painted butterfly (Vanessa cardui) defoliate local areas,
but their populations vary yearly with migrations. This butterfly occurs primarily in the
southern states and also affects other Cirsium species.
Several insects have been introduced, either accidentally or intentionally, and may be
established in different parts of the country. An Asian beetle (Altica carduorum), released in
1966, eats leaves during summer, defoliating and weakening plants but failing to kill them.
Stressed plants, however, produce fewer shoots and seeds. This beetle has not become estab-
lished due to predation. The Canada thistle stem weevil (Ceuthorhynchus litura), intentionally
released in 1971–1985, eats stems, but plants recover with new shoots. Although it does not
attack crops, it feeds on all Cirsium species, including natives. A bud weevil (Larinus planus)
eats seed heads but has little effect and also attacks native thistle species. A thistle stem gall
fly (Erophora cardui), released 1981–1985, feeds primarily on Canada thistle shoots, and a
seed-head fly (Terellia ruficauda) destroys seeds. Larvae of a European seed weevil
(Rhinocyllus conicus) prefer to eat seed heads of musk thistles but will also feed on Canada
thistle and bull thistle. A leaf-feeding tortoise beetle (Cassida rubiginosa), accidentally
 CHINESE LESPEDEZA n 349

Uses of Canada Thistle

N ative Americans adopted Canada thistle and used it for medicinal purposes to
cure mouth diseases. The Chippewa used it as an astringent, tonic, and
diuretic. Young shoots and roots may be eaten like asparagus, and the flower’s
nectar makes good honey.

introduced in 1902 and established since 1927, defoliates both Canada thistles and bull
thistles, but also feeds on artichokes. A root-feeding weevil (Cleonis pigra) and a lace bug
(Tingis ampliata) attack artichokes.
Two fungal pathogens damage Canada thistle. A widespread rust fungus (Puccinia puncti-
formis) attacks leaves of the basal rosette and new shoots, causing them to die before flower-
ing. Although introduced into Canada, its success is not yet known. Another fungus
(Sclerotinia sclerotiorum) affects the roots.

Selected References
“Canada Thistle.” Non-Native Plant Species of Alaska. Alaska Natural Heritage Program, Environment
and Natural Resources Institute, University of Alaska, Anchorage, 2006. http://akweeds.uaa
.alaska.edu/pdfs/species_bios_pdfs/Species_bios_CIAR_ed.pdf.
“Cirsium Genus.” Plant Pest and Health Prevention Services (PHHPS), California Department of Food
and Agriculture, 2010. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/winfo_table-commname.htm.
“Integrated Pest Management Manual, Thistles.” Explore Nature. National Park Service, n.d. http://
www.nature.nps.gov/biology/ipm/manual/thistle.cfm.
McClay, A. S. “Canada Thistle.” In Biological Control of Invasive Plants in the Eastern United States, by
R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-04, Morgantown, WV,
2002. http://www.invasive.org/eastern/biocontrol/17CanadaThistle.html.
Nuzzo, Victoria. “Element Stewardship Abstract, Cirsium arvense.” Global Invasive Species Team,
Nature Conservancy, 1997. http://wiki.bugwood.org/Cirsium_arvense.

n Chinese Lespedeza
Also known as: Chinese bush clover, sericea lespedeza, sericea bush clover,
silky bush clover, Himalayan bush clover, hairy lespedza
Scientific name: Lespedeza cuneata
Synonyms: Lespedeza juncea var. sericea, Lespedeza sericea
Family: Pea (Fabaceae)
Native Range. Eastern Asia, including eastern Russia, Mongolia, northern China, Korea,
Taiwan, and Japan.
Distribution in the United States. The Great Plains east to the Atlantic and Gulf coasts,
from Minnesota to Texas, New York to Florida.
Description. Chinese lespedeza is a warm-season perennial herb, with one to several stems
that usually grow 3–5 ft. (1–1.5 m) tall, giving it a shrubby appearance. In deep loamy soils, it
can reach 6.5 ft. (2 m) in height. Young plants have one stem. More stems develop from the
base of older plants, and the stems branch as they grow upward. Stems angle outward, giving
the plant a V shape. The round stems have defined ridges covered with bristly white hairs.
350 n FORBS

Young stems are light green.

Older stems turn brown as they
become somewhat woody and
fibrous, and may lose the hairs.
The compound leaves, alternate
on the stem and with very short
petioles, are trifoliate, meaning
that each leaf has three leaflets,
like clover. Each narrow leaflet
is 0.2–1 in. (0.5–2.5 cm) long,
with smooth margins. Although
the tip of each leaflet is rounded,
it ends in an awl-shaped sharp
point. It is the only species in
the Lespedeza genus with a
wedge-shaped leaf base. Small
flattened hairs, which cover the
lower surface of the leaflets, give
the leaves a grayish-green to sil-
very color.
Plants have a woody taproot
with lateral branches. The
taproot may extend 3–4 ft.
(1–1.2 m) deep into the soil.
From late July to October,
two types of flowers grow from
leaf axils of the upper or middle
leaves of the plant. Chasmoga-
mous flowers, meaning that they
open to be cross-pollinated by
honeybees and other insects,
Several cultivars of Chinese lespedeza were developed and used for appear first, in clusters of 1–4.
erosion control in the midwestern and eastern states. (Native range They are typically pea-shaped
approximated from USDA GRIN and selected references. Introduced and small, 0.3 in. (0.75 cm),
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the creamy white to pale yellow,
United States, and selected references.)
with purplish throats. The
uppermost petal, the banner, is
streaked with pink- to purple-colored veins. Cleistogamous flowers, which do not open and
are self-fertilized, appear second. They do not have showy petals, and are scattered among
the chasmogamous flowers. The numbers of each type of flower, as well as the amount of seed
produced, depends on the number of daylight hours, sun or shade location, and air temper-
atures. More cleistogamous flowers are produced when plants are mowed or grazed.
One small shiny seed, which can be tan, olive, purple, or mottled brown, is produced
from each flower. The shape and size of the seeds, ellipsoid to oval and slightly flattened,
varies according to the type of flower that produced it.
Related or Similar Species. The closely related native slender lespedeza, found in a sim-
ilar range of states, has fewer stems and is a shorter plant, typically growing 2.5 ft. (0.75 m)
tall. The stems are covered with fine white hairs. The trifoliate compound leaves are
 CHINESE LESPEDEZA n 351

A. Because plants have several stems, they resemble shrubs. B. The three leaflets are slightly hairy on the lower
surface (left). C. Chasmogamous flowers are typically pea-shaped. D. Each flower produces one oval seed.
(James H. Miller, USDA Forest Service, Bugwood.org.)

supported on 0.5–1 in. (1.3–2.5 cm) petioles, and the glabrous leaflets are as long as 2 in.
(5 cm). The flowers are pink, and the base of the banner petal is a deeper rose color.
Introduction History. Chinese lespedeza was deliberately introduced into the southern
states in the late 1800s for its useful characteristics, including bank stabilization and wildlife
cover. The plant was widely recommended in the 1940s by federal and state agencies for
erosion control and as a forage and hay crop capable of growing on poor soils. Several culti-
vars, such as ‘Common Sericea,’ ‘Arlington,’ ‘Serala,’ and ‘Interstate,’ were specifically devel-
oped for the midwestern and eastern United States. The plant’s stems, however, are only
tender and succulent until they reach 12–15 in. (30–38 cm) tall. Older stems are too woody
and coarse to be palatable.
Chinese lespedeza seeds, including the cultivars, continue to be available for sale via the
Internet. The plant is still used for erosion control along roadsides, around reservoirs, and
on strip-mine spoils. It is also used for forage and is still planted as cover for wildlife in some
parts of the United States.
Habitat. Chinese lespedeza grows in a variety of habitats, including sites where the soil has
become severely eroded and sterile and in rough or rocky locations. Because it has been delib-
erately seeded and cultivated for pasture, it is frequently found in grassy environments, such as
meadows, prairies, open woodlands, and savannas. It is also a weed in cultivated crops and fal-
low fields. It is common in disturbed sites, such as roadsides, railroad tracks, fence rows, trails,
burned sites, and any other open disturbed ground that is moist. Plants thrive on bottomland
sites, including ditches and wetland borders of ponds and swamps. It can survive flooding in
cool water for as long as 10 days, but not in warm water. The plant tolerates light to moderate
shade but will not grow in heavy shade. Plants will grow in shallow soil but do best in deep
soil, such as organic rich sands, sandy loams, or clay loam. Soils can be either neutral or very
acid. Although it is drought tolerant, Chinese lespedeza grows best with annual precipitation
of 30–35 in. (75–90 cm), and prefers dry winters and wet summers. Plants survive freezing
winter temperatures when dormant, but late spring frosts will damage young growth.
Reproduction and Dispersal. Chinese lespedeza reproduces primarily by seed but also veg-
etatively. Seeds, with no special structure for dispersal, are dispersed when animals and birds
eat the fruit and distribute the seeds. Seeds are often contaminants in hay, which is often trans-
ported long distances to new locales. Seeds remain viable for 20 years or longer, and the seed
bank is extensive. Seeds from chasmogamous, or cross-pollinated, flowers need to be scarified
before germination. Optimum temperatures for germination are 68–86ºF (20–30ºC), with a
352 n FORBS

variation between day and night. Germination is best when day temperatures are 79ºF (26ºC),
night temperatures are 72ºF (22ºC), and days have 13–15 hours of daylight.
Although old stems die back to the ground in winter, they may remain standing. New
stems sprout every spring from buds on the root crown. A plant can produce 5–30 stems
after four years of growth. Root crown buds will be stimulated to sprout after mowing, burn-
ing, or grazing. Stems can also grow from lateral buds after mowing or grazing.
Impacts. Chinese lespedeza easily escapes cultivation and is an invasive species for many
of the same reasons that it was introduced to the United States. It can grow in poor habits of
eroded, infertile soils, is resistant to prolonged drought, and is not susceptible to insects or
disease. As a long-lived perennial, it provides abundant forage and abundant seed.
Beef cattle and goats will eat the new growth, the pliable stems with low tannin content, but
they are not appropriate for dairy cattle or hogs. Tannins, which inhibit the growth of other
plants, render Chinese lespedeza unpalatable and untouched unless nothing else is available.
Tannins are more concentrated in the leaves than in the stems, and also in the upper portions
of the plants and in older plants. Other than early in the season when the shoots are tender, it
is not important food for wildlife. The plantings at field edges next to wooded areas, however,
provide nesting sites for Bobwhite Quail, Grasshopper Sparrow, Meadowlark, and Greater
Prairie Chicken. Bobwhite Quail, Wild Turkey, and other gamebirds eat the seeds.
Chinese lespedeza is an aggressive plant that invades open areas such as meadows and
grasslands in the midwestern and eastern United States. The plants crowd out and suppress
native flora, and the tannins have alleleopathic compounds that inhibit growth of nearby
native plants. Chinese lespedeza can dominate a grassland community in 3–4 years.
Mature plants begin growth earlier in the spring than do most native species. With its large
taproot, the species outcompetes native species for water and nutrients, especially in times of
drought. Their competitive ability, however, decreases when plants are continually grazed.
With a greater number of stems each year, one plant can form a large monospecific stand
that can live for more than 20 years.
Management. The potential of restoring large areas to a natural state after invasion by
Chinese lespedeza is slim, and efforts may take several years. Small infestations have a better
chance of being eradicated, although they are difficult to identify and may quickly grow into
large stands. Chinese lespedeza is difficult to control in natural areas because digging and
mowing cause too much damage to the environment. An integrated management program,
which includes burning, mowing, and herbicides, is best. Any infestations are difficult
because roadways and erosion-control sites supply a steady seed source. The best control
would be to eliminate plant and seed sales.
Physical control alone is limited. The depth of the extensive root system makes digging or
pulling out plants difficult. Mowing stands low to the ground for 2–3 consecutive years
when they are in the flower bud stage will reduce the number of seeds produced and also
decrease the vigor of plants. Burning may increase the infestation because it both scarifies
seed in the seed bank and creates bare ground for germination. Spring burns stimulate
new shoots from the root crown. Late-summer burns, however, will decrease seed produc-
tion from the current crop. Mowing, which decreases seed production, followed by herbi-
cides on regrowth may be effective.
Postemergent chemical treatments can kill mature plants, even the taproot, but they do
not affect the seedbank. Herbicides are effective when applied in early to mid-summer, but
spraying may need to be repeated for several years. Triclopyr and clopyralid afford the best
control but cannot be used in wet sites. Glyphosate is safer for wetlands. Herbicides should
be applied while the roots are still increasing reserves, before the plants flower, to be sure
 COMMON MULLEIN n 353

that the chemicals are carried to the root. One effective integrated method is to spray herbi-
cide on newly sprouting vegetation, burn the remainder, and reseed the area with annual
cereal grains or warm season grasses. Chinese lespedeza is not susceptible to 2,4-D,
picloram, or dicamba.
Biological control is limited. Three-cornered alfalfa leaf hopper (Spissistulus festinus), grass
army worms (Pseudodaleta unipuncta), grasshoppers (Schistocerca americana), and lespedeza
webworm (Tetralopha scortealis) need further research.

Selected References
Hilty, John. “Silky Bush Clover.” Illinois Wildflowers, 2002–2010. http://www.illinoiswildflowers.info/
weeds/plants/silky_bushclover.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Lespedeza cuneata.” ISSG Global Invasive Species Database. 2005. http://www.issg.org/
database/species/ecology.asp?si=270.
Remaley, Tom. “Chinese Lespedeza.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2006. http://www.nps.gov/plants/alien/fact/
lecu1.htm.
Stevens, Sandy. “Element Stewardship Abstact, Lespedeza cuneata.” Global Invasive Species Team,
Nature Conservancy, 2002. http://wiki.bugwood.org/Lespedeza_cuneta.

n Common Mullein
Also known as: Big taper, flannel mullein, velvet dock, wooly mullein, Jacob’s staff,
flannel leaf, and others
Scientific name: Verbascum thapsus
Synonyms: None
Family: Figwort (Schrophulariaceae)
Native Range. Europe and western Asia, from Russia and Afghanistan east through India
to southern China.
Distribution in the United States. Every state, including Alaska and Hawai’i.
Description. Common mullein is an erect biennial herb that grows as tall as 5–10 ft.
(1.5–3 m), including the flower stalk. In its first year, the plant grows a rosette of leaves as much
as 24 in. (60 cm) in diameter. Rosette leaves are 3–20 in. (8–50 cm) long and 1.5 in. (4 cm)
wide. Growth of the rosette continues until stopped by low temperatures in autumn. In the sec-
ond season, the plant bolts, sending up a strong flowering stem, 1.5–6 ft. (0.5–2 m) tall, which
is densely wooly with branched hairs. The stem is longitudinally ridged and covered by over-
lapping leaves. Stem leaves, oblong to lance-shaped, 4–16 in. (10–40 cm) long and usually
2–4 times longer than they are wide, are alternate or in a loose spiral around the stalk. Leaves
are largest at the bottom of the stalk, decreasing with height. Both rosette and stem leaves are
bluish gray-green, with a felt-like pubescence which makes then densely wooly and soft.
Plants grow both a deep taproot and fibrous lateral roots in the first year. Depth and size
of roots vary with location and environmental conditions. Root growth almost ceases when
the plant puts its energy into producing the flower stalk.
Single flowers, sessile or with very short pedicels, are densely packed on a terminal spike
at the top of the leafy stalk. Plants normally have one inflorescence, but it may branch into
two or three, especially if damaged. Beginning in June and usually continuing to August or
even October, flowers bloom and produce fruit progressively from the base of the inflores-
cence upward. Flowering period depends on the height of the inflorescence, which can be
354 n FORBS

8–20 in. (20–50 cm) long. The

stalk will continue to grow
and flower until stopped by
cold temperatures in autumn.
Flowers are 0.75–1.5 in. (2–
3.8 cm) in diameter and bright
sulfur yellow, rarely white, with
five round petals that are fused
at the base. Stalks, often with
fruit attached, remain standing
throughout the winter. Fruit
are small, 0.25 in. (6 mm), oval
two-celled capsules covered
with short hairs. They split
open when mature to release
many tiny brown seeds, which
are rough with wavy ridges and
deep grooves. Common mullein
is monocarpic, meaning that it
dies after flowering and matur-
ing fruit.
Related or Similar Species.
Moth mullen, also called spuri-
ous mullein or slippery mullein,
is a close relative of common
mullein. Also native to Eurasia,
it was introduced to the East
Coast and was first recorded in
Pennsylvania in 1818 and in
Michigan in 1840. It can be dis-
tinguished from common mul-
Deliberately brought to the United States in the mid-1700s, common lein by its smaller size, leaf and
mullein has a long-standng presence in every state, including Alaska flower details, and lack of hairs.
and Hawai’i. (Native range approximated from USDA GRIN and selected The basal rosette is usually 8–12
references. Introduced range adapted from USDA PLANTS Database, in. (20–30 cm) in diameter, and
Invasive Plant Atlas of the United States, and selected references.) the flowering stalk is 2–4 ft.
(0.6–1.2 m) tall. The largest
leaves are 6 in. (15 cm) long by 2.5 in. (6 cm) wide. Leaf margins are toothed and sometimes
slightly wavy or irregular. Leaves are hairless, and veins on the upper surface appear wrinkled.
The inflorescence is 0.5–2 ft. (0.2–0.6 m) long. Showy flowers, ranging in color from yellow
to pinkish white, with a purplish tinge in the center, are less densely packed on the inflorescence
than on common mullein. Flowers are not sessile but are borne singly on 0.5 in. (1.25 cm) ped-
icels. Flower centers have fine purple and white hairs on the stamens.
Introduction History. Common mullein was deliberately brought to North America in the
mid-1700s by settlers as a fish poison. Crushed seeds placed in the water prevent fish from
breathing, allowing easy capture. The plant was first recorded in the Blue Ridge Mountains
of Virginia in the 1700s. Because it was used as a fish poison in Europe and also as a medici-
nal herb, common mullein was probably introduced multiple times. The species has been in
 COMMON MULLEIN n 355

A. In the first season, plants grow as a rosette. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.)
B. Fuzzy leaves feel like velvet. (Bonnie Million, BLM, Ely District, Bugwood.org.) C. In the second season,
plants bolt and flower. (Karan A. Rawlins, University of Georgia, Bugwood.org.) D. Single flowers are sessile
on the stalk. (Steve Dewey, Utah State University, Bugwood.org.) E. Dried fruit capsules split open to release
seeds. (Bonnie Million, BLM, Ely District, Bugwood.org.)

the United States so long that it was described as native in an 1818 flora of the East Coast. It
spread quickly and was recorded in Michigan by 1839 and was a common weed in Boulder,
Colorado, in 1905. The species reached the Pacific coast and became naturalized by 1876. It
was first reported in Hawai’i in 1932 and recorded in Alaskan floras by 1968.
Habitat. As evidenced by its distribution in both Alaska and Hawai’i as well as through-
out the contiguous United States, common mullein has a wide range of environmental toler-
ances. It prefers disturbed areas with little vegetation, such as neglected meadows and
pastures, forest openings, fence rows, roadsides and other right-of-ways, and industrial
areas. It can be found in any vegetation type, including prairie, desert, chaparral, and
deciduous or coniferous forests. Mullein populations increase as rangeland deteriorates
due to heavy grazing, logging, or fires. Even digging by animals can create a small disturbed
area suitable for common mullein to invade. An early colonizer, it is often the pioneer
species on bare soils. It prefers well-drained, sandy or gravelly soils with a pH of 6.5–7.8,
but soil specifics are not important. It is intolerant of shade, but will grow in any open,
sunny area. It grows best with annual precipitation of 20–60 in. (50–150 cm) and requires
a 140-day growing season. It is found from sea level to 9,000 ft. (2,750 m) elevation, and
occasionally higher. Plants vary in size according to climate.
Reproduction and Dispersal. Common mullein reproduces solely by seed. Size of the over-
wintering rosette is important in determining the next season’s flowering. Over 50 percent of
rosettes smaller than 6 in. (15 cm) in diameter die over the winter, and rosettes smaller than
3.5 in. (9 cm) fail to flower. Individual flowers open for only a short period, from dawn to
midafternoon. If not pollinated by short-tongue or long-tongue bees, the flowers will self-
pollinate after they close. Each plant may have 200–300 fruit, with 500–800 seeds per seed
capsule, meaning that each plant can produce 100,000–240,000 seeds. Branched inflores-
cences produce even more. The form of mullein most frequently seen in Hawai’i has a fasciated
inflorescence, a twisted and compact irregular growth form, with more flowers and more seed.
With no specific adaptations for dispersal, most seeds fall within 3.3–16 ft. (1–5 m) of the
parent plant, although some are carried by wind and animals. Seeds are dispersed long dis-
tances as contaminants in soil moved for highway or building construction. They have no
dormancy requirement and remain viable in the soil for 35–100 years or longer. Seeds in soil
356 n FORBS

samples from archaeological sites dated 1300 AD were viable. It is the seed bank, not disper-
sal, that creates “instant” plants immediately following a disturbance. Seeds germinate faster
and seedlings grow more quickly on bare sites. Seeds at or near the surface germinate best,
even under diurnal temperature extremes, while those deeply buried do not. Seeds cannot
germinate, however, at temperatures below 50ºF (10ºC) or above 104ºF (40ºC). If seed ger-
minates in the fall, the plant becomes an annual and will bolt and flower the following
summer.
Impacts. Common mullein is rarely a significant weed in natural areas because it is easily
outcompeted by native plants. It is incapable of maintaining itself unless the site is repeat-
edly disturbed. It becomes a problem in bare ground or in soils with little vegetation, and
stands can cover acres or miles along a roadway. Plants create a dense ground cover that pre-
vents normal establishment of native species after disturbances. By growing on barren
ground exposed by forest fires, for example, common mullein alters the successional
sequence. It is eventually outcompeted by shrubs or other native plants in later successional
stages because mullein seeds are unable to germinate in the shade.
Common mullein, however, has invaded and colonized pristine, undisturbed meadows in
the Mono Lake and Owens Valley area of California. It also threatens sparsely vegetated alpine
sites on Hawai’ian volcanoes, from sea level to almost 15,080 ft. (4,600 m) on Mauna Kea. In
1990, the plant covered 770 sq. mi. (2,000 km2) in Hawai’i. It is found in several undisturbed
mountain communities, including subalpine grasslands dominated by alpine hairgrass, subal-
pine ohia lehua woodlands, as well as in alpine desert communities. The many bare sites on
volcanic soils are favorable to common mullein, and preferential grazing of other plants by
feral sheep and goats have probably helped it to spread. It inhibits revegetation by native
Hawai’ian species and specifically replaces the endangered silversword.
Common mullein is not a weed in agricultural crops because it does not survive repeated
cultivation. It is a host, however, for insect pests, such as the mullein leaf bug, which attacks
apples and pears in eastern states.
Management. Because common mullein is primarily a weed of bare soil surfaces, mainte-
nance of native plant cover or sowing bare sites with native plant seeds may prevent estab-
lishment. Its abundant seed production makes it difficult to totally eradicate. Any physical
control should minimize disturbance to the soil surface. Regular cultivation, such as disking
or plowing, however, controls infestations because it continually uproots seedlings. Plants
can be hand-pulled before seed sets, but stalks, flowers, and seeds should be bagged to pre-
vent seed dispersal. Plants cut below the lowest leaves will not resprout. Mowing is ineffec-
tive because the cuts are too high and the rosettes continue to grow. When mowing ceases,
plants bolt to flower. If only the flower stalk is cut, it will branch new stalks. Burning kills
both rosettes and bolted plants but, unless done selectively with a flamethrower, creates bare
soils that enhance more germination. Grazing is not an option because cattle and sheep
avoid the unpalatable hairy leaves, and livestock trampling exposes bare soil.
Applications of systemic herbicides, such as glyphosate or triclopyr, provide effective
chemical control as long as a surfactant is added to help the herbicide penetrate the thick
hairs on leaf surfaces. To prevent damage to nontarget plants, applications are best done in
early spring before native species break dormancy. Chemical applications are appropriate
on steep slopes or otherwise inaccessible areas or where physical removal would cause too
much soil disturbance. Herbicides do not need to be applied to the entire plant.
Application to the growing point in the center of the rosette is sufficient. Preemergent herbi-
cides are an option to control germination from the seed bank, but their application also
destroys seeds of desirable plants.
 COMMON MULLEIN n 357

Medicinal and Other Uses of Common Mullein

C ommon mullein has had many uses in Europe. The yellow flowers were used
for a hair dye as early as the fourth century BC. Aristotle recorded its use as
a fish poison, and Romans used the stalks, dipped in tallow, for torches. Plant
parts have been an herbal medicine for respiratory ailments, such as asthma,
tuberculosis, bronchitis, coughs, colds, congestion, and pneumonia. It was used
as a remedy for urinary tract infection, skin disease, hemorrhoids, diarrhea,
burns, earaches, warts, migraines, frostbite, and ringworm. Native Americans
readily adopted the plant. Hopi in Arizona smoked dried leaves and flowers, com-
bined with other plants, to treat mental illness. Other tribes in the Midwest and
East smoked the herb to treat respiratory problems, and concocted a cough syrup
from boiled roots. Mashed leaves were used as a poultice for bruises, wounds,
sprains, and headaches. The plant remains useful today. A methanol extract is
used against mosquito larvae, and the plant contributes to herbal eardrops for
children’s earaches. Recent studies have shown that mullein has antibacterial
and antitumor properties.

Two insects from Europe offer possible biological control. Maturing larvae of a host-
specific curculionid weevil (Gymnaetron tetrum), which was accidentally introduced to
North America sometime before 1937, eats seed capsules. Although the larvae can destroy
50 percent of a plant’s seeds, the weevil fails to limit the plant because of the number of
seeds produced. The mullein moth (Cucullia verbasci), which makes only limited use of
native species, has been tested in the United States. Two diseases, powdery mildew
(Erysiphe cichoracearum) and root rot (Phymatototrichum omnivorum), attack mullein, but also
affect commercial crops, including vegetables and cotton.

Selected References
Gucker, Corey L. “Verbascum thapsus.” In: Fire Effects Information System, USDA, Forest Service, Rocky
Mountain Research Station, Fire Sciences Laboratory, 2008. www.fs.fed.us/database/feis/plants/
forb/vertha/all.html.
Hoshovsky, Marc C. “Element Stewardship Abstract, Verbascum thapsus.” Global Invasive Species
Team, Nature Conservancy, 1986; modified 2009. http://wiki.bugwood.org/Verbascum_thapsus.
“Moth Mullein: Verbascum blattaria.” Virginia Tech Weed Identification Guide, n.d. http://www
.ppws.vt.edu/scott/weed_id/vesbl.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Verbascum thapsus.” ISSG Global Invasive Species Database, 2005. http://www.issg.org/
database/species/ecology.asp?si=695&fr=1&sts=sss&lang=EN.
Pitcairn, Michael J. “Verbascum thapsus L.” In Invasive Plants of California’s Wildlands. edited by Carla C.
Bossard, John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000.
http://www.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=87&survey
number=182.php.
Remaley, Tom. “Common Mullein.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/
veth1.htm.
358 n FORBS

n Common St. Johnswort

Also known as: Klamath weed, St. John’s wort, common goatweed, tipton weed
Scientific name: Hypericum perforatum
Synonyms: None
Family: Mangosteen (Clusiaceae)
Native Range. Europe, the Canary Islands, North Africa, and Asia, including aouthwes-
tern Asia, western Russia, Mongolia, and as far east as the China coast.
Distribution in the United States. Every state except Florida, including Alaska and
Hawai’i. Most problematic in the northern states.
Description. Common St. Johnswort is an erect perennial herb, with one stem or multiple
stems, that grows 1–4 ft. (0.3–
1.2 m) tall. Most branching is
toward the top of the plant.
Stems emerging from the lower
leaf axils are short, 0.8–4 in.
(2–10 cm), and sterile. The red-
dish or rust-colored stems are
woody at the base. Although
the stems are smooth, two lon-
gitudinal ridges, which are
covered with black glands,
make the stems two-sided. The
sessile leaves, opposite on the
stem, are narrow and lance-
shaped, 1–2 in. (2.5–5 cm) long,
with pointed tips. The green
leaves are covered with tiny
translucent glands, which can
be seen when the leaf is held to
the light. These glands contain
hypericin, a toxic substance.
Leaf margins are slightly rolled
under.
The plant has a complex root
system with a long, stout tap-
root. Many branching lateral
roots grow from the taproot, as
deep as 5 ft. (1.5 m). Rhizomes,
extending from the root crown,
grow just below the soil surface.
New shoots in the spring emerge
both from the root crown and
Common St. Johnswort is often found along roadsides and in other disturbed from the rhizomes.
ground and waste areas. (Native range adapted from USDA GRIN and From June to September,
selected references. Introduced range adapted from USDA PLANTS bright yellow star-like flowers,
Database, Invasive Plant Atlas of the United States, and selected references.) 0.75–1 in. (2–2.5 cm) in diameter,
 COMMON ST. JOHNSWORT n 359

A. The top of the stem may have several branches. (Eric Coombs, Oregon Department of Agriculture,
Bugwood.org.) B. Showy yellow flowers have many stamens. (Richard Old, XID Services, Inc., Bugwood.org.)
C. Margins of the lance-shaped leaves are slightly inrolled. (Steve Dewey, Utah State University, Bugwood.org.)
D. Fruit are clustered at the ends of stems. (Steve Dewey, Utah State University, Bugwood.org.)

grow in flat-topped clusters of 25–100. Each flower has five petals, sometimes with tiny black
dots and edges, and many yellow stamens. Fruit, a rust-brown three-part oval pod, less than
0.4 in. (1 cm) long, matures at the end of August. The capsule, which has no lobes, opens to
release many shiny dark brown seeds.
Aerial parts of the plant turn rusty red in fall, and the dead stalks may remain standing
through the winter or for several years.
Related or Similar Species. Canary Island St. Johnswort is an ornamental shrub, 13–16.4 ft.
(4–5 m) tall, with narrow leaves, 0.75–2.75 in. (2–7 cm) long. Its yellow-orange flowers are
0.5 in. (1.5 cm) in diameter, and the sepals are lined with hairs. The seed capsules are leathery.
A relatively recent invader in coastal California, its impacts are not yet fully known.
Several species of Hypericum are native to the United States. Dwarf St. Johnswort, found
growing in moist places such as riparian habitats, is an erect annual or perennial herb that
is usually 2 ft. (0.6 m) tall. Its stem is four-angled. Flowers are small, less than 0.25 in.
(0.6 cm) across, and the fruit is a single-cell capsule. Large St. Johnswort, an annual that
grows in moist sandy soil, has tiny flowers, linear leaves, and a one-compartment capsule.
Tinker’s penny is also an annual herb in wet places, growing prostrate, less than 1 ft.
(0.3 m) tall. Native species do not produce hypericin.
Introduction History. Currently cultivated in farms in Eastern Europe, common St.
Johnswort has been used medicinally for centuries. It was brought to the United States in
the seventeenth century by Rosicrucians, a group also known as the Ancient Order of the
Rosy Cross, which believes in mysticism and metaphysical connections. The plant escaped
from gardens, and by the Revolutionary War, it was common along roadsides from Maine
to Florida. Common St. Johnswort was introduced to California around 1900, and by
1940, about 7.5 million ac. (3 million ha) of rangelands in California and the Pacific
Northwest were infested.
Habitat. St. Johnswort grows best in temperate climates with dry summers and cool,
moist winters. The plant thrives in poor, dry soils in full sun and does not tolerate saturated
360 n FORBS

soils. It prefers coarse textured, gravelly or sandy soils, including sand dunes, and grows in a
wide range of acidity, pH 4.3–7.6. Common in waste areas and disturbed ground, initial
infestations are often associated with logging, fire, mining, or overgrazing. Plants also grow
in road and railroad right-of-ways and even in sidewalk cracks. The plant spreads from
those disturbed areas to natural open forests, healthy rangelands, meadows, dry pastures,
and fields.
Reproduction and Dispersal. Common St. Johnswort reproduces both sexually and veg-
etatively. Plants usually do not flower in their first year of growth. Flowers are both insect
pollinated and apomictic (needing no pollination). Every year, each plant typically produces
15,000–33,000 seeds, which are viable for 6–10 years. Wind, water, and animals carry seeds
long distances. The seed coat is gelatinous, allowing it to adhere to fur, feathers, clothes,
vehicle tires, and machinery. Seeds can also be transported in contaminated hay. After
4–6 months of dormancy, needed to break down the seed coat, seeds germinate from fall
to spring. They do best at temperatures 70–78ºF (21–25.5ºC) and on bare soil in full sun
after a heavy rain. Fire temperatures of 212–284ºF (100–140ºC) frequently stimulate more
seeds to germinate. Seedlings, however, grow slowly and are poor competitors with estab-
lished native grasses and other plants.
Buds on underground rhizomes and above-ground creeping stems may sprout and grow
roots. Independent plants subsequently separate from the parent. Vegetative propagation is
stimulated by mowing, grazing, or burning.
Impacts. By creating a dense canopy of vegetative growth early in spring, common
St. Johnswort shades out and displaces native plant species, which reduces forage for live-
stock and wildlife. Because they grow on disturbed sites, plants disrupt succession and delay
establishment of native species.
Although not palatable and only grazed when little else is available, all parts of
St. Johnswort are low to moderately toxic when eaten by livestock. The plant is toxic at all
stages of growth. The toxins remain active in dried plants and may be a contaminant in
hay or processed food. The toxin is absorbed in the intestinal tract, enters general circula-
tion, and is carried to the skin, where it is activated by ultraviolet rays. Sunlight reaching
the animal is abnormally converted into heat, causing sunburn, which damages cells. Skin
may become swollen and tender before turning red, and may be so badly burned that it peels
off in sheets. It also causes inflammation of mucus membranes, resulting in swelling, blis-
ters, and open sores, with a risk of infection. Other symptoms may include itching, fever,
rapid pulse, and diarrhea. It is especially toxic to cattle and sheep, but also to goats, horses,
and swine. It primarily affects areas with light pigmentation, such as pink or white skin,
udders and teats, and the delicate tissue around the mouth and eyes. Parts of the body that
receive more sun, such as head, neck, and back, are also affected. Poisoning can develop
after the animal has consumed 1–4 percent of its body weight or eaten it for 4–5 days.
Animals will act abnormally, and pain may prevent them from eating or drinking.
Inflammation in the eyes may affect vision. Although moving affected animals into shade
will be sufficient for mild cases, severe cases, including damaged eyes or blistered skin,
require veterinary care, including antibiotics or other medication. Although rarely resulting
in death, the toxin causes economic losses.
The accumulation of standing dry stalks presents a fire hazard in late summer and fall.
Management. Common St. Johnswort is difficult to eradicate because of its extensive root
system and abundant seeds. Maintaining rangeland in good condition will help to prevent its
encroachment. Although it must be ingested to affect animals, workers should wear gloves
 COMMON ST. JOHNSWORT n 361

St. Johnswort in Medicine

T he plant has been used for centuries in Europe for various ailments. It was a
treatment for wounds, an astringent, a sedative, and a diuretic. It was also used
to treat hysteria. The toxin, hypericin, in St. Johnswort is an antidepressant ingre-
dient in herbal medicine. Research using lab animals has indicated that hypericin
was effective in treating several viruses, including herpes and hepatitis B. In 1991,
studies were begun to test the effectiveness of very high doses of hypericin against
human immunodeficiency virus (HIV). The trial was aborted, however, when the
patients became very sensitive to sunlight and developed skin rashes. One dark-
skinned patient, however, was not affected. Because limited studies indicate that
St. Johnswort extracts interact with other medications, further research should
proceed cautiously.

and avoid touching their eyes while removing common St. Johnswort because contact with
the plant may cause skin blisters on sensitive persons.
Physical control, such as tilling or mowing, will limit seed production but stimulate veg-
etative reproduction and new growth. Burning will stimulate both seed germination and
sprouting of root buds. Repeated tillage may be effective in agricultural fields, but may also
spread root fragments. New plants or small infestations can be effectively pulled by hand,
providing that the entire plant, including the root system, is removed. All plant parts and
seeds must be disposed of offsite because they can sprout. Large stands may be mowed,
sprayed with herbicides, and reseeded to improve pasture.
The waxy leaves limit the uptake of chemical applications. St. Johnswort can be tempo-
rarily suppressed in pasture and rangeland by several herbicides, including aminopyralid,
metsulfuron, picloram, and 2,4-D. Each type is effective during a different stage in the
plant’s life cycle. Herbicides, however, often increase the palatability of St. Johnswort and
exacerbate the problem of toxicity.
St. Johnswort was the first weed to be targeted for biological control in the United States.
After being released in California in 1945 and 1946, two foliage beetles (Chrysolina hyperici
and C. quadrigemina), became established in two years. The larvae, feeding on leaves and
flowers, can completely defoliate plants. Although the beetles have reduced infestations by
97–99 percent, new outbreaks can occur locally following logging, fire, or some other type
of disturbance. In 1950, a root-boring beetle (Agrilus hyperici) was released. Larvae feed on
roots for almost a year, stunting plants and reducing flowering. The California colonies of
beetles became the source for further collection and release elsewhere. Those insects, how-
ever, do not thrive at high elevations or in cold climates. The larvae of a moth (Aplocera pla-
giata), more recently released and established, eats leaves and flowers, defoliating plants and
inhibiting seed production.

Selected References
“Common St. Johnswort.” Non-Native Species of Alaska. Alaska Natural Heritage Program,
Environment and Natural Resources Institute, University of Alaska, Anchorage, 2005. http://
akweeds.uaa.alaska.edu/pdfs/species_bios_pdfs/Species_bios_HYPE.pdf.
362 n FORBS

“Common St. Johnswort or Klamathweed.” Plant Pest and Health Prevention Services (PHHPS),
California Department of Food and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/
hypericum.htm.
“Fact Sheet 729, St. Johns Wort (Hypericin).” AIDS InfoNet, New Mexico AIDS Education and Training
Center, Health Sciences Center, University of New Mexico, 2009. http://www.aidsinfonet.org/fact
_sheets/view/729.
Goetz, Rebecca J., Thomas N. Jordan, John W. McCain, and Nancy Y. Su. “Common St. Johnswort,
Klamath Weed.” Cooperative Extension Service, School of Veterinary Medicine, Purdue
University, West Lafayette, IN, n.d. http://www.vet.purdue.edu/toxic/plant38.htm.
Jacobs, Jim. “Ecology and Management of Common St. Johnswort (Hypericum perforatum L.)” Natural
Resources Conservation Service, Invasive Species, Technical Note No. MT-14, U.S. Department of
Agriculture, Montana, 2007. http://www.mt.nrcs.usda.gov/technical/ecs/invasive/technotes/invasi
vetechnotemt14/.

n Dyer’s Woad
Also known as: Marlahan mustard
Scientific name: Isatis tinctoria
Synonyms: None
Family: Mustard (Brassicaceae)
Native Range. Grasslands and desert regions of Central Asia, especially the Caucasus
region. Possibly western China, western Tibet, and Afghanistan. Widespread in Europe.
Distribution in the United States. Eastern states, from New York south to Virginia, Illinois
and Missouri in the Midwest, and from the Rocky Mountains west to the Pacific coast.
Description. Usually a biennial, dyer’s woad may also be a short-lived perennial or a win-
ter annual. Either the seed or the rosette must undergo a period of cold temperatures before
the plant can flower. Most germination occurs in spring. The plant grows a basal rosette of
leaves and overwinters. In its second spring, the plant bolts, sending up flower stalks that
are 1–4 ft. (0.3–1.2 m) tall. If conditions are poor in the second season, the plant can con-
tinue growing as a rosette for a third year. A few seeds, however, germinate in the fall and
live as annuals, flowering and producing seed the following spring.
Leaves, both basal and stem, are bluish green, with a prominent cream-colored midrib from
base to tip. The basal rosette leaves, oblong or lance-shaped, 2–8 in. (5–20 cm) long, are slightly
succulent. They have small, rounded teeth and are covered with fine hairs, especially along the
midrib. Broad at the tip, the leaves narrow toward the base, grading into a short petiole. Leaves
on flowering stems are smaller, lance shaped, and glabrous, with entire margins. They are ses-
sile, in an alternate arrangement, and the lower part of the leaf clasps the stem with auricles.
The plant has a thick taproot, 3–5 ft. (1–1.5 m) long. Lateral roots, that grow in the top
12 in. (30 cm) of soil, branch during the second season.
Plants may have as many as 20 flowering stems. Most flower stems grow erect from the
base, but the lower branches are decumbent, paralleling the ground and turning up at the
tips. The tiny yellow flowers, 0.25 in. (0.6 cm) in diameter, grow in flat-topped, elongated
clusters on branched racemes. Typical of the mustard family, they have four narrow petals
that form a cross. Flowers are showy, crowded on both the plant and the stems. Flowers will
be blooming at the top of the raceme, while seeds are developing below. The winged fruit is
a narrow, flattened pod (silicle), 0.5–0.75 in. (1.2–2 cm) long and 0.25 in. (0.6 cm) wide,
with the tip slightly wider than the base. Pods, which hang down from the stalk on
short pedicels, are green and hairless when young, but become black or dark purplish
brown as they mature. Pods are indehiscent and cling to the stem. Each pod has one
 DYER’S WOAD n 363

brownish-yellow cylindrical
seed, 0.1 in. (3 mm) long. Fruit
matures in late spring to early
summer, and plants die after
they produce seed.
Related or Similar Species.
Although many mustard species
have showy yellow flowers,
none have a similar accumula-
tion of black seed pods lining
the old flower stems.
Introduction History. Dyer’s
woad was brought to colonial
Virginia in the late seventeenth
century, where it was cultivated
for blue dye. It is a more recent
invader of western states, intro-
duced in the early 1900s in
contaminated alfalfa seed, both
to northern California and to
Utah. In 1934, a specimen was
noted in the pharmaceutical
garden at the University of
Montana, and by 2006, dyer’s
woad infested at least 12 coun-
ties in that state. The first iden-
tification in Washington, in
1986, was along a railroad.
Although that stand was eradi-
cated in 1992, the site is still
monitored. The plant has the
capacity to continue expanding By reducing forage for livestock, dyer’s woad threatens pastures and
its range in the United States. rangeland in western states. (Native range approximated from USDA
Habitat. Although dyer’s GRIN and selected references. Introduced range adapted from USDA
woad can infest both disturbed PLANTS Database, Invasive Plant Atlas of the United States, and selected
references.)
and undisturbed communities,
it is often found in rugged, inac-
cessible terrain that is usually gravelly, rocky, or sandy. The plant will also grow in loose
alkaline soil, and its long taproot enables it to thrive in the arid steppe regions of the West.
The species requires little nitrogen and can thrive on nutrient-poor soil. Woad may first
appear in gravel pits and waste areas, or along road, levee, and railroad right-of-ways, before
spreading into well-vegetated rangeland, pasture, forest, and wildland. Plants may also
invade row crops and orchards. They can grow at 3,000–8,000 ft. (1,000–2,500 m) eleva-
tion, usually in full sun.
Reproduction and Dispersal. Rosettes grow quickly after snowmelt, and seeds develop
4–8 weeks after the flower stalks begin to grow. Each plant usually produces 350–500 seeds.
About 95 percent of the indehiscent seed pods fall to the ground within 2 ft. (0.6 m) of the
parent plant, but the winged pods may be blown several feet, especially over a snow crust.
364 n FORBS

A. Dyer’s woad is bushy with tall flowerstalks. B. Fine hairs highlight the midribs on the basal rosette leaves. C.
Each seed pod has one seed. D. Flowering stem, immature green seed pods, and mature black pods. (Steve
Dewey, Utah State University, Bugwood.org.)

Seeds may be transported long distances along roads, trails, or railways. The pods float in
water, and seeds may be carried by animals, both attached to fur and within the gastrointesti-
nal tract. Seeds may be a contaminant in crop seeds, livestock bedding, or in hay or other live-
stock feed. Most seeds are viable for only a short term, but intact pods that are buried in the
soil may remain viable for 10 years. Seed pods contain a water-soluble chemical that inhibits
germination, and seeds remain in the seed bank until rainfall is sufficient to wash it away,
ensuring that the seedling has enough moisture for growth. Spring germination may be more
common than fall germination because melting winter snows leach the chemical out of the
seed. Seeds germinate under a wide range of temperature, 37–77ºF (3–25ºC).
Although the root crown has dormant buds that can sprout if the top part of the plant is
damaged, plants do not reproduce vegetatively.
Impacts. The biggest threat of dyer’s woad is to rangelands and pastures, where it can
form dense infestations, outcompete native plants, and reduce forage. It grows rapidly, out-
competing native plants for spring moisture. Currently infesting thousands of acres of
western rangeland, dyer’s woad lowers the quality and capacity of rangeland. It has been
estimated to reduce grazing capacity of range by 38 percent. It also competes with crops,
costing millions of dollars in diminished crop yields. Although sheep will eat the young
rosettes when nothing else is available, dyer’s woad is unpalatable to livestock. An accumu-
lation in the soil of the chemical which inhibits dyer’s woad germination may be alleleo-
pathic to other species, but it is not toxic to animals.
Management. At least in some areas of the western states, dyer’s woad appears to be in
the introductory stages of invasion. The goal is to eradicate it before it spreads further, and
any method must be repeated because seeds are persistent in the seed bank for several years.
Physical control can be accomplished by intensive labor of hand-pulling or digging out
plants, including the taproot and root crown, because latent buds on the root crown will
sprout. Because fewer than 25 percent of young rosettes survive, compared with 80 percent
of mature rosettes, removal efforts should be concentrated on mature rosettes and flowering
plants, after plants have bolted but before seed is produced. Removing young rosettes would
duplicate their natural mortality. Hand-pulling, however, should be repeated in 3–4 weeks
to remove the later-maturing plants. Hand removal may be the only method possible on dif-
ficult terrain. Because even green pods may set seed, all pods on plants should be burned or
removed to a landfill. Cropland may be kept free of dyer’s woad by a combination of tilling,
 DYER’S WOAD n 365

Successful Eradication—but a Lot of Work!

V olunteer labor from the Montana Dyer’s Woad Cooperative Project, which was
established in 1984, had, by 2007, significantly reduced the dyer’s woad popu-
lation in that state, from 13 infested counties down to 4, and from 480 ac. (194 ha)
down to 6.4 ac. (2.6 ha). After eight years of volunteer labor in Utah, a population
of woad was reduced by 95 percent.
Source: Jacobs and Pokorny, 2007.

crop rotation, and herbicides. Tilling in spring will kill new seedlings and rosettes. Seedlings
that emerge in summer or fall cannot flower and produce seed until they undergo a cold
period. Those seedlings can be tilled or sprayed with herbicide. Mowing plants in orchards,
if repeated when new stems sprout from the root crown, will prevent seed development.
Although sheep may eat the young rosettes in early spring when nothing else is available,
grazing offers little control.
The choice of chemical treatments is dependent on the plant’s stage of growth.
Applications of metsulfuron or chlorsulfuron effectively prevent seed production when
applied to stems in the late bloom stage. Imazapic may also be applied to bolting rosettes.
Limiting seed production will decrease the number of rosettes the following spring. The
most effective herbicide on the seedling to rosette stage is 2,4-D, but it does not control
the plant after flowering begins.
The only known biological control is a rust fungus thought to be Puccinia thlaspeos, a
native to Eurasia. First discovered in southern Idaho in 1978, it has since spread to other
states. It is a systemic, entering the plants through leaves. Although it takes 3–9 months
for the plant to succumb, the rust kills seedlings and young rosettes and completely prevents
seed production on most infected plants. It has spread naturally to new populations, and
deliberate transfer is being investigated.

Selected References
“Dyer’s Woad.” Weed of the Week. U.S. Department of Agriculture, Forest Service, Invasive Plants,
2006. http://www.invasive.org/weedcd/pdfs/wow/dyers_woad.pdf.

Uses of Dyer’s Woad

D yer’s woad has long been used for dye and medicine. It was cultivated as a
source of blue dye in Europe until it was replaced in the 1600s by indigo from
the East Indies. Cultivation ceased in England in 1930. The source of the blue dye is
indigotine, present in the leaves and extracted through a fermentation process.
Although woad has a history of medicinal use, and may hold promise for future
treatments, no clinical trials have been conducted regarding its safety. Root extracts
are a rich source of the anticancer chemical glucobrassicin. Plant parts have been
used in Chinese medicine for a wide variety of ailments, including throat infections,
mumps, encephalitis, gastroenteritis, hepatitis, headaches, and fever. Chinese stud-
ies indicate that dyer’s woad is antibacterial, antiviral, and antiparasitic.
366 n FORBS

“Dyers Woad (Isatis tinctoria L.).” Written Findings of the State Noxious Weed Control Board—Class A Weed,
State of Washington, 1999. www.nwcb.wa.gov/weed_info/Written_findings/Isatis_tinctoria.html.
Jacobs, Jim, and Monica Pokorny. “Ecology and Management of Dyer’s Woad (Isatis tinctoria L.).”
Invasive Species Technical Note No. MT-10. Natural Resources Conservation Service, U.S.
Department of Agriculture, 2007.
Roberts, Teresinha. “Woad Facts.” 2010. http://www.woad.org.uk/html/woad_facts.htm.

n Fig Buttercup
Also known as: Lesser celadine, pilewort, bulbous buttercup, small crowfoot
Scientific name: Ficaria verna
Synonyms: Ranunculus ficaria, Ficaria ranunculoides
Family: Buttercup (Ranunculaceae)
Native Range. Europe,
northern coast of Africa, and
western Russia.
Distribution in the United
States. Upper midwestern and
northeastern states, from Wis-
consin east to New Hampshire,
south to Virginia, and west to
Missouri, Illinois, and Texas.
Also in Oregon and Washington.
Description. Fig buttercup is
an herbaceous perennial plant
with a short life cycle, growing
in late winter and spring. The
species has variable morphol-
ogy due to several cultivars
and varieties. Shiny, dark-
green leaves begin to emerge
in winter, forming a low-
growing basal rosette. De-
pending on the variety or sub-
species, plants are either erect
or reclining, 4–12 in. (10–
30 cm) tall. The tender, fleshy
leaves, supported by long
petioles, are kidney- to heart-
shaped, 0.7–2 in. (1.8–5 cm)
in both length and width. Leaf
tips are rounded, and margins
are usually entire or wavy. The
glabrous, fleshy stems have
closely spaced nodes where
Ornamental varieties of fig buttercup are sold by many plant nurseries
throughout the country. (Native range adapted from USDA GRIN and bulblets may develop.
selected references. Introduced range adapted from USDA PLANTS The root system is both fi-
Database, Invasive Plant Atlas of the United States, and selected references.) brous and tuberous. The leaves
 FIG BUTTERCUP n 367

A. Butter yellow flowers bloom in early spring. B. Edges of the heart-shaped leaves may be wavy. C. New sprouts
grow from clusters of tubers. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology University of Connecticut,
Bugwood.org.)

and flower stalks grow from a cluster of tubers. Many turions, which are young scaly shoots,
grow on the roots.
Plants flower in late winter or early spring, March to May, with one terminal flower on
each stem. The delicate flower stalks grow taller than the leaves of the plant. More flowers
are produced in high light conditions and on plants with larger tubers. The flowers are sym-
metrical, about 1 in. (2–3 cm) in diameter with 8–12 glossy yellow petals. They are slightly
darker in the center and fade to white as they age. In May, each flower may produce a globu-
lar fruiting head, about 0.25 in. (0.6 cm) in diameter, composed of several pubescent
achenes (dry, one-seeded fruits).
After flowering, plants die back, and by early June, most are dormant, reduced to thick
tubers in the ground.
Related or Similar Species. At least nine cultivars and 100 varieties of fig buttercup have
been developed, including a double-flower form. Some varieties have leaves with silvery or
whitish mottling or black spots. Most buttercup species have upright stems, while fig butter-
cup may have either upright or decumbent stems, meaning that the stems lie parallel to the
ground, but with the tip upright.
Marsh marigold, native to wetlands in eastern United States, is a more robust plant with
larger, lighter green leaves. Its flower stalks, at least 8 in. (20 cm) tall, grow upright and sup-
port a number of flowers. It has no petals, but the 5–9 sepals are a deep yellow and resemble
petals. The fruit is a follicle or dry seed pod. Marsh marigold produces no tubers or bulblets
and never grows as a continuous carpet.
Celandine, also called greater celandine, grows much taller and has distinctly lobed
leaves. Celandine poppy is a tall native perennial with a hairy stem and deeply lobed, large
leaves. Both plants are in the poppy family and have flowers with four petals. Ground ivy,
also known as gill-over-the-ground and creeping Charlie, is a creeping perennial ground-
cover in the mint family. It is distinguished from fig buttercup by having hairy leaves and
lavender asymmetrical flowers.
Introduction History. Fig buttercup was introduced to the eastern United States as an
attractive ornamental, probably in the mid-1700s. Many varieties are widely available from
nurseries, catalogs, and Internet sales.
Habitat. Fig buttercup grows best in the moist alluvial soil of herbaceous wetlands and
forested floodplains, where it forms large dense patches that carpet the ground. It also occu-
pies some upland sites, including grassy meadows and early successional forests. The spe-
cies is less frequently found in drier soils, such as embankments, roadsides, and open
woodlands. Although it may tolerate dry sites during summer dormancy, seasonally wet
368 n FORBS

areas provide plants with moisture in the spring when they are actively growing. In the same
manner, it grows in sunny sites in wooded areas in spring, but is dormant in summer when
the canopy provides shade. It also grows in disturbed sites, such as abandoned fields, pas-
tures, roadsides, and vacant lots, and is a weed in gardens and lawns. Some evidence exists
that fig buttercup prefers sandy and slightly alkaline soil.
Reproduction and Dispersal. Although each flower may produce 10–15 achenes, fig but-
tercup seeds are rare because most abort before reaching maturity. Approximately 60 percent
of seeds that are produced are viable. No information is available regarding a seed bank.
Most reproduction is vegetative, by tubers and bulblets. Small cream-colored bulblets, each
of which can produce a new plant, grow in stem axils or nodes. Plants have many fleshy, finger-
like tubers with several turions, meaning shoots that are ready to develop new plants. Tubers
and turions survive for years and are easily transported. Tubers may be uprooted and scattered
by digging or burrowing animals. Tubers may also be dispersed in floodwaters or as a contam-
inant in transported soil, or by human activity, such as weed-pulling or deliberate planting.
Impacts. Fig buttercup is an aggressive plant that spreads rapidly, creating monocultures
that cover acres of ground. Plants begin growth early in spring, in sunlight before the leaf can-
opy of trees and shrubs develops. Because fig buttercup emerges first and is a vigorous spring
grower, it displaces native plants, especially those with a spring-flowering life cycle, including
but not limited to bloodroot, wild ginger, spring beauty, harbinger-of-spring, twinleaf,
squirrel-corn, trout lily, trillium, and Virginia bluebells. As a result, the structure of the native
community is altered. These native spring ephemerals are important sources of nectar and pol-
len for native pollinators, such as bees, small beetles, and flies. These native plants also pro-
duce fruit and seeds, which are food for native species of insects and other wildlife.
Management. Fig buttercup’s abbreviated life cycle allows management opportunities in
only a short window of time when it is growing. Because it is difficult to remove all bulblets
and tubers, physical control, such as pulling or digging out plants, is practical only on small
infestations. Such methods also cause soil disturbance, which creates sites for more invasion.
Chemical treatment is preferable for controlling or eradicating fig buttercup. Glyphosate,
a systemic herbicide, will kill tubers, but the treatment should be repeated the following
year. Herbicides should be sprayed on the leaves in late winter or early spring, either before
plants flower or when approximately 50 percent of plants are in the bloom stage. Spraying
before native ephemerals begin to grow ensures that few to no desirable plants will be dam-
aged. Native amphibians, such as frogs and salamanders that can absorb toxins through their
skin, are also less likely to be harmed by early spring applications of herbicides because they
have yet to come out of hibernation.
No biological controls are known or are being investigated.

Selected References
“Fig Buttercup.” Invasive Plant Atlas of New England (IPANE), University of Connecticut, 2009. http://
nbii-nin.ciesin.columbia.edu/ipane/icat/browse.do?specieID=89.
Huebner, Cynthia, Cassandra Olson, and Heather Smith. “Lesser Celadine.” Invasive Plants Field and
Reference Guide: An Ecological Perspective of Plant Invaders of Forests and Woodlands-Supplement 2.
USDA Forest Service, State and Private Forestry Technical Publication 05-04, 2008. http://
na.fs.fed.us/pubs/misc/ip/ip_field_guide_supp2_lr.pdf.
“Lesser Celandine.” Oregon Department of Agriculture Plant Division, Noxious Weed Control, n.d.
http://www.oregon.gov/ODA/PLANT/WEEDS/profile_lessercelandine.shtml.
Swearingen, Jil.“Fig Buttercup.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2010. http://www.nps.gov/plants/alien/fact/
rafi1.htm.
 GARLIC MUSTARD n 369

n Garlic Mustard
Also known as: hedge garlic, sauce-alone, jack-by-the-hedge, poor-man’s mustard
Scientific name: Alliaria petiolata
Synonyms: Alliaria officinalis, A. alliaria, Sisymbrium alliaria, S. officinalis, Erysium alliaria
Family: Mustard (Brassicaceae)
Native Range. Southern Europe, northern Africa, and western Russia. Possibly
Southwest Asia as far east as India and western China.
Distribution in the United States. Most abundant in the Midwest and New England, from
North Dakota east to Maine, south to Georgia, and west to Oklahoma, Colorado, and Utah,
and the Northwest coast. Also in Alaska.
Description. Garlic mustard,
so named because the dark
green leaves emit a garlic smell
when crushed, is a biennial
herb, meaning that it has a
two-year life cycle. In the first
year, the plant grows a low
rosette of leaves, 4 in. (10 cm)
tall and 2.4–4 in. (6–10 cm) in
diameter by fall. The plant
remains green and continues to
grow in winter during snow-
free periods when temperatures
are above freezing. In the sec-
ond summer, plants send up a
flowering stem, 2–3.5 ft. (0.6–
1 m) tall. First-year leaves are
round to kidney-shaped, 2.4–4
in. (6–10 cm) in diameter, with
scalloped edges. The alternately
arranged stem leaves are tri-
angular or heart-shaped, 1.2–
3.2 in. (3–8 cm) both long and
wide, and coarsely toothed.
They become gradually smaller
toward the top of the stem. All
leaves grow on pubescent
petioles, 0.4–2 in. (1–5 cm)
long.
Plants usually have one or two
flowering stems, with few or no
branches. Small white flowers,
Garlic mustard grows in a number of habitats, which vary according to
0.3 in. (7 mm) in diameter, geographic region. (Native range adapted from USDA GRIN and
appear in late April and May. selected references. Introduced range adapted from USDA PLANTS
Flowers, in button-like clusters, Database, Invasive Plant Atlas of the United States, and selected
usually grow on a terminal references.)
370 n FORBS

A. Flowering stems may be more than 2 ft. (0.6 m) tall. (Chris Evans, River to River CWMA, Bugwood.org.)
B. The four petals of each flower form a small cross. (Chris Evans, River to River CWMA, Bugwood.org.)
C. Rosette leaves are distinctly kidney-shaped. (Ohio State Weed Lab Archive, The Ohio State University,
Bugwood.org.) D. Triangular stem leaves have pointed tips. (Linda Haugen, USDA Forest Service,
Bugwood.org.) E. Fruit are very slender seed pods. (James H. Miller, USDA Forest Service, Bugwood.org.)

raceme, but they can also occur on short racemes in leaf axils. The four narrow petals form the
shape of a cross.
Fruit, which develops in May and June, is a four-sided slender pod called a silique, 1–2.5 in.
(2.5–6.3 cm) long. Siliques, averaging 9–19 per plant, grow erect on short, stout pedicels
0.2 in. (.5 cm) long. Each silique contains 10–20 tiny black, cylindrical, ridged seeds,
arranged in two rows. By June, most plants are dead, but the erect flower stalks with dry
pale-brown seedpods remain through the summer, sometimes still holding seeds.
Related or Similar Species. Several white-flowered native herbs, such as toothworts,
sweet cicely, and early saxifrage, grow alongside garlic mustard. First-year garlic mustard
plants may be confused with other rosette species, especially violets, white avens, and tooth-
wort species in the Midwest and Northeast, and with fringecup and piggyback plant in the
West.
Although the effect dissipates toward autumn, garlic mustard can be distinguished by the
garlic odor of its foliage. Plants have a slender white taproot with a distinctive crook shape
close to the soil surface.
Introduction History. Because garlic mustard can be used as a potherb or in salad, espe-
cially the rosette leaves in winter and early spring, it was probably introduced multiple times
by settlers for food or medicine. It was first recorded in 1868 in Long Island, New York. By
1991, it had spread to 28 midwestern and northeastern states. It is considered a gourmet
ingredient, and seeds can be ordered off the Internet.
Habitat. Garlic mustard grows in a wide range of habitats, most often in deciduous forest
communities with partial shade. It is found in both full sun and shade, upland forests, ripar-
ian forests on floodplains, roadsides, trails, forest edges, oak savannas, and urban areas. It is
less frequent in woodlots surrounded by cropland. Although rarely found under conifers in
the midwestern states, it invades those habitats in Ontario, Canada. Its habitat varies with
geography. Garlic mustard is river-associated in the Northeast and along railroad right-of-
ways in the Midwest. It grows in riparian habitats and on waste ground on the Great
Plains and along hiking trails in the eastern Rocky Mountains. Disturbed areas are most
 GARLIC MUSTARD n 371

vulnerable to rapid invasion and dominance, but garlic mustard can also spread into healthy
communities. It grows best in rich, moist soil, preferably calcareous, and does not tolerate
high acidity. It can grow on sand, loam, clay, limestone, and sandstone, but is rare on peat
or mucky soils. Flooding during the growing season may limit its extent.
Reproduction and Dispersal. Although damaged plants may sprout new flowering stems
from the root crown, garlic mustard reproduces exclusively by seed. Plants develop rapidly
in their second spring, flowering, setting seed, and dying by late June. Flowers may be pol-
linated by bees and flies, but are also self-fertile, enabling one plant to populate an entire
site. Each plant can produce over 500 seeds, allowing an extensive seed bank to accumulate
in the soil. The seed pods burst when ripe, expelling seeds several feet from the parent plant,
but humans and wildlife are responsible for long-distance dispersal. Rodents and birds also
contribute to seed dispersal. Because seeds are easily transported by water, floodplains are
vulnerable to invasion. Abundant seeds enable garlic mustard to rapidly invade sites after a
disturbance. While browsing other plants, white-tailed deer trample the soil, inadvertently
providing more disturbed area for garlic mustard.
Seeds remain viable for five or more years but require 8–22 months of dormancy before
germinating. Seeds germinate in early spring, which may be late February to mid-May,
depending on geography. While native plants require light to germinate, garlic mustard
has an advantage because it is able to germinate in both sun and shade. Although initial
seedling density can be 500 or more per sq. ft. (5,000 per m2), only 2–4 percent survive
to flower the following spring.
Impacts. Because it grows in early spring and late fall when native plants are dormant, gar-
lic mustard is one of the few plants able to invade the understory of forests in the United
States. It dominates the herbaceous layer in woodlands, eliminating native plants and ani-
mals. Garlic mustard outcompetes native wildflowers, such as spring beauty, wild ginger,
bloodroot, Dutchman’s breeches, hepatica, toothworts, and trillium, which also complete
their life cycles in the spring. When these wildflowers are displaced, so is the wildlife that
depend on them for pollen, nectar, foliage, fruits and seeds. Garlic mustard threatens the
existence of native garden white butterflies. It appears to be toxic to the butterfly eggs because
they fail to hatch when laid on that plant. The rare West Virginia white butterfly is dependent
on two species of toothwort, crinkleroot, which is also called twoleaf toothwort, and cutleaf
toothwort, as hosts for their eggs, plants which are often displaced by garlic mustard.
Management. Eradication or control of garlic mustard is difficult due to the enormous
numbers of seeds produced and their long-term viability. The best control is prevention
and early detection. The goal should be to exhaust the seed supply, and sites should be
monitored for seedlings for at least five years.
Any physical removal of plants should minimize soil disturbance and be done before
seeds mature. Hand removal is effective for small infestations or especially when garlic mus-
tard threatens the survival of native plants, but the entire root system must be removed to
prevent resprouting. Plants in larger areas can be cut at ground level every year until the seed
bank is exhausted. If plants are cut too high on the stem, flowers will grow from leaf axils.
All plant fragments should be removed from the site, especially if flowers or seedpods are
present. Fire can be used in fire-adapted communities, but the ground disturbance opens
the surface to seeds and encourages germination. If fire is used, sites must be burned annu-
ally for 3–5 years to deplete the seed bank.
Chemical control is appropriate for heavy infestations where few native plants remain.
Glyphosate is best applied to dormant rosettes in late fall or early spring, any time the tem-
perature is above 50ºF (10ºC), and when native plants are still dormant.
372 n FORBS

Possibilities of biological control are still being investigated. Garlic mustard has no natu-
ral enemies in the United States, but at least 69 species of insects and 7 species of fungi are
associated with the plant in Europe. Five weevils and one flea beetle are being studied. Two
weevil species (Ceutorhynchus alliariae and C. roberti) are effective in reducing seed produc-
tion. Adults eat the leaves, and the larvae develop inside the stems and leaf petioles. The lar-
vae of a root-mining weevil (C. scrobicollis) first mine the petioles and rosette leaves but do
most of their feeding on the root crowns. Although the larvae of a fourth weevil
(Ceutorhynchus constrictus) eat developing seeds, they fail to provide control. A fifth weevil
(Ceutorhynchus theonae), more recently discovered in Russia, also attacks seeds and may be
more effective. Impact of the flea beetle (Phyllotreta ochripes) is not yet known. Adults eat
the leaves, while larvae eat the roots, rosettes, and bolting plants.

Selected References
Blossey, Bernd, Victoria A. Nuzzo, Hariet L. Hinz, and Esther Gerber. “Garlic Mustard.” In Biological
Control of Invasive Plants in the Eastern United States, by R. Van Driesche et al. USDA Forest
Service Publication FHTET-2002-04, Morgantown, WV, 2002. http://www.invasive.org/eastern/
biocontrol/29GarlicMustard.html.
“Garlic Mustard—Alliaria petiolata.” Southeast Exotic Pest Plant Council (SEEPPC) Invasive Plant
Manual. Invasive Plants of the Eastern United States, 2003. http://www.invasive.org/eastern/eppc/
garlicmustard.html.
Nuzzo, Victoria. “Element Stewardship Abstract, Alliaria petiolata.” Global Invasive Species Team,
Nature Conservancy, 2000; updated 2009. http://wiki.bugwood.org/Alliaria_petiolata.
Rowe, Pamela, and Jil M. Swearingen. “Garlic Mustard.” Weeds Gone Wild: Alien Plant Invaders of
Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://www
.nps.gov/plants/ALIEN/fact/alpe1.htm.

n Giant Hogweed
Also known as: Giant cow-parsnip, cartwheel-flower
Scientific name: Heracleum mantegazzianum
Synonyms: Heracleum caucasicum, H. giganteum, H. panaces, H. speciosum, H. tauricum
Family: Carrot (Apiaceae)
Native Range. Caucasus region of Southwest Asia.
Distribution in the United States. New England, northern midwestern states, and
Washington and Oregon.
Description. Giant hogweed is an herbaceous perennial that often reaches 10–20 ft. (3–
6 m) tall at the flowering stage. The plant is a rosette of leaves during its first year of growth,
and leaf clusters become larger as the plant becomes older. After accumulating enough
energy, over 2–5 years, plants bolt and flower in midsummer. The stout stems, 2–4 in. (5–
10 cm) in diameter, are coarse and ridged, with prominent white hairs, especially at the
nodes. The stems and leaf stalks, which are both hollow, have dark reddish-purple spots.
Both the leaf stalks and stems have small pustules (blister-like spots), each with a single
sturdy bristle, that contain toxic sap. Leaves are alternate on the stem and have various
shapes and sizes at different life stages. Young leaves at the base of the plant in spring are
ternately compound, meaning that divisions each have three leaflets. As the plant grows,
those leaves become larger, as wide as 5 ft. (1.5 m), with deeply incised lobes. Some leaves
have measured almost 10 ft. (3 m) wide. Higher on the stem, the leaves are smaller, and
 GIANT HOGWEED n 373

the lobes are even more deeply

cut. The lower surface of the
leaves is densely covered with
short, stiff hairs.
Young plants develop a tap-
root in their first year. Lateral
roots also develop, and the
extensive root system gives the
large plant stability. Foliage dies
to the ground in winter, and
buds on the tuberous root stalk
sprout new leaves each spring.
Plants flower in June and
July. The large inflorescence, as
wide as 2.5 ft. (.75 m), is a
broad flat-topped, compound
umbel, with many small white
or greenish-white flowers. The
small dry fruit, flattened, win-
ged, and oval in shape, are
approximately 0.4 in. (1 cm)
long. Each contains one seed.
Seeds are marked with brown
swollen resin canals. Plants are
believed to be monocarpic and
die after flowering, although
the dead stems often remain
standing.
Related or Similar Species.
Although several native species
in the carrot family resemble
giant hogweed, the size of the Although currently present in few states, giant hogweed may become
plant makes it distinct. Com- more expansive due to its unique qualities as an ornamental. (Native
mon, or cow, parsnip is a range adapted from USDA GRIN and selected references. Introduced
smaller plant, usually less than range adapted from USDA PLANTS Database, Invasive Plant Atlas of the
United States, and selected references.)
6 ft. (1.8 m) tall. The hollow,
ridged stems are usually less than 2 in. (5 cm) in diameter at the base and lack reddish pur-
ple spots. The palmately compound leaves, very pubescent on the undersides, are deeply
incised but only 1 ft. (0.3 m) wide. The white flower umbels are rounded on top and 6–
12 in. (15–30 cm) wide. Purplestem angelica grows 4–6 ft. (1.2–1.8 m) tall, with a smooth,
hairless purple stem and doubly compound leaves that are not dissected. Its white flowered
inflorescence is round, not flat-topped, and 8–12 in. (20–30 cm) wide. Wild parsnip is 2–5
ft. (0.6–1.5 m) tall, with a yellow-green ridged stem. Yellow flowers are in flat umbels, 4–6
in. (10–15 cm) wide, and the yellow-green compound leaves have 5–11 leaflets. The watery
sap from wild parsnip may cause a photosensitive reaction.
Two alien species in the carrot family, both widely naturalized, may also appear similar.
Queen Anne’s lace is small, 1–3 ft. (0.3–1 m) tall, with thin hairy stems. Flowers grow in a
flat umbel, 3–6 in. (7.5–15 cm) wide, and the 6 in. (15 cm) long leaves are finely dissected.
374 n FORBS

A. Giant hogweed can grow 2–3 times the height of a person. (Terry English, USDA APHIS PPQ, Bugwood.org.)
B. The inflorescence is a large umbel. (Terry English, USDA APHIS PPQ, Bugwood.org.) C. Large stalks are
hollow. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology, University of Connecticut, Bugwood.org.) D.
Large leaves are deeply lobed. (Donna R. Ellis, University of Connecticut, Bugwood.org.) E. Dried fruit, each
with one seed, have prominent resin canals. (USDA APHIS PPQ Archive, USDA APHIS PPQ, Bugwood.org.)

The smooth, purple stems of poison hemlock reach 4–7 ft. (1.2–2 m) tall. The flowers
are half-round umbels, 2–8 in. (5–20 cm) in diameter, and the compound leaves have fine,
linear leaflets. Common elderberry, in the honeysuckle family and native to the United
States, is a shrub or tree 2–10 ft. (0.6–3 m) tall with a woody gray-brown stem. White
inflorescences are flat, 4–6 in. (10–15 cm) wide, and compound leaves have 5–7 leaflets,
each 2–6 in. (5–15 cm) long.
Introduction History. Because of its unique size, giant hogweed was probably brought to
the United States as an ornamental. In 1917, it was found growing in a garden in New York,
and it was reported in Canada around 1950. The dried fruit is used as a spice, called golpar,
in Iranian cooking, and seeds are occasionally sold in ethnic food stores.
Habitat. Giant hogweed is most common on disturbed sites, such as vacant lots, unculti-
vated fields, and along roads and railroads, but may also invade intact communities such as
grasslands. It thrives in rich, moist soils in riparian zones along rivers and streams, where
water easily distributes seeds. It also occupies ravines and ditches. Giant hogweed grows
best in sunny locations and can be overgrown by native species in shady sites. When planted
as an ornamental, it may escape to vacant lots or open space between residential
communities.
Reproduction and Dispersal. Giant hogweed reproduces by seed. Although it takes sev-
eral years before plants grow flower stalks, one plant can produce 27,000–50,000 seeds.
Most seeds fall near the parent, and natural dispersal is primarily by wind and water. The
fruit is able to float for three days and may be carried as far as 6 mi. (10 km) distant. Seeds
are also dispersed by birds that eat the fruit. Human activity also aids in the spread of giant
hogweed. Seeds may escape from gardens or be contaminants in topsoil deposited along
right-of-ways. Dried inflorescences are frequently used in decorative flower arrangements.
Because they require a period of cold weather, seeds will not germinate in the summer they
are produced, and seeds are viable for more than seven years. Germination begins in early
 GIANT HOGWEED n 375

spring and continues all summer, but the plant requires moisture to become established.
Although plants usually die after flowering, they may offset, producing new plants from
their root crowns.
Impacts. The plant’s clear watery sap contains glucoside phototoxins, which react with
sunlight. Brushing against the bristles or against broken stems or leaves will deposit the toxic
sap onto the skin. The result is severe dermatitis, making the plant a serious health hazard in
urban and suburban areas. In the 1970s, several children in Great Britain were affected after
using the hollow stems as toys. Direct contact with the sap increases the skin’s sensitivity to
the ultraviolet radiation in sunlight, often leading to severe burns that may require months
to heal. Symptoms appear 15–20 hours after contact with sap and exposure to sunlight.
Affected skin may redden and swell, often developing blisters, which break open to infection
and frequently cause scarring. Contact of the sap with the eyes can cause either temporary or
permanent blindness. It is advised to avoid breathing smoke from fires that contain giant
hogweed fuel. Even birds and other animals have been reported to have experienced skin
injuries from contact with the plant.
Due to its abundant seed production, giant hogweed colonizes rapidly. Its larger leaves
create a dense canopy that suppresses native riparian species, negatively affecting associated
wildlife. Because the large plants die back to the ground in winter, the resulting exposure of
bare soil may increase soil erosion along stream banks and on steep slopes. Other soil
changes include a potential increase in nutrients because of the large amount of decaying
biomass each year. Economic impacts include not only money spent to control infestations,
but also funds spent for medical care.
Management. Because of varying leaf morphology, plants in the rosette stage may go
unnoticed, enabling the plant to grow and spread. After establishment, infestations may
require five years to eradicate. Control measures should be repeated and sites frequently
monitored, with the goal of both depleting root reserves and eliminating the seed bank.
Anyone working with or around giant hogweed should avoid direct contact with the plant
by wearing protective clothing. Work done after sunset will minimize the possibility of sun-
burns. If contact occurs, the skin should be covered to reduce sunlight exposure and
immediately washed with soap and cold water.
Physical control is the most commonly used method. If nothing else can be done,
removal of flowers will prevent seed development. Plants can be dug out or cut below
ground level. Although difficult to do, taproots should also be removed because root buds
will sprout. All seeds should be destroyed by burning or taken to a landfill. Mechanical
brush cutters can be used on large populations, just after the peak of flowering but before
seed sets. Resprouts can be cut a few weeks later. Plants that are cut after seeds mature, how-
ever, will not resprout. Although mowing stimulates the buds on the root stalk, repeated
mowing may exhaust the plant’s nutrient reserves. Grazing by cattle and pigs, which suffer
no ill effects from eating the foliage, may stress the plant and limit flowering. Trampling also
damages plants.
Effective chemical control of giant hogweed can be achieved by applications of glyphosate
or triclopyr, which will be absorbed into the roots. The best time for application on first-year
plants is before they reach 2 ft. (0.6 m) tall or in the fall, after the first frost. Other herbicides,
including 2,4-D, TBA, MCPA, and dicamba, kill the aerial parts of the plant but not the
roots.
No biological control is known. Although the toxic sap may be a deterrent to most
insects, the caterpillars of some butterflies do feed on it. Several insect species found on giant
hogweed in Switzerland are being investigated.
376 n FORBS

Selected References
“Giant Hogweed.” Environment and Natural Resources Institute. Alaska Natural Heritage Program.
University of Alaska, Anchorage, 2005. http://akweeds.uaa.alaska.edu/pdfs/potential_species/bios/
Species_bios_HERMAN.pdf.
“Giant Hogweed—Heracleum mantegazzianum.” Agricultural Program, Cornell University Cooperative
Extension, Wyoming County, NY, n.d. http://counties.cce.cornell.edu/wyoming/agriculture/
resources/ipd/giant_hogweed.htm.
Marrison, David L., and David J. Goerig. “Giant Hogweed.” Horticulture and Crop Science, Ohio State
University Extension Fact Sheet. Columbus, n.d. http://ohioline.osu.edu/anr-fact/hogweed.html.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Heracleum mantegazzianum.” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=418&fr=1&sts.
Nice, Glenn, Bill Johnson, and Tom Bauman. “The Infamous Giant Hogweed.” Purdue Extension Weed
Science. Purdue University, West Lafayette, IN, 2004. http://www.btny.purdue.edu/weedscience/
2004/articles/gianthogweed04.pdf.

n Goutweed
Also known as: Snow-on-the-mountain, bishop’s weed, bishop’s goutweed, ground elder
Scientific name: Aegopodium podagraria
Synonyms: None
Family: Carrot (Apiaceae)
Native Range. Most of Europe and northern Asia to eastern Siberia.
Distribution in the United States. Eastern half of the United States, from Maine south to
Georgia and west to Minnesota and Missouri. Also the Northwest coast, Washington and
Oregon, east to Idaho and Montana. It is either absent or uncommon in the drier western
states.
Description. Goutweed is an herbaceous perennial commonly used as a groundcover.
Although sometimes called a vine, it does not have a climbing or twining habit. It grows as
a mat, but with leaves and flowers held upright. Leaves are basal, meaning that the leaf stalks
are attached directly to the rhizomes in the soil. The hollow stems holding the leaves grow
1 ft. (0.3 m) tall. The compound leaves are triternate, in three groups of three leaflets. The
lower leaflets may be fused together, reducing the number of leaflets in the compound leaf.
Leaflets are oval with a pointed tip and toothed or lobed edges, with no pattern to the teeth
or the lobes. Leaves are medium green or pale gray-green, with each leaflet 1–3 in. (3–8 cm)
long. The variegated form has bluish-green leaves with a white, creamy edge. Patches may be
mixed because the variegated form often reverts to green. Goutweed is deciduous. After yel-
lowing in August, leaves drop off the stem following the first frost in autumn.
The root system is shallow, but extensive, with both main and lateral roots. The rhizomes
are long, white, and branching, forming roots at each node. Horizontal rhizomes may
develop vertical shoots at the end of the growing season. Rhizomes can be as short as 2 in.
(5 cm) or as long as 9.8 ft. (3 m), but only 0.08 in. (2 mm) in diameter.
Flower stems, reaching 2–3 ft. (0.6–0.9 m) tall, appear from May through August,
depending on geographic location. The leafy stalks support flat-topped, compound umbels
of small, five-petaled flowers. Flower clusters are 2–4.5 in. (6–12 cm) in diameter. Leaves,
however, are much more numerous than flower stalks, especially in deep shade, and the
white flowers are almost camouflaged by the white leaf variegation. Tiny brown, elongate
seeds, resembling carrot seeds, ripen in late summer.
 GOUTWEED n 377

Related or Similar Species.

Aegopodium podagraria ‘Varie-
gatum’ is the variegated variety,
with no difference in habitat or
invasive characteristics except
that it may be less aggressive
than the green form.
Introduction History. Bro-
ught to the United States by
European immigrants as an orna-
mental, goutweed was well estab-
lished by 1863. The plant’s name
stems from the once-held belief
that it could be used to cure gout.
The leaves were used as a salad
ingredient in parts of Russia.
Goutweed was uncommon in
New England in the early 1980s
and is currently expanding its
range, especially in Vermont and
Massachusetts. The plant is still
readily available in nurseries and
is one of the most common
deciduous groundcovers in
northern U.S. climates.
Habitat. Goutweed’s natural
history in North America has
not been well studied, but its
habitat preferences can be
extrapolated from its native
range. Its habitat in Europe is
primarily deciduous forests, Goutweed colonizes disturbed sites and deciduous forests, primarily in
especially riparian sites, but it is the eastern states. (Native range adapted from USDA GRIN and selected
also found in shrublands, wet- references. Introduced range adapted from USDA PLANTS Database,
lands, and grasslands. Although Invasive Plant Atlas of the United States, and selected references.)
most often found on disturbed
sites in North America, such as logged forest areas, abandoned fields, and pastures, it can also
invade closed-canopy forests. In Europe, goutweed grows under the canopy of several types of
deciduous forest trees, including ash, oak, beech, maple, and elm. It also grows in mixed for-
ests where spruce, aspen, and linden are common, and is found beneath willows, birch, and
alder. These same genera dominate the northern forests of the United States, which are suscep-
tible to invasion. Goutweed is tolerant of poor soils and a range of soil acidity. It grows best on
moist, well-drained soils, but may also tolerate saturated soils on floodplains. Lack of sufficient
soil moisture may be a limiting factor in its extent. Goutweed grows well in part sun to full
shade. Leaves will scorch if the site is hot, sunny, and dry. In Europe, it grows where annual
precipitation is 19.5–32.8 in. (495–832 mm), and at elevations up to 3,488 ft. (1,063 m).
Reproduction and Dispersal. Although goutweed is capable of producing viable seed, it
spreads primarily by vegetative means, growing new stems from rhizomes and root
378 n FORBS

A. Flower stems may be held high above the foliage. (Richard Old, XID Services, Inc., Bugwood.org.) B. Leaflets
are in sets of three. (Richard Old, XID Services, Inc., Bugwood.org.) C. Leaves of the variegated form have white
edges. (Richard Old, XID Services, Inc., Bugwood.org.) D. Plants have an extensive rhizome system. (Leslie J.
Mehrhoff, Ecology and Evolutionary Biology University of Connecticut, Bugwood.org.) E. Flower clusters are
flat-topped umbels. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology University of Connecticut,
Bugwood.org.)

segments. Established plants can expand by rhizome extension into both sunny and shady
areas, and a clump can increase its diameter by more than 2 ft. (0.6 m) a year. Patterns of
invasion are often clumped due to growth by rhizomes. Goutweed invades new areas by
intentional transplants and by improper disposal of yard waste containing rhizomes.
The fragrant flowers are pollinated by many insects, including beetles, bees, and small
flies. Seeds have no specific dispersal mechanism and simply fall from the plant. They
require a cold period and germinate the following spring, but do not seem to remain in
the soil as a seed bank. Seedlings need disturbed sites and bright light and may not survive
shady locations. Plants, however, rarely produce seeds, and seedlings are rare.
Impacts. Most deciduous forests in the eastern United States are vulnerable to invasion by
goutweed. Because goutweed is widely planted as an ornamental, infestations usually come
from abandoned or poorly kept gardens. Moist forests near old residential neighborhoods or
farmsteads are especially susceptible. Goutweed is aggressive, dominating the herb layer in
forests and displacing native plants, which reduces species diversity. By forming a dense
groundcover, it inhibits the germination and subsequent establishment of native tree spe-
cies. Direct effects on wildlife are not known, but deer seem to avoid grazing it.
Management. Because goutweed is limited in its extent, it may be possible to eradicate it
before it spreads into new territory. Any site where it has been eradicated or controlled
should be closely monitored for a few years to ensure that regrowth from rhizomes can be
eliminated as it develops. Once goutweed is controlled, the area should be reseeded with
native plants or noninvasive ornamentals, depending on the site, to avoid other invasive spe-
cies from taking advantage of the bare or disturbed site. Reseeding or plantings may also be
important to prevent soil erosion on the bare site. If used as a garden plant, goutweed should
be planted in a restricted area—for example, confined by concrete, such as a sidewalk or
foundation, to prevent unwanted spread by rhizomes.
Physical means of control will rarely kill the plant. Digging it up breaks apart the root sys-
tem and invigorates the plant, causing resprouting. Hand-pulling may be adequate for small
patches, but all the rhizomes and stolons must be removed. Plant debris should be left to
dry, then bagged and disposed of because any piece of root will grow. Frequent mowing
may keep goutweed from spreading along roadsides or into gardens and lawns. Covering
 HALOGETON n 379

plants with black plastic when leaves begin to emerge in spring will prevent the plant from
photosynthesizing and deplete its nutrient supply. Plastic should be left on all summer. An
alternative method is to mow the leaves before covering with plastic. The plants, therefore,
have used some of their reserves in growth but have no way of obtaining more. Covering
plants in mid- to late-summer is ineffective because the roots have already acquired nutrient
reserves.
Chemical control requires a systemic herbicide, such as glyphosate. Repeated applica-
tions may be necessary to deplete the nutrient reserves. Contact herbicides only kill the
leaves, and the plant resprouts from its extensive rhizome system within two months.
Only new leaves can be killed by contact herbicides. By late summer, the cuticle on the
old leaves is too thick to allow the herbicide to penetrate. Mowing in late summer, however,
will stimulate new growth, which can then be sprayed.
No biological control is available, because goutweed has no significant diseases or pests.

Selected References
“Aegopodium podagraria ‘Variegatum.’ ” Ohio State University, n.d. http://hcs.osu.edu/hcs/TMI/
Plantlist/ae_raria.html.
Garske, Steve, and David Schimpf. “Goutweed.” Weeds Gone Wild: Alien Plant Invaders of Natural
Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2005. http://www.nps.gov/
plants/alien/fact/aepo1.htm.
“Goutweed Aegopodium podagraria L.” Weed of the Week. U.S. Department of Agriculture, Forest
Service, Invasive Plants, 2006. http://www.na.fs.fed.us/fhp/invasive_plants.
Waggy, Melissa A. “Aegopodium podagraria.” In Fire Effects Information System. U.S. Department of
Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, 2010.
http://www.fs.fed.us/database/feis/.

n Halogeton
Also known as: Saltlover
Scientific name: Halogeton glomeratus
Synonyms: Anabasis glomerata
Family: Goosefoot (Chenopodiaceae)
Native Range. Deserts and grasslands of southern Asia, including Kazakhstan,
Uzbekistan, Turkmenistan, Afghanistan, and Pakistan, and eastward into southern Russia,
Mongolia, and western China.
Distribution in the United States. The northern Great Plains, from Nebraska to the
Dakotas, and all states west of the Rocky Mountains.
Description. Halogeton is an annual herbaceous plant that is semi-succulent. Plants vary
in size, usually 3–18 in. (7.5–45 cm) tall and equally as wide. Several stems emerge from the
base of the plant, spreading outward before bending to grow vertically. These main stems
have many secondary branches, which are often tinged with red or purple. Stems grow only
a few inches tall where stands are dense, but they can be 2 ft. (0.6 m) tall where plants are
widely spaced. The fleshy, succulent leaves are alternate and sessile on the stem. Leaves
are cylindrical, smooth and short, 0.2–0.9 in. (0.4–2.2 cm) long. At the end of the blunt leaf
tip is a conspicuous but delicate needlelike spine. The foliage is glabrous, but leaf axils have
tufts of long, cotton-like interwoven hairs. Plants have a dull bluish green tone in spring and
early summer, but change color later in the season. By late summer, stems may turn pink or
380 n FORBS

reddish, or both stems and

leaves may become pink, red,
salmon, or yellow. Leaves are
deciduous or dried by the time
seeds mature.
Plants slowly grow a taproot
as deep as 20 in. (50 cm), and
lateral roots radiate as far as 18
in. (45 cm) in all directions
Beginning in June and con-
tinuing through September,
dense clusters of small flowers
form in most of the leaf axils.
Halogeton has two types of flo-
wers, which produce two differ-
ent kinds of seeds. Neither
flower type has petals. The ma-
jority of flowers, 0.08–0.12 in.
(2–3 mm) in diameter, have five
fan-shaped, membranous or
transparent sepals, narrow at
the base and wider at the top.
The fan-shaped sepals are
greenish yellow, sometimes tin-
ged with red. Smaller flowers
have brown, toothlike sepals
that look like bracts. Seeds
mature from July to October.
Winged dark-brown or black
seeds are produced in late
summer. The black seeds are
Halogeton is toxic to livestock on western rangeland. (Native range enclosed by the fan-shaped
adapted from USDA GRIN and selected references. Introduced range sepals, in clusters dense enough
adapted from USDA PLANTS Database, Invasive Plant Atlas of the to cover and hide the stem.
United States, and selected references.) Wingless light tan or brown
seeds mature in early summer.
They are completely enclosed by the brown toothlike sepals. Plants that germinate
and mature late in the season produce black seeds only.
Related or Similar Species. Young plants may resemble Russian thistle, another invasive
species in arid regions of the western United States (see Forbs, Prickly Russian Thistle), but
Russian thistle leaves are linear. It also does not have white hairs in the leaf axils, and its seeds
are cone-shaped. Burning bush, also known as summer cypress or kochia, is an annual bushy
forb growing 1–7 ft. (0.3–2.1 m) tall. Its taproot and resistance to insects and disease make it
desirable as a forage crop for livestock. Dark green when young, plants became red as they
mature. Its pubescent leaves and lack of a stiff spine at the tip differentiate it from halogeton.
Introduction History. Halogeton was introduced to northern Nevada in the early 1930s,
most likely as a contaminant in crop seed. The first herbarium specimen was collected in
Nevada in 1934.
 HALOGETON n 381

A. Plants are usually less than 1.5 ft. (0.5 m) tall. (Joseph M. DiTomaso, University of California–Davis,
Bugwood.org.) B. Small succulent leaves end in a short spine. (Bonnie Million, BLM, Ely District,
Bugwood.org.) C. What appear to be petals are membranous sepals. (Clinton Shock, Oregon State University,
Bugwood.org.)

Habitat. Halogeton is called saltlover because it does best in alkaline and saline soils, pH
of 8–9 with a salt content of at least 5,800 ppm. It grows in the semiarid and high-desert
regions of the West that receive 5–13 in. (127–330 mm) of annual precipitation, approxi-
mately 60–70 percent of which falls as snow. It can be found on any soil texture, from clay
to loamy sands, which may or may not have a hard pan of calcium carbonate. The plant
commonly grows around dry lakebeds or alkaline flats, at 2,500–7,000 ft. (750–2,150 m)
elevation. It primarily invades disturbed sites in saltgrass, salt desert shrub, big sagebrush,
desert grasslands, and pinyon and juniper woodland ecosystems. It often grows in associa-
tion with other alien weeds, such as cheatgrass, clasping pepperweed, and Russian thistle.
It also grows with native shrubs, such as greasewood and shadscale saltbush, and with
native grasses, such as wheatgrass, needlegrass, Indian ricegrass, squirreltail, and galleta
grass. It is typically found in areas associated with livestock, such as stock trails, bedding
or feeding areas, watering sites, and overgrazed land. Halogeton is also common in
human-disturbed areas, such as roadsides, gravel pits, airstrips, railroad right-of-ways, and
abandoned farms, mines, and townsites. Numbers of plants and severity of infestations
depend on sporadic precipitation, which triggers germination.
Reproduction and Dispersal. Halogeton reproduces only by seed and is a prolific pro-
ducer. One inch of stem can produce 75 seeds (35 per cm), and one large plant can produce
more than 100,000 seeds. An infestation may produce 200–400 lbs. of seed per acre (222–
449 kg per ha). Seeds have several means of dispersal. The winged black seeds are easily
wind-dispersed. Plants also break at the base of the stem and become tumbleweeds, pieces
of which can be transported long distances when caught on vehicles. Road equipment, such
as graders, also distributes stems and seeds. Dust devils can carry dry stems and seeds as far
as 2 mi. (3.2 km). Seeds are resistant to animal intestinal tracts and are often deposited in
manure. Two types of seeds with different germination requirements ensure regeneration
of the plant. The black seeds drop off the plant by early November, have no dormancy
requirement, and germinate in the first growing season after maturing. The brown seeds stay
on the dried plant until January or February and remain viable in the soil seed bank for 10 or
more years. They require a three-month dormant period of cold temperatures, slightly above
freezing, before germination. Both types of seeds contain a coiled embryo. Either type germi-
nates quickly after a rainfall, from February through mid-August, with a peak in April.
382 n FORBS

Impacts. Although halogeton is not a competitive plant, it infests millions of acres of dis-
turbed or overgrazed land in the western states. Once established, it can prevent regrowth of
desirable native plant species because it alters the chemistry of the soil. Halogeton plants
take salt from deeper layers in the soil and deposit it on the soil surface when they die.
Soils become more alkaline and less permeable to water, which causes more evaporation.
The salts also inhibit microorganism activity.
Halogeton was first recognized as toxic in 1942. As a halophyte, the plant manufactures
large amounts of oxalic acids due to its excessive uptake of sodium ions. The large quantities
of the oxalate in the leaves and stems are poisonous to livestock. Usually unpalatable
because of its bitter taste, hungry or thirsty livestock will eat the plant if nothing else is avail-
able. It is especially toxic to sheep, and thousands of animals have died after ingesting the
plant. Signs of poisoning, which may occur as little as two hours after eating 12–18 oz.
(340–510 g) of the plant, include weakness, staggering, rapid and shallow breathing, drool-
ing, muscle spasms, and eventually coma and death. No treatment is available. The amount
of oxalates varies by season and part of the plant, but is more concentrated in actively grow-
ing plants. Sheep may become adapted to the oxalates if they are slowly given gradually
increasing amounts. The oxalates cause calcium to be lost from the blood, and sheep are fre-
quently fed calcium-enriched feed to counteract the loss.
Management. The best management is prevention, through maintenance of vigorous stands
of pasture and range grasses and adoption of good grazing practices. All soil disturbances should
be held to a minimum, including road construction and off-road vehicle use. To prevent poison-
ing, livestock should be kept away from infestations, especially after precipitation occurs
because the moisture will trigger germination from the seed bank. It is possible to eradicate
small, isolated patches. Sites should be monitored once or twice a year so seedlings may be killed
before they have a chance to produce seed. This should be done for at least 10 years to exhaust
the seed bank. For extensive stands, the cause of soil disturbance must be eliminated or reduced
and the site replanted with grasses, forbs, or shrubs that can compete with halogeton.
Physical removal of small plants or seedlings can be accomplished by hand-pulling. If
flowering has begun, plants must be removed from the site to ensure that no seed develops.
Mowing close to the ground will reduce, but not eliminate, seed production because short
branches missed by mowing will flower. Prescribed burns are a poor choice because halo-
geton is one of the first plants to regrow after a fire. Although the fire may destroy existing
plants and the seed bank, new seeds will be recruited from elsewhere to the now-exposed
site. Seeding or encouraging perennials that can compete with halogeton may decrease its
dominance. Although most wheatgrass species do not grow well on salty soils, a hybrid, such
as A. desertorum cv ‘Hycrest,’ is adapted to saline conditions and can eliminate halogeton.
Chemical applications of glyphosate can be used for spot treatment of small patches
before flowering occurs, and repeated throughout the summer as new seedlings emerge.
Metsulfuron is a better choice for more extensive infestations in pastures and rangeland

A Russian Spy Plot?

D uring the Cold War, it was speculated that Russian spy planes introduced halo-
geton as a biological weapon.
 ICE PLANT AND CRYSTALLINE ICE PLANT n 383

because it does not harm grasses. In field tests, dicamba and picloram were much less effec-
tive. Spraying must be repeated for 6–10 years to exhaust the seed bank.
Although potential insects have been identified in Central Asia, biological control is not
yet available. A stem-boring moth (Coleophora parthenica) from Pakistan was introduced to
the United States, but it failed to become established.

Selected References
Dewey, Steve. “Halogeton glomeratus.” In Invasive Plants of California’s Wildland, edited by Carla C.
Bossard, John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000.
California Invasive Plant Council, 2006–2010. http://www.cal-ipc.org/ip/management/ipcw/pages/
detailreport.cfm@usernumber=53&surveynumber=182.php.
“Halogeton (Halogeton glomeratus).” Plant Pest and Health Prevention Services (PHHPS), California
Department of Food and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/
halogeton.htm.
Pavek, Diane S. “Halogeton glomeratus.” In Fire Effects Information System. U.S. Department of
Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, 1992.
http://www.fs.fed.us/database/feis/plants/forb/halglo/all.html.

n Ice Plant
Also known as: Highway ice plant, pigface, sea fig, Hottentot fig
Scientific name: Carpobrotus edulis
Synonyms: Mesembryanthemum edule
Family: Stone Plant (Aizoaceae)

n Crystalline Ice Plant

Also known as: Common ice plant, ice plant
Scientific name: Mesembryanthemum crystallinum
Synonyms: Cryophytum crystallinum
Family: Stone Plant (Aizoaceae)
Native Range. Coastal areas of southwest Africa.
Distribution in the United States. Ice plant grows in California, from Eureka south into
Baja California, and in Florida. Crystalline ice plant is in California, Arizona, and Pennsylvania.
Description. Ice plant: Ice plant is a large, creeping, mat-forming perennial herb, which is
sometimes referred to as a shrub. Stems are 0.3–0.5 in. (8–13 mm) in diameter and as long
as 6.5 ft. (2 m). Stems become woody after several years growth. The very succulent leaves
are arranged opposite on the stem and are much longer, 1.6–4 in. (4–10 cm), than they
are wide, 0.2–0.5 in. (5–12 mm). They are either straight or slightly curved, triangular in
cross section, with one edge of the triangle finely serrated. Large water-filled cells in the
leaves often give them a shiny appearance. Leaf color ranges from dull green to bright green,
with red, orange, or purple margins. More sun exposure increases coloring in the leaves.
Flowering occurs all year, but the bloom peak is in late spring or early summer. The large
flowers, 2.5–6 in. (6.5–15 cm) in diameter, are terminal and solitary, rising from 0.4–0.8 in.
(1–2 cm) long pedicels. Dozens of narrow linear petals, pink, yellow, or yellowish white and
1.2–1.6 in. (3–4 cm) long, radiate from a center filled with many yellowish stamens. Flowers
become pinker as they age. The edible fruit is a fleshy sphere or oval, 1–1.2 in. (2.5–3 cm)
384 n FORBS

long, initially green but ripen-

ing to purplish red. They do
not split at maturity but remain
on the plant for several months.
Each fruit contains numerous
tiny black seeds.
Crystalline Ice Plant: Although
crystalline ice plant has different
morphological characteristics, it
causes many similar problems
as an invasive. It is found from
San Francisco Bay south to Baja,
including the Channel Islands
and as far as 8 mi. (13 km)
inland. A low-growing, annual
or biennial herb, it spreads over
the ground on trailing stems.
The succulent stems and leaves
are green to reddish. Stems
branch frequently and are usu-
ally less than 3.3 ft. (1 m) long.
Leaves, growing 0.8–8 in.
(2–20 cm) long on short
petioles, are oval- to spoon-
shaped, with wavy margins.
Leaves, stems, and fruit are
covered with crystal-like projec-
tions that resemble tiny water
bubbles. The majority of flowers
bloom from March through
June, but because germination
Both species of ice plant are considered invasive in coastal California. takes place all year, some flowers
(Native range adapted from USDA GRIN and selected references. can be found all year as well. The
Introduced range adapted from USDA PLANTS Database, Invasive Plant flowers are small, 0.3–0.4 in. (7–
Atlas of the United States, and selected references.) 10 mm), and stalkless, with
many narrow white petals radiat-
ing from the center. Petal color changes to pink as the flowers age. Flowers have numerous sta-
mens. After the fruits ripen in June through August, the plants dry from the base upward and
die. Dried fruits burst to release many seeds. Germination begins with the first fall rains and
fog and continues all winter. Seedlings grow rapidly until spring, but stop vegetative growth
during the hot, dry summer. All parts of the plants are edible.
Related or Similar Species. The genus Carpobrotus contains about 30 species, most of
which are called ice plant. The common name ice plant refers to many other plants as well,
such as Delosperma cooperi and Lampranthus spp.
Sea fig, also called Chilean iceplant although it is native to South Africa, is a close relative
that is smaller and less aggressive. Its magenta flowers are 1.5–2.5 in. (3.8–6.4 cm) in diam-
eter. The two species of Carpobrotus hybridize naturally along the California coast, resulting
in pink hybrid flowers intermediate in size. Hybrid plants are invasive.
 ICE PLANT AND CRYSTALLINE ICE PLANT n 385

A. Ice plant mats can cover large areas. (Richard Old, XID Services, Inc.) B. The large ice plant flowers have many
narrow petals. (Joseph M. DiTomaso, University of California–Davis, Bugwood.org.) C. Flowers of crystalline ice
plant are smaller, and the oval leaves have wavy margins. (Joseph M. DiTomaso, University of California–Davis,
Bugwood.org.) D. The long, succulent ice plant leaves are triangular. (Richard Old, XID Services, Inc.)
E. Crystalline ice plant leaves are covered with tiny bubble-like projections. (Joseph M. DiTomaso, University
of California–Davis, Bugwood.org.)

Introduction History. Ice plant was intentionally introduced to California in the early
1900s and used to stabilize soil along railroad tracks. It was subsequently planted along
highways for the same purpose until the 1970s. Thousands of acres were planted in
California. It is still promoted and sold in nurseries as an attractive garden plant, particularly
along the coast due to its tolerance of salt.
Crystalline ice plant was probably brought to California in the 1500s, in sand used for
ship ballast. The California Department of Transportation used it for erosion control and
landscaping in the 1950s, but stopped in 1969 when it was realized that it failed to stabilize
soil on steep slopes and died during a hard freeze.
Habitat. Ice plant tolerates a range of environmental conditions, such as soil moisture and
nutrient content. It grows predominantly in coastal habitats, including dune scrub, prairie,
bluffs, salt marsh, and maritime chaparral. Because it is intolerant of frost, it does not extend
far inland or above 500 ft. (150 m) elevation. The plant’s invasive ability varies with the type
and health of native plant communities. Seeds are most easily established on disturbed sites.
Rodent disturbances in coastal prairie, for example, open bare soil patches. In a healthy,
undisturbed community, the native prairie grasses can outcompete ice plant seedlings.
Crystalline ice plant grows mostly on saline coastal soils, such as beaches, sage scrub,
grasslands, estuaries, salt marshes, and bluffs. It easily spreads onto bare soil and disturbed
sites. Grazing of coastal grasslands and continued natural erosion of coastal bluffs provide
disturbed ground that encourages its spread. Its intolerance of frost limits its distribution.
Reproduction and Dispersal. Ice plant reproduces and spreads both sexually and vegeta-
tively. Each plant produces an abundance of fruit, each containing hundreds of seeds. The
fruit are eaten by several mammals, including deer, rabbits, and rodents, all of which are sig-
nificant in dispersal of seeds, because scarification in a digestive tract aids germination. Long
trailing stems branch frequently and grow shallow, fibrous roots at every node in contact
with the soil. All stems and shoots grow roots, and continue to grow when separated from
the parent plant. Vegetative growth is extensive.
386 n FORBS

Rabbits and mice eat the fruit and disperse the seeds of crystalline ice plant. It does not
reproduce vegetatively.
Impacts. Ice plant is invasive in Mediterranean climates similar to its native range, especially
in California and Australia, where it often forms monospecific stands. It lowers biodiversity by
either directly competing with native plants, both seedlings and mature shrubs, or by altering
environmental conditions. Ice plant makes soils more acidic, and buildup of organic matter
in sandy soils or dunes allows other nonnative plants to also invade. It has the biggest impact
on native plants in drought years, and is quicker than natives to invade burned clearings.
Specific endangered or sensitive native plants threatened by ice plant include Wolf’s primrose,
pink sand verbena, and beach layia. Because ice plant has no dormant season, stems can
increase their length more than 3 ft. (1 m) each year, even though it is somewhat limited due
to grazing by rabbits. Stems grow over one another and form impenetrable mats blanketing
the ground up to 19.5 in. (50 cm) thick. Individual plants eventually coalesce to cover areas
as large as 165 ft. (50 m) in diameter. Because ice plant stabilizes coastal sand dunes more
quickly than native plants do, dune buildup is decreased. Areas inland from the dunes may
be subject to more wave erosion because of fewer large, protective dunes.
Crystalline ice plant outcompetes native species because it is better able to withdraw
moisture from the soil. Salts that accumulate in plant tissues return to the soil at the death
of the plant, which increases the salt content on the surface beyond the tolerance of many
native grasses and forbs. Levels of soil nitrate are higher under crystalline ice plant, which
inhibits the growth of native grass seedlings.
Management. Hand-pulling is an effective physical way of controlling small plants or
populations, but it is expensive and labor intensive. Larger areas may be more easily
removed using tractors or other machinery. Because fragments of ice plant will grow roots
and sprout, all root and stem pieces must be removed from the soil. Reseeding with native
species or mulching the area may help to suppress growth of any remaining pieces.
Prescribed burning is not useful because of the high water content in both types of ice plant.
Control by grazing is limited because leaves are too salty and astringent and the stems too
fibrous and woody to be palatable to livestock.
Because herbicides have detrimental effects on nontarget plants, chemical control can
damage native plants. Foliar spraying is best done in winter when native plants are dormant,
but the evergreen ice plant is not. Although glyphosate will kill plants, sprouts may reoccur
for several months. Chlorflurenol, used along roadsides, is also effective.
No biological controls are available. Ice plant scale insects (Pulvinariella mesembryanthemi
and P. delottoi) do some damage, but not enough to affect growth. Parasitism by a dodder
(Cuscuta sp.) also fails to do enough harm.
Although no specific research has been conducted on eradication of crystalline ice plant,
some of the same methods may be applicable. Because crystalline ice plant is an annual,
however, efforts should target reducing seed production.

Selected References
Albert, Marc. “Carpobrotus edulis.” In Invasive Plants of California’s Wildlands, edited by Carla C.
Bossard, John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000.
http://www.cal-ipc.org/ip/management/ipcw/online.php.
Randall, John M. “Mesembryanthemum crystallinum.” In Invasive Plants of California’s Wildlands, edited
by Carla C. Bossard, John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California
Press, 2000. http://www.cal-ipc.org/ip/management/ipcw/online.php.
 JAPANESE KNOTWEED n 387

Whittemore, Frank. “Why Is the Ice Plant Bad?” Garden Guides, 1997. http://www.gardenguides.com
/88524-ice-plant-bad.html.

n Japanese Knotweed
Also known as: Japanese bamboo, crimson beauty, Mexican bamboo,
Japanese fleece flower, reynoutria
Scientific name: Fallopia japonica
Synonyms: Polygonum cuspidatum, Pleuropterus cuspidatus, Pleuropterus zuccarinii, Polygonum
zuccarinii, Reynoutria japonica
Family: Buckwheat (Polygonaceae)
Native Range. Eastern Asia,
including China, Taiwan,
Japan, and Korea.
Distribution in the United
States. Most of the lower 48
states and Alaska. Especially
prominent in the eastern half
of the country, from Wisconsin
east to Maine, south to Loui-
siana and Georgia, and west to
South Dakota and Oklahoma.
Stands are scattered in mid-
western and western states.
Description. Japanese knot-
weed is an herbaceous peren-
nial that appears woody
because of its upright stems that
grow 3–10 ft. (1–3 m) tall with
little side branching. Although
it superficially resembles bam-
boo because of its straight
growth of cane-like hollow
stems, it is not related and can
be distinguished by its leaf
shape. The round stems, which
are 1 in. (2.5 cm) or more in
diameter, are smooth and
sturdy. Swollen nodes, where
leaves are attached to the stem,
occur every 3–5 in. (7.5–
12.5 cm). Characteristic of all
members of the buckwheat
Japanese knotweed, which was used for erosion control, is more
family, the stems have a mem- prevalent in the eastern states. (Native range adapted from USDA GRIN
braneous sheath encircling the and selected references. Introduced range adapted from USDA PLANTS
nodes. In Japanese knotweed, Database, Invasive Plant Atlas of the United States, and selected
the sheath is often reduced to a references.)
388 n FORBS

A. Japanese knotweed can grow 10 ft. (3 m) tall. (Richard Old, XID Services, Inc., Bugwood.org.) B. Leaves have
smooth margins and a pointed tip. (James H. Miller, USDA Forest Service, Bugwood.org.) C. The rhizome system
is extensive. (Ohio State Weed Lab Archive, The Ohio State University, Bugwood.org.) D. Small flowers grow on
panicles at the ends of stems. (James H. Miller, USDA Forest Service, Bugwood.org.) E. Papery, winged fruits
contain small seeds. (Barbara Tokarska-Guzik, University of Silesia, Bugwood.org.) F. Swollen nodes line the
smooth stems. (James H. Miller, USDA Forest Service, Bugwood.org.)

red ring around the stem, which sometimes makes the stem appear reddish brown. The
broadly oval leaves vary in size, but are usually 6 in. (15 cm) long and 3–4 in. (7.5–
10 cm) wide, with smooth margins and a pointed tip. Leaves are alternate and grow on the
upper parts of the stems.
The extensive root system, with rhizomes as long as 65 ft. (20 m), enables Japanese knot-
weed to form thick colonies. Rhizomes extend as deep as 7 ft. (2 m) into the soil, where they
tap a wide source of water and nutrients. Each root crown produces 30–50 shoots, and indi-
vidual plants can be 8–15 ft. (2.4–4.5 m) in diameter. Large clumps consisting of several
plants can cover several hundred square feet or several acres and can dominate an entire
shoreline. Intolerant of freezing temperatures, above-ground parts of the plants quickly die
after frost. The bare, brown stalks remain upright all winter. Rhizomes survive and resprout
rapidly in spring, as early as April or as late as June, depending on geographic region. Young
sprouts resemble red asparagus. Because of root reserves, plants grow fast, as much as 1 ft.
(0.3 m) each week, and if they resprout early in spring, they can reach 10 ft. (3 m) by June.
In late summer to early fall, August to September, greenish-white or cream-colored
flowers appear in panicles, or branched sprays near the ends of the stems. Flowers are tiny,
0.1 in. (3 mm), but the panicles are 3–6 in. (8–15 cm) long. Although all flowers have ves-
tigial parts of both sexes, male and female flowers grow on different plants. Male inflores-
cences are upright, while female inflorescences are drooping. Pollination is done by bees
and other insects, and fruit appears only two weeks after flowering. The fruit are small
(0.25–0.4 in.; 6–10 mm), papery, and winged, containing very small, shiny black seeds,
0.15 in. (4 mm) long.
Related or Similar Species. Giant knotweed and Bohemian knotweed, which is a hybrid
between giant knotweed and Japanese knotweed, look similar to Japanese knotweed except
for tiny hairs on the undersides of the leaves. Giant knotweed grows to 15 ft. (4.5 m) tall,
with heart-shaped leaves and greenish flowers. Hybrid seeds are fertile. Nomenclature of
knotweeds is in dispute, but the invasive effects are the same regardless of name.
Introduction History. Japanese knotweed was from brought from Japan to the United
Kingdom in 1825 as an ornamental, and then introduced to the United States from Great
 JAPANESE KNOTWEED n 389

Britain in the late 1800s. Used for erosion control and landscape screening because of its
dense network of rhizomes, it became invasive in natural habitats after escaping cultivation
in gardens. By 1894, it had naturalized near Philadelphia, Pennsylvania; Schenectady, New
York; and Atlantic Highlands, New Jersey. In 1910, it was advertised in seed catalogs, but by
1938, information was published on how to eradicate it from gardens.
Habitat. Japanese knotweed is tolerant of many conditions, including full shade, high
temperatures, drought, and floods. Its growth rate depends on soil quality and nutrients,
and the species is more common in regions with high rainfall. The species grows on a variety
of soils, including silt, loam, and sand, under saline, alkaline, or acid conditions. It thrives
along streams, rivers, and other low-lying areas, but also invades disturbed ground, such
as old homesites, abandoned farmland, ditches, roadsides, railroad tracks, and utility line
corridors. It needs damp soil to become established, but grows in dry sites as well. It grows
best in sun, and poorly in shade. In its native range, Japanese knotweed is the dominant pio-
neer on volcanic slopes and disturbed sites.
Reproduction and Dispersal. New plants arise from seed, broken stems, and rhizome
nodes. Although seeds are the normal means of reproduction in its native Japan, they are
rarely produced in the United States, because fertile males are rare and clones are usually
all male or all female. Seeds do germinate, however, when produced. Stands expand pre-
dominantly by vegetative means. Any fragment of the roots, rhizomes, or stems, as small as
0.5 in. (1.25 cm), will grow into a new plant. Even internode pieces will sprout.
Fragments are relocated by water, wind, mud, as contaminants in fill dirt, or as discarded
cuttings from urban gardens. Optimal sprouting takes place from rhizomes just below the
soil surface, but sprouts are capable of emerging from rhizomes buried as deep as 3.3 ft.
(1 m).
Impacts. Specific problem areas include hundreds of acres of wetlands in western
Pennsylvania, creeks in suburban Philadelphia, and Rock Creek National Park in
Washington, D.C. Because of its propensity to grow from rhizomes that can extend up to
65 ft. (20 m) away from the parent plant, Japanese knotweed forms dense, monoculture
thickets that exclude native vegetation and reduce biodiversity. Rapid growth in spring, over
3 in. (8 cm) per day, enables plants to shade out native species. Growing best in full sun-
light, it is less likely to invade undisturbed forest and shady areas. Japanese knotweed is
especially a threat to riparian areas, where it destroys habitat for fish and other wildlife
and limits recreational opportunities. Not only can it survive extensive flooding, but it rap-
idly colonizes bare ground left by floodwaters. It pulls nitrogen out of the soil but does not
return it in leaf fall, thereby altering the soil and aquatic nutrient exchanges. In the Pacific
Northwest, Japanese knotweed threatens salmon due to loss of insects and shady areas
because it outcompetes native plants. By providing shade and habitat for animals, native
trees are an important part of the natural riparian environment. Native plants also stabilize
the ground against erosion. Woody debris from native trees disrupts stream flow, resulting
in the creation of pools of water for salmon.
Japanese knotweed causes monetary loss because vigorous rhizome growth can break
foundations and road beds.
Management. It is probably not possible to eliminate Japanese knotweed everywhere.
Management should be selective, as funds and personnel permit. Treated sites should be
monitored frequently for regrowth. At minimum, areas 20 ft. (6 m) away from the original
plants should be examined for sprouts, which can then be pulled before they become estab-
lished. The origin of stands along the Hoh River in Washington State was traced upstream to
a single ornamental planting from which pieces escaped cultivation. Therefore, any
390 n FORBS

management along waterways must begin upstream, because broken pieces carried down-
stream will grow into new colonies.
Complete physical removal may only be possible with small plants or small infestations.
It is imperative that all parts of even a small plant be dug up and properly disposed of.
Removing even a small patch of Japanese knotweed may take as long as three years.
Digging plants is not practical, because the roots extend very deep and pieces break off.
Repeated cutting or mowing stands, before regrowth reaches 6 in. (15 cm), at least three
times a year for 2–3 years, will deplete root supplies and weaken the stand. Careless mow-
ing, however, may actually increase the size of the stand, because pieces will grow, and
one plant can quickly give rise to hundreds. Burning does not kill the plant, and the only
benefit is that it opens access to dense stands for other means of control. Covering cut or
mowed stands with black plastic will prevent plants from conducting photosynthesis,
thereby depriving them of nutrients. Covering should remain in place for at least one year
and be frequently checked for sprouts, both around the edges and through the material.
Japanese knotweed has been observed growing through 2 in. (5 cm) of asphalt and can
more easily penetrate even thick plastic. Grazing of young shoots by sheep, goats, cattle,
and horses will not kill plants but serves the same purpose as mowing, to weaken the
plant.
Chemical control is possible with repeated applications and is more effective combined
with cutting or mowing. Glyphosate or triclopyr may be applied to cut stumps within one
half-hour after cutting, before the wound begins to heal, or may be directly injected into
each stem. Because they are labor intensive, neither method is feasible for a large stand.
Picloram is a selective herbicide but cannot be used near water. Applications of herbicide
can be used in cold weather as long as the ground is not frozen. Systemics work best in
early fall, when plants are actively moving nutrients into the roots. Foliar applications
are effective on new growth or anytime the plant is in full leaf, but is best done before
flowering.
Using selections from a list of potential insects and pathogens from Japan, initial research
on biological control of Japanese knotweed has been conducted in the United Kingdom,
where the plant is also a serious invasive. Two insects, a leaf-chewing chrysomelid beetle
(Gallerucida bifasciata) and a sap-sucking psyllid (Aphalara itadori), were imported into
U.S. quarantine in 2007 for testing.

Beneficial Uses of Japanese Knotweed

W idely planted as an ornamental, Japanese knotweed may have other beneficial

uses. Dried roots of the plant are steeped for tea in Asia, and the plant shows
promise for medicinal use. It is a major source of resveratrol, the same substance that
is in red wine, which is often sold as a nutritional supplement. Experiments with mice
show that extracts from the plant may be anti-inflammatory, helping to prevent
ulcers or to promote healing of burns. Compounds from the roots also seem to be
beneficial in several ways. They inhibit growth of certain bacteria, may protect
against cancer, and prevent formation of blood clots. Roots, with their concentra-
tions of emodin, also sold as a nutritional supplement, are also used in Asia as a natu-
ral laxative.
 KAHILI GINGER n 391

Selected References
“Integrated Pest Management Profile, Japanese Knotweed.” Department of Ecology, State of
Washington, 2007. http://www.ecy.wa.gov/programs/wq/plants/weeds/aqua015.html
Mehrhoff, L. J., J. A. Silander Jr., S. A. Leicht, E. S. Mosher, and N. M. Tabak. “Japanese Knotweed.”
Invasive Plant Atlas of New England (IPANE), University of Connecticut, 2003. http://
www.ipane.org.
Nice, Glenn. “Japanese Knotweed, (Polygonum cuspidatum).” Extension Weed Science, Purdue
University, 2007. http://www.btny.purdue.edu/weedscience/.
Remaley, Tom. “Japanese Knotweed.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/ALIEN/
fact/faja1.htm.
Seiger, Leslie. “Element Stewardship Abstract, Polygonum cuspidatum.” Global Invasive Species Team,
Nature Conservancy, 1991. http://wiki.bugwood.org/Polygonum_cuspidatum.

n Kahili Ginger
Also known as: Awapuhi kahili, kahili, kahila garland lily, wild ginger
Scientific name: Hedychium gardnerianum
Synonyms: None
Family: Ginger (Zingiberaceae)
Native Range. Lower slopes, up to 4,100 ft. (1,250 m), of the Himalayas in Nepal and
eastern India.
Distribution in the United States. Hawai’i.
Description. Kahili ginger is a non-woody perennial herb with thick, branching rhi-
zomes. From May through July every year, buds on the rhizomes produce many new leafy
shoots, which can be as tall as 5–6.5 ft. (1.5–2 m). The large oblong to lance-shaped leaves,
8–20 in. (20–50 cm) long and 4–6 in. (10–15 cm) wide, are shiny and alternate. The upper
surface is glabrous, while the lower surface is sparsely pubescent. Tips are pointed, and
leaves have no petioles. The ligules, where the leaf attaches to the stem, are membraneous,
0.8–1.6 in. (2–4 cm) long, and pubescent, but leaf sheaths are glabrous.
The fleshy rhizomes, which are segmented into units 4 in. (10 cm) long and 1.5 in.
(4 cm) in diameter, grow in an intertwined, solid mat that can be 3.3 ft. (1 m) thick in the
soil. A distant relative of the ginger used in cooking, the rhizome interiors are pale, with a
faint ginger smell and taste.
Plants in Hawai’i flower in July and August. The erect inflorescence is a large spike, 10–
18 in. (25–45 cm) long, and cylinder shaped, with many lemon yellow zygomorphic flow-
ers. The corolla is a slender tube, 3–3.5 in. (8–9 cm) long. Yellow structures that resemble
petals, but are sterile stamens called staminodes, are 1.6–2 in. (4–5 cm) long. One long sta-
men, usually red but sometimes yellow, conspicuously extends beyond the yellow corolla.
Flowers are very fragrant. After the flowers drop off, the fruiting spike remains upright with
fleshy orange fruits, each 0.6–0.8 in. (1.5–2 cm) long. When mature, usually October
through December, the seed capsules open on the stalk to expose small, shiny red, fleshy
seeds, which become gray as they age.
Related or Similar Species. Hedychium species readily hybridize, confusing the taxonomy
of yellow ginger and white ginger. Yellow ginger, also known as cream lily, is native to
eastern India and Nepal at elevations of 1,650–2,600 ft. (500–800 m). A significant pest in
Hawai’i, it grows in forests, at forest edges, and along riverbanks. Stems grow 3.3–10 ft.
(1–3 m) tall. The plant’s elliptical to lance-shaped leaves, 8–20 in. (20–50 cm) long and
392 n FORBS

1.5–4 in. (4–10 cm) wide, are

more narrow than those of
kahili ginger and more verti-
cally oriented, pointing up.
Leaf tips are pointed, and leaf
sheaths are slightly pubescent.
The flower spike is smaller, 6–
8 in. (15–20 cm) long and 1.2–
2.4 in. (3–6 cm) wide, with
creamy to pale-yellow fragrant
flowers, sometimes reddish yel-
low at the base. Green bracts
below the corolla are distinctly
imbricate (overlapping). Yellow
ginger reproduces both by seed
and by rhizome buds.
White ginger, also known as
ginger-lily, is native to the tropi-
cal wet regions of southeastern
China and southern Asia, from
India east to Vietnam and south
to Malaysia and Indonesia.
Although it can be found in full
sun, it prefers partial shade,
humus-rich soils, and water-
logged, but not submerged, soils.
Its stems reach 3.3–10 ft. (1–3 m)
tall. Leaf blades are lance-shaped,
8–16 in. (20–40 cm) long and
1.8–3 in. (4.5–8 cm) wide, gla-
brous on top and thinly hairy
Kahili ginger grows best in wet or mesic environments in Hawai’i. (Native beneath. Elliptical flower spikes,
range adapted from USDA GRIN and selected references. Introduced 1.5–8 in. (10–20 cm) long and
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the 1.6–3 in. (4–8 cm) wide, have
United States, and selected references.) fragrant white flowers, some-
times tinged with yellow at the
base. Seeds are bright red. Precise correlation of seed production and elevation is not known,
but seeds do not develop at Kalopa State Recreation Area, on the Big Island, at 2,000 ft.
(600 m). In Hawai’i Volcanoes National Park, seeds are produced only below an elevation of
4,000 ft. (1,200 m). Because it produces fewer seeds, white ginger spreads primarily by rhizomes.
Introduction History. Kahili ginger was introduced to every island in the Hawaiian chain
as an ornamental by Chinese immigrants in the early 1900s. It has since escaped from culti-
vation. When first recorded in Hawai’i Volcanoes National Park, it was kept in check by feral
pigs (see Volume 1, Vertebrates, Mammals, Feral pig). Since the pigs were fenced out of the
park in the 1970s, the ginger now grows uncontrolled.
Habitat. Kahili ginger thrives in both bright light and in dense shade. Although it prefers
fertile soils and mesic to wet conditions, including areas with poor drainage, it can also grow
on soils with low fertility, in very little soil, or in thick humus. It even grows successfully in
 KAHILI GINGER n 393

A. New shoots grow from the dense mat of rhizomes. B. Both the inflorescences and leaves are large. C. Flowers
are zygomorphic. D. The fruit, fleshy at first, dries on the stalk. (Forest and Kim Starr, U.S. Geological Survey,
Bugwood.org.)

moss-covered forks of trees or on branches. Plants can be found in agricultural land, coast-
land, natural rainforests, montane forests, forest plantations, grasslands, riparian sites, wet-
lands, and shrublands. Kahili ginger dominates early succession in sites destroyed by fire.
Both yellow ginger and white ginger are usually found at the forest edges, not in dense
shade beneath the canopy.
Reproduction and Dispersal. Reproduction is both sexual and vegetative, Kahili ginger
spreads by seeds, rhizomes, intentional plantings, and careless disposal of garden trim-
mings. All rhizome pieces sprout stems, and also grow outward to expand the size of the
clump. Each clump has several flower heads, and each flower head produces 20–600 seeds.
Fewer seeds are produced in low light under forest canopies. Dense infestations in Hawai’i
may have 200 seeds per sq. ft. (2,000 per m2). Dispersal over longer distances to new areas
occurs when birds eat the fruit and expel the seeds. The two birds primarily responsible for
dispersal are both introduced species, the Japanese White-eye and the Red-billed Leiothrix.
Although passage through the digestive tract may enhance germination of seeds, most seeds
are viable for only one year.
Impacts. Because kahili ginger grows quickly, lives at least 70 years, and sprouts from
rhizomes when stems are removed, it has the potential to invade all wet habitats in Hawai’i
below 5,500 ft. (1,700 m) elevation. Kahili ginger does not need disturbance and can invade
healthy, closed-canopy forests, where plants thrive because the species is shade-tolerant. It
outcompetes native plants for light, space, nutrients, and moisture, displacing the under-
story of native mosses, ferns, and shrubs. Because seedlings of native plants cannot grow
through the thick rhizome mat, kahili ginger prevents forest regeneration, with the result
that the multilayered forest is replaced by dense, monospecific thickets.
Infestations of kahili ginger change the habitat for wildlife, especially for birds. Results
from a study conducted in Hawai’i Volcanoes National Park indicated that populations of
Japanese White-eye (see Volume 1, Vertebrates, Birds, Japanese White-Eye), a nonnative
bird, increased in ginger-dominated forests.
Although the rhizome system is extensive, the shallow roots beneath the rhizome mat do
not grip the soil well. Due to the weight of the rhizomes mass, the entire plant often falls as a
unit from steep, wet slopes. Such erosion downgrades water quality and causes siltation. The
rhizome mat can block stream flow, altering hydrology. The density of the rhizome mat pre-
vents water from infiltrating and percolating to the water table, reducing the forests’ useful-
ness as watersheds.
394 n FORBS

Management. Eradication of kahili ginger is very difficult, usually requiring a combina-

tion of physical removal and herbicide application. Physical removal includes digging out
small stands or small plants and cutting off top growth of larger plants. All rhizomes must
be removed when plants are dug out. Because fruit consumed by birds is a major means of
dispersal, inflorescences should be pulled off before seeds mature. If seeds are mature, care
must be taken to properly dispose of the flower heads. Because the cut stems and leaves do
not sprout, they can be composted. Fire is inefficient because neither stalks nor roots burn
well. Additionally, most sites are too ecologically sensitive to attempt control by burning.
Kahili ginger is not poisonous and is palatable to livestock.
Chemical control is necessary for large plants. Herbicides applied directly to exposed rhi-
zomes after foliage is removed is the most effective means of control, although it may take
12–15 months for rhizomes to die. Treated rhizomes will not sprout and can be left in place
to decay. Metsulfuron-methyl is the most effective herbicide sprayed on foliage, but it will
also damage nontarget native plants. The most efficient method is to chop down the stalks
and spray the regrowth when it reaches 20–24 in. (50–60 cm) tall. Glyphosate is also an
effective foliar spray. Kahili ginger may also be controlled by imazapyr, amitrol, and triclo-
pyr ester.
Glyphosate is effective on rhizomes of white ginger after the shoots have been removed.
White ginger is also sensitive to foliar application of picloram and to metsulfuron, and to
some extent to triclopyr. Yellow ginger is extremely sensitive to metsulfuron applied to cut
rhizomes.
A bacterium (Ralstonia solanacearum), which causes bacterial wilt, shows good potential
as a biological control agent for kahili ginger. This bacterium is host-specific and does not
harm either white ginger or yellow ginger. Experiments have shown that spraying with a
bacteria solution, which is easily produced, decreased fruit, stunted growth, wilted leaves,
and promoted rhizome decay. The bacterium is now established in soils at Hawai’i
Volcanoes National Park and is expanding its range by means of soil water and insect
movements.

Selected References
Buddenhagen, Chris. “Ecology of Hedychium gardnerianum.” ISSG Global Invasive Species Database,
2006. http://www.issg.org/database/species/ecology.asp?si=57&fr=1&sts=&ang=EN&ver=print
&prtflag=false.
Csurhes, Steve, and Martin Hannan-Jones. Pest Plant Risk Assessment, Kahili Ginger, White Ginger, yellow
Ginger. Department of Primary Industries and Fisheries, Queensland Government, Brisbane,
Australia, 2008. http://dpi.qld.gov.au/documents/Biosecurity_EnvironmentalPests/IPA-Kahili
-Ginger-Risk-Assessment.pdf.
“Wild Ginger.” Pest Facts 4. Northland Regional Council, New Zealand, n.d. http://nrc.govt.nz/upload/
2398/Plant%20Pests%2004%20-%20Wild%20Ginger.pdf.

“Kahili” Sounds Like It’s Native to Hawai’i

K ahili ginger got its Hawaiian name because the flowering stalks were thought
to resemble the feathered staffs used by Hawaiian royalty to show their status
and lineage.
 LEAFY SPURGE n 395

n Leafy Spurge
Also known as: Wolf’s milk, Fuitour’s-grass
Scientific name: Euphorbia esula
Synonyms: Euphorbia virgata, Tithymalus esula, Galarhoeus esula, and others
Family: Spurge (Euphorbiaceae)
Native Range. Europe and Asia, including western Russia, the Caucasus region,
Southwest Asia, Mongolia, China, and Korea.
Distribution in the United States. Most of the United States except the southern states.
From the West Coast east to New Mexico, through the Great Plains States, the Midwest,
and into the Mid-Atlantic states and New England.
Description. Leafy spurge is
an erect branching perennial
herb reaching 2–3.5 ft. (0.6–
1 m) tall. Plants become woody
at the base when mature. The
smooth, or sometimes hairy,
stems often occur in clusters
arising from one root. All parts
of the plant exude a white latex,
or white milky sap, increasingly
so as the plant ages. Leaves are
stalkless and variable in both
shape and size. They may be lin-
ear, lance-shaped, or oval, 1–4
in. (2.5–10 cm) long and 0.25
in. (6 mm) wide. Tips are either
pointed or rounded, and leaf
margins are smooth or slightly
wavy. Leaves are alternate on
the stem, but so closely spaced
that they may seem to be oppo-
site or even whorled. Leaves on
seedlings are similar to those on
mature plants, but smaller, and
leaf pairs are rolled or folded
together. The bluish-green color
gives the plant a slightly frosted
appearance. Stems die back at
the onset of the cold season,
turning yellow or red before
leaves drop. Plants need approx-
imately 42 days of chilling tem-
A common invader of grassland or rangeland, leafy spurge may be
peratures to break dormancy accidentally dispersed by farm machinery. (Native range approximated
the following spring. from USDA GRIN and selected references. Introduced range adapted
Leafy spurge has a complex from USDA PLANTS Database, Invasive Plant Atlas of the United States,
root network that is tough and and selected references.)
396 n FORBS

A. The many branches make the plant appear shrubby. (Steve Dewey, Utah State University, Bugwood.org.)
B. Tiny floral parts are surrounded by cup-shaped bracts. (Norman E. Rees, USDA Agricultural Research Service,
Bugwood.org.) C. Leaves are often linear. (Steve Dewey, Utah State University, Bugwood.org.) D. Flower clusters
grow at the end of stems. (George Markham, USDA Forest Service, Bugwood.org.) E. Leafy spurge (right) is larger
and more leafy than the ornamental cypress spurge. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology,
University of Connecticut, Bugwood.org.) F. The complex root system is tough. (Steve Dewey, Utah State
University, Bugwood.org.)

woody. Long horizontal roots and short feeder roots grow within 12 in. (30 cm) of the soil
surface. Several vertical roots extend as far as 10–20 ft. (3–6 m) into the soil, occasionally
reaching 30 ft. (9 m) deep. Long, old roots become corky and drought tolerant. The exten-
sive root system stores an abundance of nutrient reserves, and the roots and root crowns
have numerous regenerative buds. First-year shoots, which are more abundant at the edges
of clone infestations, do not yet have a woody crown.
What appear to be yellow-green flowers, the showy part, are bracts at the base of the flow-
ers. The tiny flowers are reduced to the basics. Each female flower consists of one pistil, and
each male flower consists of one stamen. What looks like a flower is actually one female sur-
rounded by 11–21 males, all inside a cuplike bract. The bract clusters begin to develop in late
May and early June, with the flowers following in mid-June. Plants may flower again in the fall.
Flowers have abundant pollen and nectar and are pollinated by at least 60 insects in the
United States. Fruits are smooth, round, three-celled capsules 0.01 in. (3 mm) in diameter.
The tiny seeds, one to a cell, are oval, smooth and gray-brown to purple with yellow flecks.
Related or Similar Species. Leafy spurge is variable in its native range, and many regional
biotypes exist in the United States, confusing the taxonomy of these spurges. Leafy spurge
may be a hybrid or a series of hybrids. Toothed spurge, also known as serrate spurge, and
eggleaf spurge are very similar to leafy spurge. Madwoman’s milk, also called sun spurge,
is an annual. Caper spurge, or gopher plant, may be either an annual or a biennial. The more
recently introduced Geraldton carnation weed is a perennial, as is the ornamental cypress
spurge. Several of these spurge species are invasive or potentially invasive.
Introduction History. First recorded in Massachusetts in 1827, leafy spurge is believed to
have arrived in the United States as a contaminant in seed in the early 1800s. Introduction
 LEAFY SPURGE n 397

from Russia, in ship ballast and seed contamination in grain, probably occurred multiple
times. By the early 1900s, the plant had reached as far west as North Dakota. It is currently
spread with commercial seed, forage, or hay, and by fragments attached to farm or road
equipment.
Habitat. Leafy spurge has wide environmental tolerances. It is found in subtropical to
subarctic climates, and xeric to mesic sites. In its native range, leafy spurge occupies grass-
lands, hedges, and ruderal habitats. In the United States, it is most common in grassland
and rangeland. It easily invades disturbed prairies, savannas, pastures, abandoned fields,
and roadsides, but it also is found in undisturbed prairies and woodlands. Although it grows
in moist to dry habitats, leafy spurge is most aggressive in dry areas, where it encounters less
native competition. It is present on all soil types but grows best in coarse-textured soil.
Plants tolerate flooding for as long as four months, and occupy stream and ditch banks.
Although it can grow in shade, leafy spurge prefers sunlight. Fewer flowers and shoots are
produced under shady conditions.
Reproduction and Dispersal. Because leafy spurge reproduces both sexually and asex-
ually, it grows and spreads quickly. Stands usually start by seed, then expand vegetatively.
Reproduction in sandy soils is predominantly vegetative due to a small soil seed bank,
approximately 20 seeds per sq. ft. (200 per m2). In clay soils, with approximately 200 seeds
per sq. ft. (2,000 per m2), 60 percent of sprouts emerge from seeds. Times of emergence,
flowering, seed set, and germination vary with geographic location.
Seeds mature about 30 days after flowering occurs, usually in late July. Each flower stem
can produce 250 seeds. The seed capsules open explosively, ejecting the seeds as far as 15 ft.
(4.5 m) away from the parent plant. Long-distance dispersal is aided by water and wildlife.
Seeds that survive the digestive tract of sheep and goats are spread through their manure.
Because no dormancy or cold period is required, seeds are able to germinate as soon as they
mature. Seeds have a high germination rate, 99 percent in the first two years, and they
remain viable in the soil for 7–9 years. Optimal temperatures for germination are 68–85ºF
(20–30ºC), preferably with alternating freezing and thawing combined with periods of dark-
ness. With sufficient moisture, seeds germinate throughout the growing season. Although
many seedlings appear to die, they resprout from root buds that developed soon after germi-
nation. Stems from seedlings and root buds usually do not flower the first year.
Plants also reproduce from the roots and root crowns. Root buds are primarily within
12 in. (30 cm) of the soil surface, but root pieces 9 ft. (2.8 m) deep can grow new plants.
Buds remain viable for 5–8 years. Any root fragment, even as small as 0.5 in. (1.25 cm),
can grow into a new plant. Clones extend their cover vegetatively at the rate of several feet
per year. Damage to aerial plant parts stimulates the root buds.
Impacts. Leafy spurge is very aggressive and a major problem in drier western states.
Over five million acres (2 million ha) in the United States and Canada are infested. It can
completely take over large areas, forming dense patches that replace native grasses and forbs
by shading the ground and tapping water and nutrients. Because it can germinate when tem-
peratures are close to freezing, leafy spurge is one of the first plants to emerge in spring. Its
thick litter layer smothers the ground, and the decomposing plant tissue exudes a toxin that
prevents other plants from growing. Severe infestations reduce both biodiversity and carry-
ing capacity of rangeland. Studies on the Great Plains indicate that numbers of species of
native grassland birds decline where leafy spurge is dense.
The sap causes skin irritation, blisters, or severe rashes in humans, and can seriously
damage the eyes. Because of the white sap, which affects the digestive tract in a similar man-
ner, it is unpalatable or even toxic to native herbivores, such as elk, deer, and antelope.
398 n FORBS

Significant ingestion can be fatal to cattle, which will not eat leafy spurge unless nothing else
is available or the plant is a contaminant in hay.
Leafy spurge infestations are responsible for significant economic losses, and can reduce
crop yields by 10–100 percent. A conservative estimate of monetary loss in 1979 due to cost
of control and loss of productivity was $10.5 million. A 1990 study concluded that the
direct financial impact in Montana, North Dakota, South Dakota, and Wyoming was
$40.2 million, with a secondary impact of $89 million. Economic impact in the Dakotas,
Montana, and Wyoming is estimated at $144 million a year in crop and animal losses and
control expenses
Management. The best control of leafy spurge is early detection and elimination. Once it
has gained a foothold, its deep root system and multiple buds make it almost impossible to
totally eradicate. The abundant reserves in the root system enable the plant to survive
repeated removal of aerial parts. Sites should be monitored for regrowth for at least 10 years
after supposed eradication.
Physical means may control the growth and spread, but it is highly improbable that
plants can be eradicated because most physical removal does not affect the roots except to
stimulate bud growth. Very small stands may be removed by hand-pulling or digging, but
every piece of root must be removed. Gloves provide protection from toxic sap. Physical
removal, however, disturbs the soil and other plant species, exposing more areas that may
become infested. Continuous grazing by sheep and goats can reduce the plant’s density by
adversely affecting reserves and seed production, but root crown buds will continue to
sprout for several years after grazing stops. Sheep and goats should also be quarantined after
grazing leafy spurge to prevent the transport of seeds in their manure. Cultivation or disking
of infested fields is inadvisable because it spreads root pieces. Fire is generally not successful
because roots and root crowns will resprout, but it does remove litter and other dead plant
material. It may also encourage native bunchgrasses to grow and compete with leafy spurge.
Fire may provide good control if repeated for at least 5–10 years.
Any chemical treatment must be repeated over several years in order for the reserves in
the roots to be depleted. Timing is critical. Chemical applications work best when combined
with grazing, but can be detrimental to beneficial insects, including biological control
agents. Leafy spurge can purge unwanted chemicals, such as herbicides, from its surface root
system. The rest of the root regenerates after the soil concentration of herbicide has dissi-
pated. Systemics, such as glyphosate or 2,4-D, are best applied in June on the developing
flowers and seeds, or in mid-September when plants are transporting nutrients into the
roots. Picloram is the most effective herbicide and, in combination with 2,4-D, can provide
90–95 percent control after five years. Research shows that chemical treatment in the fall
combined with a spring burn reduces seed production, which in turn reduces the infesta-
tion. Native grasses respond well when herbicide is selectively applied to leafy spurge.
The potential for biological controls is good. Twelve insect species, natural enemies from
Europe and China, have been released in the United States since 1965, but few have estab-
lished populations. These insects may help to manage infestations, allowing native plants
to effectively compete with leafy spurge. Larvae of the hawkmoth (Hyles euphorbiae) eat the
leaves and flowers but have succumbed to predation by ants, carabids, and mammals. The
larvae of three clearwing moths (Chamaesphecia spp.) feed on roots but also failed to estab-
lish. A longhorn beetle (Oberea erythrocephala) has become established in Montana. The
adults eat stems and roots while larvae feed on the root crown and roots. Four species of flea
beetles (Aphthona spp.) now have established populations and have been the most successful
in defoliating plants. Adults feed on stems, leaves, and flowers, but more important, the
 MUSK THISTLE n 399

Successful Eradication

R esults of biological control, documented by before and after photos by Team

Leafy Spurge, illustrate that insect pest introductions can be successful in as lit-
tle as two years. After 40 years of herbicide use failed to limit dense leafy spurge
infestations on his rangeland, a rancher in Montana introduced flea beetles and
is now a firm supporter of biological control. Densities of leafy spurge in other sites
have been reduced from almost 50 percent cover down to 12 percent, allowing
native grasses to recover.
Source: Team Leafy Spurge, n.d.

larvae eat the root hairs and roots, opening them to fungal infections. The short-tip gall
midge (Spurgia esulae) causes galls to form, which prevents flowering.

Selected References
Biesboer, David D. “Element Stewardship Abstract, Euphorbia esula.” Global Invasive Species Team,
Nature Conservancy, 1996; modifed 2009. http://wiki.bugwood.org/Euphorbia_esula.
“Euphorbia Genus.” Plant Pest and Health Prevention Services (PHHPS), California Department of
Food and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/euphorbia.htm.
“Leafy Spurge, Euphorbia esula L.” 2005. Alaska Natural Heritage Program. Environment and Natural
Resources Institute, University of Alaska, Anchorage, 2005. http://akweeds.uaa.alaska.edu/pdfs/
potential_species/bios/Species_bios_EUES.pdf.
“Leafy Spurge, Euphorbia esula L.” Weed of the Week. U.S. Department of Agriculture, Forest Service,
Invasive Plants, 2006. http://www.na.fs.fed.us/fhp/invasive_plants.
Nowierski, R. M., and R. W. Pemberton. 2002. “Leafy Spurge.” In: Biological Control of Invasive Plants in
the Eastern United States, by R.Van Driesche et al. U.S. Department of Agriculture, Forest Service
Publication FHTET-2002-04, Morgantown, WV, 2002. http://www.invasive.org/eastern/biocontrol/
14LeafySpurge.html.
Team Leafy Spurge. “Leafy Spurge Biological Control Information and Photo Resource Gallery”, n.d.
http://www.team.ars.usda.gov/v2/photos.html.

n Musk Thistle
Also known as: Nodding thistle, plumeless thistle
Scientific name: Carduus nutans
Synonyms: C. nutans ssp. leiophyllus, C. nutans ssp. macrocephalus, C. nutans ssp. macrolepis,
Carduus armenus, C. thoermeri, C. coloratus, C. schischkinii, and others
Family: Sunflower (Asteraceae)
Native Range. Western and central Europe, north to Scotland, east to Yugoslavia, and the
Ukraine. Also the western part of Asia from the Caucasus east to western China.
Distribution in the United States. Every state except Maine, Vermont, Florida, Alaska,
and Hawai’i. Most problematic in the Northwest Coast, the Rocky Mountains, Great
Plains, Upper Midwest, and central Atlantic states.
Description. Musk thistle is an aggressive biennial herb. In their first year of growth,
seedlings develop a large flat-lying basal rosette, up to 5 ft. (1.5 m) in diameter. The plant
400 n FORBS

flowers in the second year,

rarely in the first year, sending
up a flower stalk 1.5–6 ft.
(0.5–1.8 m) tall. Plants have
considerable variation in leaf
size, spine length, flower head
diameter, hairiness, and shape
of bracts. Leaves and stems are
spiny. The coarsely lobed leaves
are dark green, with a smooth
waxy surface, prickly toothed
margins, and a yellowish-white
spine at the tip. Leaves in the
basal rosette are lance-shaped
and can be as large as 16 in.
(40 cm) long and 6 in. (15 cm)
wide. The stem leaves are alter-
nate and smaller, 3–6 in. (7.5–
15 cm) long, with light-green
midribs and a white margin.
The base of each leaf extends
down the full length of the stem
to form a narrow, spiny wing-
like structure.
Many small roots develop in
the fall after seeds germinate. A
long, fleshy taproot, which
may be branched, develops in
the spring. Corky and hollow
near the soil surface, the taproot
may extend to a depth of 16 in.
Musk thistle grows in a variety of environments, but is absent from deserts, (40 cm) or more.
shady forests, and coastal areas. (Native range adapted from USDA GRIN Plants begin to bolt in
and selected references. Introduced range adapted from USDA PLANTS March, and flowers appear from
Database, Invasive Plant Atlas of the United States, and selected references.) May through September. Flower
stalks, one or more rising from
the root crown, are multi-branched. Large, showy disc-shaped flower heads, 0.8–3 in.
(2–7.5 cm), occur singly at the tips of stems. The base of the flower head resembles a pine
cone, with many large, lance-shaped bracts, tipped with spines. The outer bracts are broad,
as wide as 0.4 in. (1 cm), with long, flat spine tips. The soft, narrow inner bracts may be
purplish and are only sparsely spiny. Hundreds of tiny red-purple tubular flowers, each
1.5–3.5 in. (4–9 cm) long, comprise the disc flower. The short, 0.8 in. (2 cm) pedicels bend
at maturity, causing the flower heads to droop at a 90-degree angle to the stem. Seeds mature
about one month after flowering occurs. Each plant produces thousands of smooth straw-
colored seeds, 0.1–0.2 in. (3–5 mm), with cream-colored plumed bristles.
Related or Similar Species. The taxonomy of musk thistle is not clear, because plants
exhibit considerable variation in morphological characteristics. Carduus nutans may be one
species with four subspecies or three separate species. The genus Carduus has several
 MUSK THISTLE n 401

A. Flowering plants can be as tall as 6 ft. (1.8 m). (Norman E. Rees, USDA Agricultural Research Service,
Bugwood.org.) B. Musk thistle flowers (right) are distinctly different from bull thistle flowers (left). (Steve
Dewey, Utah State University, Bugwood.org.) C. Plants grow as a rosette in their first season. (Richard Old,
XID Services, Inc., Bugwood.org.) D. Stems are covered with sharp prickles. (Steve Dewey, Utah State University,
Bugwood.org.) E. Seedheads are filled with plumed bristles. (Steve Dewey, Utah State University, Bugwood.org.)

additional species similar to musk thistle, all of which are erect plants with prickly winged
stems and leaves. Common names may be confusing because they are not always distinctive.
Scientific names provide more clarity. No Carduus species are native to the United States.
Plumeless thistle (Carduus acanthoides) is a winter annual or biennial herb growing as tall
as 5 ft. (1.5 m). Its basal leaves are 4–8 in. (10–20 cm) long. This plant is distinguished by its
erect flower heads less than 1 in. (2.5 cm) in diameter and narrow, lance-shaped hairy
bracts. It is more abundant in the northern states. Musk thistle and C. acanthoides readily
hybridize, resulting in plants with intermediate characteristics.
Italian thistle (Carduus pycnocephalus), from the Mediterranean region, and slender-
flowered thistle (Carduus tenuflorus), from central Europe, are primarily winter annuals
reaching 6.5 ft. (2 m) tall. C. pycnocephalus has narrow cylindrical heads 0.6 in. (1.5 cm)
across, in clusters of 2–5 at the ends of branches. Its leaves are covered with wooly hairs.
Flowerheads of C. tenuflorus occur in clusters of 5–20. C. pycnocephalus and C. tenuflorus
are currently found in only a few states but are expanding their ranges. Because C. tenuflorus
is self-pollinating, these two species rarely hybridize.
Other introduced thistle genera are also invasive. Cirsium species are differentiated from
Carduus species by pappus hairs which are divided and feathery. Native to southwestern
Asia, Canada thistle (Cirsium arvense; see Forbs, Canada Thistle) is a perennial with creeping
roots that give rise to new plants. It has small unisexual flower heads, with male and female
flowers on different plants. Its stems and flower heads are smooth rather than prickly. The
upper leaf surfaces of bull thistle (Cirsium vulgare), a coarse biennial native to Europe and
North Africa, are covered with stiff bristly hairs, making them rough to the touch.
Although the leaves of Scotch thistle (Onopordum acanthium), a plant native to Scotland,
are tipped in spines, the plant is covered with silvery white cottony wool rather than with
bristles.
402 n FORBS

Starthistles (Centaurea spp.; see Forbs, Yellow Starthistle), native to southern Europe and
southwestern Asia, are distinguished by long, sharp spines surrounding the flower head.
Several related genera, such as Centaurea, Saussurea, and Cirsium, have species native to
the United States that are similar to Carduus.
Introduction History. Although musk thistle was brought to the eastern United States in
the early 1800s, probably as a contaminant in ship ballast, it arrived in the midwestern states
about 1900. It was recognized as a serious weed in the 1940s, and by the 1970s had spread
to 42 states.
Habitat. Musk thistle has a wide range of environmental tolerances, but does best in well-
drained fertile alluvial soils, of any texture. It grows poorly on acidic or nutrient-deficient
soils, but is somewhat tolerant of salinity. Although musk thistle grows in open natural
areas, such as meadows, prairies, and deciduous forests, it heavily infests disturbed areas,
such as sites subjected to landslides, flooding, fire, heavy grazing, or trampling. It is also fre-
quently found in pastures, fields, annual grasslands, rangeland, roadsides and other right-
of-ways, stream or ditch banks, and waste areas. It does not grow in very wet or very dry
sites or in shade, and is absent from deserts, dense forests, and coastal environments.
Although foliage is killed by a hard frost, the spiny stems remain erect. Musk thistle grows
from sea level to 10,000 ft. (3,000 m) elevation.
Reproduction and Dispersal. Musk thistle reproduces only by seed, and plants may be
either cross-pollinated or self-pollinated. One plant can quickly become an infestation.
The number of seed heads on each plant depends on site conditions, 1–18 on poorer sites,
increasing to 24–46 on favorable sites. One 6 ft. (1.8 m) tall plant, however, had 643 seed
heads! Plants will continue to flower and set seed until the soil moisture is depleted, producing
more seeds in climates with spring and summer rainfall. One flower head can produce as
many as 1,500 seeds, and one plant can produce as many as 120,000. The number of seeds
per flower head decreases to as little as 25 later in summer. Most seeds fall within 165 ft.
(50 m) of the parent plant, but some are transported long distances by wind, small mammals,
birds, or water. The plumed seeds can be windblown for miles. Seeds become sticky when
moistened, enabling them to adhere to surfaces. Human activity, involving farm animals,
machinery, and vehicles, aids the spread of musk thistle seeds. Seeds may also be a contami-
nant in hay and crops. Although most seeds germinate within three years, some seeds remain
viable in the soil for more than 10 years, enabling the plant to rapidly invade areas after a dis-
turbance. Because seeds usually need a cool winter period, 40 days at temperatures below
50ºF (10ºC), in order to germinate, most seeds germinate from late summer to spring, in moist
soils. Those that germinate in late fall may live as annuals and flower the subsequent summer.
Although musk thistle reproduces only by seed, many buds on the root crown can sprout
after the top portion of the plant is damaged.
Impacts. Infestations of musk thistle degrade grasslands and inhibit succession. Plants
frequently form large, dense colonies that crowd out native plants and suppress growth of
forage plants by competing for light, nutrients, and moisture. Musk thistle is unpalatable
to livestock and wildlife. Animals avoid grazing near the prickly plants, and selective grazing
of native plant species decreases competition for the thistle. One plant per 16 sq. ft. (1 per
1.5 m2) can reduce pasture yields by more than 20 percent. Musk thistle has alleleopathic
properties. Its seeds and dead leaves inhibit germination and growth of pasture grasses. At
the bolting stage, the larger rosette leaves begin to decompose, possibly releasing inhibitors
to other plants. Musk thistle threatens rare or sensitive species such as the Sacramento
Mountain thistle in New Mexico and sand dune thistle, which is native to the sand dunes
on the Great Lakes shoreline.
 MUSK THISTLE n 403

In contrast, seeds are important feed for songbirds, and flowers are attractive to insect
pollinators.
Management. The best control is to maintain good range management and ensure that
seed, hay, bedding, and equipment are clean of contaminants. Infected areas should be
avoided. Care should be taken during any type of management to minimize soil disturbance,
which triggers germination from the seedbank. Eradication may take 15 or more years.
Because seedlings are intolerant of competition, reestablishment of native vegetation is
effective.
Physical control, such as mowing or burning, reduces seed production but will not elimi-
nate infestations because new stems will emerge from the root crown. A single mowing is
ineffective, because stands have plants of different heights and flower and seed develop-
ment. Tilling, or any method which severs the root 2–4 in. (5–10 cm) below the soil surface,
will kill plants but is not practical in crop or natural areas. Small plants in small populations
can be hand-pulled all year, but pulling is best done before seeds develop. Any flower heads
present should be bagged and disposed of to prevent seed dispersal.
Chemical control will be only temporary if the conditions that caused the infestation are
not changed. Preemergents are ineffective. Postemergence compounds, such as glyphosate,
triclopyr, dicamba, 2,4-D, or picloram, work well when sprayed on foliage, especially when
applied to the rosette stage in spring before flowering takes place. Chlorpyralid, which is
selective to sunflower, buckwheat, and pea families, may harm crops.
Because musk thistle reproduces only by seed, research for biological control has concen-
trated on seed-eating insects in southern Europe. Of the several species released in the
United States, two have become established and are effective. A thistlehead-feeding weevil
(Rhinocyllus conicus) was released in 1969. The insect hatches inside the flower head and
feeds on developing florets. It works best where limited grazing allows grasses to compete
with the thistles and can reduce thistle density by 90 percent in 5–6 years. It also attacks
Scotch thistle. Because this weevil also feeds on several native Cirsium thistles, however,
and can significantly reduce their seed production, its release in areas with Cirsium species
requires extra care.
A rosette-feeding weevil (Trichosirocalus horridus), which only occasionally feeds on
native Cirsium species, was released in 1974. The larvae feed on meristem tissues in the
rosette, causing the crown tissue to die. It can reduce musk thistle infestations by 96 percent
within six years and works best in conjunction with Rhinocyllus conicus. It also attacks other
Carduus species, such as Canada thistle, bull thistle, and Scotch thistle. The establishment of
two other insects, a root-crown fly (Cheilosia corydon) and a thistle-rosette weevil (Psylliodes
chalcomera), neither of which eat native Cirsium, is unknown. A second thistle-rosette wee-
vil, Ceutorhynchus trimaculatus, was found to be destructive to native Cirsium species and
was not released.

Selected References
Gassmann, A., and L. T. Kok. “Musk Thistle (Nodding Thistle).” In Biological Control of Invasive Plants
in the Eastern United States, by R. Van Driesche et al. USDA Forest Service Publication FHTET-
2002-04, Morgantown, WV, 2002. www.invasive.org/eastern/biocontrol/18MuskThistle.html.
“Musk Thistle.” Plant Pest and Health Prevention Services (PHHPS), California Department of Food
and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/carduus.htm.
“Musk Thistle (Carduus nutans L.) and Related Species.” Environment and Natural Resources Institute,
Alaska Natural Heritage Program, University of Alaska, Anchorage, 2005. http://akweeds.uaa.alaska
.edu/pdfs/potential_species/bios/Species_bios_CANU.pdf.
404 n FORBS

Remaley, Tom. “Fact Sheet: Musk Thistle.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas.
Plant Conservation Alliance’s Alien Plant Working Group, 2005. http://www.nps.gov/plants/alien/
fact/canu1.htm.
Zouhar, Kris. “Carduus nutans.” In: Fire Effects Information System, U.S. Department of Agriculture,
Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, 2002. www.fs.fed.us/
database/feis/plants/forb/carnut/all.html.

n Perennial Pepperweed
Also known as: Tall whitetop, dittander, giant whiteweed, slender perennial peppercress,
broadleaved pepperweed
Scientific name: Lepidium latifolium
Synonyms: Cardaria latifolia
Family: Mustard (Brassicaceae)

n Hoary Cress
Also known as: White top, perennial peppergrass, pepperweed whitetop, heart-podded
hoary cress
Scientific name: Cardaria draba
Synonyms: Cochlearia draba, Lepidium draba, Nasturtium draba
Family: Mustard (Brassicaceae)
Native Range. Pepperweed is native to Europe, the northern coast of Africa, and
Southwest and Central Asia, from Turkey and southwestern Russia to eastern China.
Hoary Cress comes from Europe, European Russia, and Southwest and Central Asia, from
Turkey to western China.
Distribution in the United States. Pepperweed grows in coastal New England, midwest-
ern and Great Plains states, from Illinois west to Nebraska, Kansas, and Texas, and in all
states west of the Rocky Mountains. Especially invasive in the western states and still
expanding its range. Hoary Cress grows in all states except in the Southeast. Most invasive
west of the Rocky Mountains. Neither species is in Alaska or Hawai’i.
Description. Perennial Pepperweed: Pepperweed is an herbaceous, perennial herb typically
1.6–5 ft. (0.5–1.5 m) tall, but it can reach more than 8 ft. (2.4 m) in height. The aerial parts
die back to the soil surface in winter. Stems, very woody at the base, may remain erect for
several years in dry areas, forming impenetrable thickets. New leaves emerge from the root
in March, creating rosettes on the soil surface for several weeks before the plants bolt to
flower. In late April, stems elongate quickly, and may reach 1.6 ft. (0.5 m) by June 1 and
3.3 ft. (1 m) by June 15, branching at the top. Leaves are glabrous and waxy, green to
gray-green, sometimes with reddish spots. Leaves are elliptical and vary in size according
to location on the plant. Rosette leaves, 4–12 in. (10–30 cm) long and 1–3 in. (2.5–8 cm)
wide, have long petioles, 1.2–6 in. (3–15 cm), and serrate margins. Stem leaves are alternate,
lance-shaped or oblong, and tapered at the base, with entire or slightly toothed margins.
Leaf petioles become shorter with stem height, until leaves on the upper part of the stem
are sessile. Leaves also become successively shorter up the stem. Both the rosette leaves
and lower stem leaves senesce as the stems grow taller.
Pepperweed has a large semi-woody root crown and many rhizomes, which are long,
thick, and almost branchless. Although 20 percent of the root biomass is in the top 4 in.
 PERENNIAL PEPPERWEED AND HOARY CRESS n 405

(10 cm) of soil, and 85 percent

in the top 24 in. (60 cm), roots
can extend more than 10 ft.
(3 m) deep. Roots are phreato-
phytic, meaning that they grow
down to the water table.
Plants flower from May
through August. Small, cross-
shaped flowers, about 0.1 in.
(3 mm) wide, with four white
petals form dense clusters on
panicles, 5–6 in. (13–15 cm)
wide, at the stem tips. The fruit,
which matures in August and
September, is a small, 0.08 in.
(2 mm), two-chambered pod
called a silicle. The silicle is
round or slightly flattened, with
long hairs. It is attached to the
plant by a pedicel longer than
the fruit. Each chamber produces
one tiny reddish-brown seed, but
stressed plants frequently abort
the seeds. The fruit do not readily
release the seeds and remain on
the stem until the following
spring. Stems become dried
stalks in mid to late summer.
Hoary Cress: Hoary cress is a
smaller plant, an herbaceous
perennial with erect stems
growing 6–20 in. (15–50 cm) Perennial pepperweed grows best in wetlands and is often found along
tall. Plants usually have one irrigation canals. (Native range adapted from USDA GRIN and selected
stem, which may occasionally references. Introduced range adapted from USDA PLANTS Database,
branch near the top. Basal Invasive Plant Atlas of the United States, and selected references.)
leaves, growing in a rosette, are
1.5–4 in. (4–10 cm) long and lance-shaped. Stem leaves are smaller, 1.5–3 in. (4–8 cm)
long, and broadly oval. They are alternate and sometimes have toothed margins. The stem
leaves are sessile, and their broad bases clasp the stem. Both stems and leaves, blue-green
to gray-green, are softly hairy at the base of the plant and become less hairy with height.
Hoary cress has an extensive root system with abundant nutrient reserves. Rootstocks
may extend 6 ft. (1.8 m) deep, and long rhizomes have numerous buds which create new
shoots.
Plants flower from April to June, and seeds mature from June to August. Flowers develop
in terminal, branching racemes of flat-topped clusters. The four narrow, short petals
forming a cross are white. Glabrous seed pods, called silicles, are heart-shaped at the base
and supported by slender pedicels 0.2–0.6 in. (6–15 mm) long. Each flat silicle has 2–4
seeds.
406 n FORBS

Related or Similar Species.

Although 75 species of Lepidi-
um, both annual and perennial,
grow in North America, no
natives grow aggressively or are
invasive. Two alien invasive
species of Cardaria can be dis-
tinguished by the shape of their
seed pods. Lens-pod hoary
cress has circular pods that are
glabrous and fairly flat. Pods of
hairy whitetop are spherical
and covered with fine hairs.
Introduction History. Pep-
perweed was accidentally intro-
duced to California in the early
1900s, perhaps as a contami-
nant in sugar beet seed from
Europe. It was first reported in
Massachusetts in 1924 and in
Connecticut in 1933, with an
unknown source. Hoary cress
was first recognized in Cali-
fornia in 1876, probably intro-
duced as a contaminant in ship
ballast or alfalfa. Similar intro-
ductions took it to the south-
western United States in 1910
and to New York State in 1898.
Habitat. Pepperweed is
capable of growing in environ-
Hoary cress is invasive in sunny sites and in dry pastures and crops. ments as different as Donner
(Native range adapted from USDA GRIN and selected references. Summit in the Sierra Nevada,
Introduced range adapted from USDA PLANTS Database, Invasive Plant California, at 8,500 ft. (2,600 m)
Atlas of the United States, and selected references.)
and in the coastal salt marshes of
San Francisco Bay. Although
adapted to salinity and also found on salt pans in the Intermountain West, it does not
require salty soils. It prefers wetlands, such as riparian areas, floodplains, and coastal
marsh, but can also colonize seasonally dry meadows. It is often found in alfalfa fields,
rangeland, native wet hay meadows with sedges and rushes, and along the irrigation canals
and ditches that carry water to agricultural fields. Pepperweed does not grow in upland
soils or where drought is prolonged. It dominates in areas that have been disturbed or
are already weedy, such as roadsides, overgrazed pastures, and abandoned cropland.
Although hoary cress is found in a variety of environments, it grows best in open, sunny
grassland sites with neutral or alkaline soils. It is most often found on land disturbed by
grazing or cultivation, such as fields, pastures, waste areas, and feed lots, and along road-
sides and irrigation ditches. It is a more significant pest in the dry western states and is not
as problematic in areas that receive higher rainfall. It is found at more than 8,000 ft.
 PERENNIAL PEPPERWEED AND HOARY CRESS n 407

A. Stem leaves of perennial pepperweed are supported by petioles. (Joseph M. DiTomaso, University of
California–Davis, Bugwood.org). B. Perennial pepperweed flowers grow in dense panicles. (Leslie J. Mehrhoff,
Ecology and Evolutionary Biology University of Connecticut, Bugwood.org.) C. Pepperweed plants frequently
form dense thickets. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology University of Connecticut,
Bugwood.org.) D. Stem leaves of hoary cress are sessile and clasp the stem. (Joseph M. DiTomaso, University
of California–Davis, Bugwood.org.) E. Hoary cress flower clusters have flat tops. (Chris Evans, River to River
CWMA, Bugwood.org.) F. Hoary cress seed pods are flattened pods with heart-shaped bases. (Joseph M.
DiTomaso, University of California–Davis, Bugwood.org.)

(2,500 m) elevation in parts of the mountainous west and survives freezing temperatures
and snowfall.
Reproduction and Dispersal. Perennial Pepperweed: Pepperweed reproduces both sexu-
ally and vegetatively. Seed production is variable, perhaps in response to variations in sea-
sonal rainfall. A mature root crown may produce several flowering stems. Although more
flowers develop in dry years, each produces fewer seeds. One acre of plants can produce
more than 6.5 billion seeds (16 billion per ha) every year. Seeds readily germinate in wet
sand or mud, and seedlings can produce flowers and seeds in their first year of growth.
Seedlings, however, are rare in dense infestations. Because the seeds have no hard coat, their
viability is short term. Absence of a seed bank may account for the few seedlings found in
the wild.
Pepperweed creates large monospecific colonies from spreading rhizomes, and any small
root piece will sprout a new plant. Root fragments can tolerate dry conditions, allowing
them to be transported and become established elsewhere. Root fragments or seeds can be
dispersed by water, or as contaminants in soil or hay bales, including those used for erosion
control.
Hoary Cress: Hoary cress is a prolific producer of seeds, 1,000–5,000 seeds per plant,
with 80 percent viability. More seeds are produced in wet years, and seeds are viable for
three years. They are dispersed in hay and forage, in soil adhering to farm equipment, or
by water and wind. Such transport may be responsible for long-distance dispersal, but local
infestations increase by rhizome extensions and sprouts. Stands rapidly increase in
408 n FORBS

disturbed areas, and one plant can cover an area 12 ft. (3.6 m) in diameter after one year of
growth. Root or rhizome pieces may also be carried long distances. Seedlings often appear
after grass fires, which stimulate dormant seeds to sprout.
Impacts. Closed-canopy thickets, with 4–8 stems per sq. ft. (40–80 per m2) of soil can
obscure new spring growth of both pepperweed and other species. By sprouting from its
root after the last spring frost, usually before native species emerge, pepperweed outcom-
petes native plants for moisture and nutrients as well as for light. The basal rosettes may per-
sist all winter in regions with no frost. A dense litter layer, from old stems and leaves, can be
deep enough, 4 in. (10 cm) or more, to block light and prevent emergence of native annuals.
Even where pepperweed plants have been eradicated, the litter layer must also be removed
to encourage sprouting of native species. Pepperweed plants interfere with regeneration of
willows and cottonwoods, which is the normal succession on disturbed riparian sites.
Because it is a phreatophyte, pepperweed roots pull salt as well as water from greater depths.
The salt is then deposited on the surface, where it alters the soil chemistry and favors the
growth of halophytes.
Pepperweed displaces sedges and rushes in wetlands, which are important nesting sites
for waterfowl and shorebirds. It also displaces endangered animal species, such as the salt
marsh harvest mouse, a rodent endemic to the San Francisco Bay Area in California.
Mixed in hayfields, pepperweed lowers the protein content of the hay used for livestock.
Reports of horses being poisoned after ingesting pepperweed are being evaluated.
Hoary cress can grow into extensive mats, to the exclusion of all other plants. It displaces
forage and is toxic to livestock. Invasions of cropland reduce agricultural yields of grain,
alfalfa, and orchard crops.
Management. Eradication of pepperweed or hoary cress is difficult. The perennial roots
may remain dormant in the soil for years, and sites must be continually monitored for early
detection of new growth. Because plants are only green early in the season, they are easy to
miss. Monospecific stands are most visible when plants are in flower or fruit. All root frag-
ments must be removed or killed because even small pieces 1 in. (2.5 cm) long can grow into
new plants. All litter must also be removed. No nonchemical management methods for pep-
perweed are effective except long-term flooding. It is not known, however, whether plants
will reestablish after flooding subsides. Because pepperweed is most common on disturbed
sites, it is important to restore the native vegetation to prevent re-infestation. Different man-
agement methods may be appropriate for large monospecific stands compared with smaller,
scattered populations.
Many physical control methods such as disking are inappropriate for native meadows or
other natural environments. Other areas, such as marshes, riverbanks, and fencelines, also
cannot be easily mowed or disked. Frequent mowing to maintain short stubble and prevent
flowering is also not practical in natural environments. Although livestock will not eat pep-
perweed when the stems begin to elongate, the newly formed rosette leaves in early spring
are palatable. Old stems may be mowed in pastures to expose the new growth for grazing.
Grazing may control pepperweed growth, but plants recover when the grazers are removed.
Any type of cultivation is usually inadvisable because it cuts rhizomes into small pieces that
will resprout, resulting in stronger and larger root crowns. Although prescribed fires may be
used to destroy the litter layer, release nutrients, and stimulate new growth, burning does
not harm the root system of either pepperweed or hoary cress.
Chemical applications of herbicides, such as 2,4-D, may kill the aerial parts of the plant,
but new shoots will arise from the rootstocks the following spring. Repeated applications,
however, may eventually deplete the root reserves. Herbicides are best when applied to
 PRICKLY RUSSIAN THISTLE n 409

flowerbuds in the early flowering stage. The most effective herbicide, chlorsulfuron, which
both kills the foliage and remains in the soil to be absorbed by the roots, provides the best
long-term control. Metsulfuron-methyl is more selective and does not harm grasses, but
more research is needed. Imazapyr is also effective but is less selective, resulting in more
bare ground. If the site is appropriate, mowing or disking at the flowerbud stage followed
by herbicide application to new shoots is effective. Few herbicides are appropriate for use
in the wetlands and riparian areas favored by pepperweed, and many are not approved for
use on cropland. Glyphosate can be used in wetlands or riparian sites where other herbi-
cides are prohibited. Sites treated with glyphosate rebound with better plant diversity than
sites treated with other herbicides. Disking also aids diversity because it stimulates germina-
tion of the seed bank of other species. Herbicides applied in spring to areas that cannot be
mowed or disked can simulate mowing by removing aerial parts of plants. Herbicides can
then be applied to resprouts. Dormant, perennial roots require that the area be subjected
to spot treatment for several years.
Any biological control must be host-specific. The mustard family includes many valuable
crops, such as canola, mustard, cabbage, and kale, and several Lepidium species native to the
United States, which are threatened or endangered. Although white leaf rust (Albugo)
decreases seed production of pepperweed in the western states, it does not harm the root
system. Several insects eat the flowerbuds, which also decreases seed production, but the
roots are unaffected.

Selected References
Bossard, Carla, and David Chipping. “Cardaria draba.” In Invasive Plants of California’s Wildlands, edited
by Carla C. Bossard, John M. Randall, and Marc C. Hoshovksy. Berkeley: University of California
Press, 2000. http://www.cal-ipc.org/ip/management/ipcw.
Howald, Ann. “Lepidium latifolium.” In Invasive Plants of California’s Wildlands, edited by Carla C.
Bossard, John M. Randall, and Marc C. Hoshovksy. Berkeley: University of California Press, 2000.
http://www.cal-ipc.org/ip/management/ipcw.
Renz, Mark J., and J. M. Randall, eds. “Element Stewardship Abstract, Lepidium latifolium.” Element
Stewardship Abstract. Global Invasive Species Team, Nature Conservancy, 2000. http://
wiki.bugwood.org/Lepidium_latifolium.
Young, James A., Debra E. Palmquist, Robert S. Blank, and Charles E. Turner. “Ecology and Control of
Perennial Pepperweed.” 1995 Symposium Proceedings, California Exotic Pest Plant Council. http://
www.cal-ipc.org/symposia/archive/pdf/1995_symposium_proceedings1795.pdf.

n Prickly Russian Thistle

Also known as: Tumbleweed, Russian tumbleweed, saltwort, common saltwort, windwitch,
witchweed, prickly glasswort
Scientific name: Salsola tragus
Synonyms: Salsola kali, S. australis, S. iberica, S. kali var. tenuifolia, S. kali ssp. tragus, S. kali
ssp. ruthenica, S. pestifer
Family: Goosefoot (Chenopodiaceae)
Native Range. Semiarid steppes of southern Russia and western Siberia. Possibly native
to the Caucasus region and Southwest Asia and as far east as Mongolia and northern China.
Distribution in the United States. Every state except Alaska and Florida. Most invasive in
western states.
410 n FORBS

Description. Although the

mature plant looks like a bushy
shrub, prickly Russian thistle is
a summer annual herb that
reaches 1–5 ft. (0.3–1.5 m) tall
and as much as 6 ft. (1.8 m)
diameter. Seedlings and small
plants resemble pine seedlings
due to their finely-dissected,
needlelike leaves. Subsequent
young leaves are fleshy and soft,
about 1 in. (2.5 cm) long, and
only weakly spine tipped.
Slender, flexible stems on juve-
nile plants are red or purple
striped, either glabrous or
pubescent. Immature plants are
taller than they are wide, with
lateral branches shorter than
the main stem. Mature plants
are usually equal in height and
width. The plant’s many bran-
ches, which curve upward,
become woody and spiny as
the plants mature. Stalkless
leaves, blue-green and alternate
on the stem, are very narrow,
0.5–2.4 in. (1.3–6 cm) long,
and awl-shaped or cylindrical.
Leaves may be glabrous or
covered with a short, stiff
Dried plants of prickly Russian thistle, tumbleweeds, are commonly seen pubescence. Older leaves be-
in western states. (Native range adapted from USDA GRIN and selected come stiff and are succulent or
references. Introduced range adapted from USDA PLANTS Database, sometimes leathery, usually
Invasive Plant Atlas of the United States, and selected references.) with a pointed tip or spine at
the end.
Prickly Russian thistle has a large and spreading root system. Lateral roots are as long as
6 ft. (1.8 m), and the taproot extends 6 ft. (1.8 m) deep. Both types of roots enable the plant
to draw moisture from a wide area.
Plants usually bloom from July through October, although they will continue to flower
and produce seed until temperatures fall below 25ºF (−4ºC). The inconspicuous small flow-
ers grow solitary in leaf axils near the upper parts of the branches or in a terminal spike.
They have no petals, but the five sepals are greenish, white, or pink. Two shiny, papery,
awl-shaped bracts at the base of each flower are each tipped with a spine. Five stamens
extend beyond the sepals. Flowers are wind pollinated, both crossing and self-fertile. The
small fruit, 0.4 in. (1 cm), is enclosed in the calyx, which has five wing-like appendages,
one from the back of each sepal. The fan-shaped sepal wings are translucent pinkish to
deep red with minutely toothed margins. Each fruit contains only one seed. The dark
 PRICKLY RUSSIAN THISTLE n 411

A. Plants have a shrubby appearance but are herbaceous annuals. (Joseph M. DiTomaso, University of
California–Davis, Bugwood.org.) B. Inconspicuous flowers, with sepals but no petals, grow in leaf axils.
(Joseph M. DiTomaso, University of California–Davis, Bugwood.org). C. Seeds are dispersed as the dried plant,
the tumbleweed, blows in the wind. (Richard Old, XID Services Inc., Bugwood.org.) D. Seedlings may be
mistaken for tiny pine tree seedlings. (Phil Westra, Colorado State University, Bugwood.org.)

greenish-brown coiled embryo, or plantlet, is visible through the translucent seed coat. After
flowering and setting seed, the plant dries completely.
Related or Similar Species. The taxonomy for Salsola is confused, with no firm consensus
on either scientific name or common name. S. tragus is possibly three or more similar species
with different flower sizes and shapes. Several Salsola species are listed as noxious or
invasive.
First reported in 1958, slender Russian thistle is widespread, from the western intermon-
tane basins east to the Midwest and New England, and is expanding its range. Slender
Russian thistle seeds have been found as a contaminant in commercial bird seed. Plants grow
to 3.3 ft. (1 m) tall, and stems have green and white striations. Dark green leaves, 2 in.
(5 cm) long, on young plants are pliable, somewhat succulent to leathery, with a soft bristle
at the tip. When those leaves are shed, they are followed by short, stiff, spiny leaves about
0.5 in. (1.25 cm) long. Bracts, which are lance-shaped and spine tipped, strongly overlap
and completely cover the flowers and the fruit. Sepal wings are either absent or very narrow.
Slender Russian thistle is a noxious weed in California.
Barbwire Russian thistle closely resembles and is often confused with prickly Russian
thistle, but is shorter, usually 1.6 ft. (0.5 m) tall. It rarely hosts the leafhopper, and is found
in desert habitats, such as the Mojave Desert and east to Utah at elevations of 2,300–5,900 ft.
(700–1,800 m). Although believed to have been introduced to the southwestern United
States around 1900, it was not recognized until 1967. The immature plant is wider than it
is tall, with lateral branches longer than the main stem. Stems are thick, somewhat rigid,
with no purple striations. Leaves of barbwire Russian thistle, which do not differ between
young and mature plants, are yellow-green, fleshy, thick, and stiff, with a spine at the tip.
They are 0.2–1.2 in. (0.5–3 cm) long and often covered with short hairs. This species flow-
ers 2–3 weeks before prickly Russian thistle does, and although flower bracts are similar, the
sepal tips are stiff and spine like. Sepal wings resemble those of prickly Russian thistle but
are larger. Barbwire Russian thistle plants do not become tumbleweeds, and most seed falls
near the parent plant. Barbwire Russian thistle hybridizes with prickly Russian thistle, pro-
ducing plants with intermediate characteristics.
412 n FORBS

Oppositeleaf Russian thistle is a summer annual that grows to 1.6 ft. (0.5 m) on saline
mudflats and saltmarshes in the San Francisco Bay area of California. The glabrous stems
may be either green or red. The plant does not dry out but remains fleshy when mature.
Flowers and fruit of shrubby Russian thistle are similar to those of prickly Russian thistle,
but this species is a shrubby perennial. It has oblong to oval leaves with rounded tips, with
no spines or sharp points. Halogeton (see Forbs, Halogeton), which grows less than 1 ft.
(0.3 m) tall, can be mistaken for young prickly Russian thistle plants except for the
cottony-white hairs in its leaf axils.
Introduction History. Prickly Russian thistle was accidentally introduced to North
Dakota in 1873 by Ukrainian immigrants as a contaminant in flaxseed. It spread rapidly,
not only by its natural method of blowing in the wind, but also in contaminated seed,
threshing machinery, and railroad cars. Removal of prairie grasses by farmers provided a
smoother surface for the rolling dried skeletons. By 1900, prickly Russian thistle had
reached the West Coast.
Habitat. Prickly Russian thistle can be found anywhere the ground has been disturbed,
including dryland agricultural fields, roadsides and other right-of-ways, pastures, river bot-
toms, forest edges, and vacant lots. It also invades sandy beaches and dunes, both inland and
along the coast. It grows in a variety of soils, including alkali, but needs a loose texture and
cannot establish on hard-packed substrate. The plant is very drought tolerant and is
common in the semiarid western states, especially on overgrazed range and pasture. It will
invade areas after dry spells or where vegetation has been cleared. It grows from below sea
level in Death Valley to over 8,500 ft. (2,600 m) elevation.
Reproduction and Dispersal. Prickly Russian thistle reproduces solely by seed, and one
large plant typically produces 250,000 seeds. At maturity, the plant dries and breaks at the
base. Seeds are dispersed as the skeleton ball of woody branches, called a tumbleweed, is
blown by the wind. Often the path can be traced by a trail of seedlings that emerge the fol-
lowing spring. The tumbleweeds persist for at least a year, usually piled against an obstruc-
tion. Seeds usually remain viable for 6–12 months, and rarely for two years. Seeds germinate
in spring under a wide range of temperatures, 28–110ºF (-2–43ºC). Most seeds germinate in
March and April when temperatures are optimal, 45–95ºF (7–35ºC), but germination can
occur all year if daytime temperatures are above freezing. Seedlings are easily killed by frost.
Little rainfall (0.3 in. or 7.5 mm) is needed to stimulate germination, and seedlings develop a
taproot within 12 hours. Because seeds have no protective coat, they have no stored food
reserves. Each seed is a tiny embryonic plant, coiled inside a thin membrane, giving it the
advantage of rapid growth.
Impacts. Russian thistle is generally not competitive with other species. Seedlings are
readily shaded out by other plants. It colonizes open ground, depending on rapid growth
to gain access to deep soil moisture, thereby threatening reestablishment of native plant eco-
systems. Infestations in agricultural land reduce crop yields, especially of alfalfa and small
grains, by depleting soil moisture. Prickly Russian thistle is a host plant for the sugar beet leaf-
hopper, a carrier for curly-top virus, which affects sugar beets, potatoes, tomatoes, and beans.
Masses of tumbleweeds are a fire hazard. Entire plants can be carried by the wind as flam-
ing balls, both spreading the fire and making firefighting dangerous. Large tumbleweeds
blowing across highways are a hazard to drivers because they block vision and cause drivers
to swerve to avoid the plant. Tumbleweeds may also block stream channels and roads.
Management. Maintenance of rangeland and limiting disturbance is a key to prevention.
Care should be taken to avoid loosening the soil in abandoned areas. Prickly Russian thistle
is rarely a problem on well-tended sites where healthy stands of plants, such as perennial
 PRICKLY RUSSIAN THISTLE n 413

Russian Thistle as Food

B efore they become woody and spiny, young plants are grazed by cattle and
sheep. During the Dust Bowl in the 1930s, Russian thistle was fed to beef cattle
because no other feed was available. Young Russian thistle is also a minor part of
the diets of mule deer and elk, and tender shoots are important for bighorn sheep
and pronghorn. Seeds are eaten by birds, such as Scaled Quail and Gambel’s Quail,
and by many small mammals, such as prairie dogs and mice.

grasses, resist invasion. Plants may be successfully removed from one area, but unless the
land is revegetated, adjacent regions provide a seed source for new invasions. Seedlings
may be especially numerous along fences and other obstructions where plant skeletons
accumulate, and all sites should be continually monitored for new infestations.
Physical removal of large plants is difficult because of the spines. Although mowing large
plants stimulates new branches to grow closer to the ground, repeated mowing may provide
some control in reducing seed production. Plants may be cut, mowed, or tilled before seeds
develop, but control must be repeated until the seed bank is depleted. Mowing is effective to
eliminate seedlings. Burning is not an option because Russian thistle thrives on disturbed
sites.
Chemical control is most effective in the seedling stage. Different herbicides should be
selected for different types of sites, such as roadside versus crop or pasture. Several preemer-
gents are effective if used along roadsides in the fall. Postemergents, such as dicamba, glyph-
osate, triclopyr, and others, are more effective on seedlings than on mature plants. Few
herbicides will kill plants after they reach the spiny stage. The herbicide 2,4-D causes plants
to become tough and leathery, making them even more difficult to manage. Several applica-
tions may be necessary as seedlings continue to emerge throughout the spring and summer.
Because populations may develop resistance to specific herbicides, changing types is
recommended.
Biological control has had limited success. Two insects, a leaf-mining moth (Coleophora
klimeschiella) and a stem-boring moth (C. parthenica), released in the 1970s, did not provide
sufficient control. The host-specific blister mite (Aceria salsolae) from the Mediterranean
region stunts plants by killing the growing tips of young stems, preventing the development
of flowers. It attacks only S. tragus and its close relatives. Other possibilities include a moth
(Gymnancyla canela), whose caterpillers eat developing seeds and stems. Central Asia is
being explored for other natural enemies.

Selected References
Morisawa, Tunyalee. “Element Stewardship Abstract, Salsola kali.” Global Invasive Species Team,
Nature Conservancy, 1988; modified 2009. http://wiki.bugwood.org/Salsola_kali.
Orloff, S. B., D. W. Cudney, C. L. Elmore, and J. M. DiTomaso. “Pest Notes: Russian Thistle.” UC ANR
Publication 7486. Agriculture and Natural Resources, Statewide IPM Program, University of
California, Davis, 2008. http://www.ipm.ucdavis.edu/PMG/PESTNOTES/pn7486.html.
“Salsola genus Part 1, Russianthistle or Common Russianthistle, Spineless Russianthistle, Barbwire
Russianthistle.” Plant Pest and Health Prevention Services (PHHPS), California Department of
Food and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/salsola.htm.
414 n FORBS

Smith, Lincoln. “Biological Control of Russian Thistle (Tumbleweed).” Proceedings of the 60th
Conference of the California Weed Science Society, Monterey, CA: 90-93, January 28–30, 2008.
http://www.cwss.org/proceedingsfiles/2008/90_2008.pdf.

n Purple Loosestrife
Also known as: Purple lythrum, rainbow weed, salicaire, spiked loosestrife
Scientific name: Lythrum salicaria
Synonyms: None
Family: Loosestrife (Lythraceae)
Native Range. Eurasia, including all of Europe and east through China and Korea to
Japan. Also from the northern coast of Africa.
Distribution in the United States. All of the United States except Florida, Alaska, and
Hawai’i. The heaviest infestations are in the northeastern states, where continental glaciation
left abundant wetland sites. West of the Mississippi River, where territory is generally dryer,
distribution is scattered and primarily found around water reclamation projects.
Description. Purple loosestrife is an erect perennial herb, 1.5–10 ft. (0.5–3 m) tall, with
4–7 stems per plant. Green to purple stems are square or octagonal, often branching to make
the plant bushy. Leaves are opposite or in whorls of three, which seem to clasp the stems.
Leaves are stalkless, lance-shaped to oval, 1.5–2.3 in. (4–6 cm) long, and heart-shaped or
rounded at the base. Leaf edges are entire, not toothed or lobed. Plants are covered with fine
hairs. Leaves on the aerial shoots die back in fall, but the woody stems remain erect for
2–3 years, contributing to the density of the stand. New shoots in spring emerge from the
rootstocks, not from the dead aerial shoots.
Purple loosestrife has both a taproot and spreading rootstock. Mature plants may grow
30–50 stems from one rootstock. The large woody taproot with rhizomes forms a dense
mat in the soil.
Very showy purple or magenta flowers, with 5–7 petals each 0.4 in. (1 cm) long, develop
all summer, from June to September. They are arranged in whorls on terminal spikes which
are as small as 3 in. (7.5 cm) or as tall as 3 ft. (1 m). The calyx of each flower is tubular and
hairy, with 12 thin vertical ridges on the outside. Flowers may also be light pink or white.
The fruits, or seed capsules, which remain on the plant during winter, are 0.1 in.
(3–4 mm) long. Each capsule contains an average of 120 tiny seeds, with 900 seed capsules
per plant.
Related or Similar Species. Cultivars of purple loosestrife and European wand looses-
trife vary little from the wild plant and can still be purchased from nurseries. Although
labeled as “infertile,” they produce viable seed and cross freely with non-cultivars of purple
loosestrife and native Lythrum species. European wand loosestrife is a perennial subshrub
that reaches 20–40 in. (50–100 cm) tall. Its glabrous stems distinguish it from purple loos-
estrife. Leaf bases are wedge shaped, and leaf margins are coarsely toothed. It also has fewer
flowers, only 2–3 per stem.
Because of the showy flower spike, purple loosestrife can be confused with native flow-
ers, such as fireweed, swamp verbena, Canada germander, or blazing star, when viewed
from a distance. Close examination, however, will reveal distinct differences for positive
identification.
Introduction History. First reported in North America in 1814, purple loosestrife was
intentionally brought to the United States in the early 1800s for both ornamental and herbal
 PURPLE LOOSESTRIFE n 415

uses, and also inadvertently in

ship ballast. The plant spreads
rapidly via canals and other
water transportation systems.
Habitat. Purple loosestrife is a
riparian plant, inhabiting fresh-
water wetlands such as lake-
shores, marshes, wet meadows,
river and stream banks, and
pond, reservoir, or ditch edges—
areas that would naturally sup-
port cattail marshes, sedge mead-
ows, and bogs. Because of its
massive seed output and seed
bank, purple loosestrife quickly
reseeds and grows onto disturbed
or degraded wetland sites, such as
exposed banks during dry
years, or where bulldozing, silt
deposition, cattle trampling, or
dredging have created bare surfa-
ces. When not found in homog-
enous stands, purple loosestrife
grows in association with broad-
leaf cattail, white cattail, common
reed (see Graminoids, Common
Reed), cordgrass, bulrushes, and
sedges. The plant tolerates many
environmental conditions, but its
one requirement is moisture. It
grows best in soils with high
organic content but is found on a Purple loosestrife, with attractive purple flower spikes, commonly grows
wide range of substrate, including in wet sites around lakes and marshes. (Native range adapted from
clay, sand, muck, and silt. Al- USDA GRIN and selected references. Introduced range adapted from
USDA PLANTS Database, Invasive Plant Atlas of the United States, and
though full sun is preferred, plants
selected references.)
survive in 50 percent shade.
Reproduction and Dispersal. Although the plant resprouts from root and stem fragments,
its major means of reproduction and spread is by seed. Attracted by abundant nectar, several
types of bees and butterflies pollinate the flowers. Because purple loosestrife has a long
flowering season, the top of the spike is often still in flower while the lower seed capsules
are mature. A large plant from a single rootstock can produce 2–3 million seeds per year.
Plants in open areas produce more seeds than plants in dense stands. When ripe during
June to September, the seed capsules burst, expelling the seeds. Seeds may remain viable
for 20 years. Ability to germinate, however, decreases after two years of storage in water.
The buoyant seeds are disseminated primarily by wind and water, but also by birds, animals,
and humans. Seeds often are embedded in mud on the feet of waterfowl and other animals
or carried on clothing. Beekeepers have been known to deliberately sow seed in order to
have a good source of nectar.
416 n FORBS

A. Lance-shaped leaves are opposite on the stem. (Steve Dewey, Utah State University, Bugwood.org.) B. Three
whorled leaves clasp the stem. (Ohio State Weed Lab Archive, The Ohio State University, Bugwood.org.)
C. The inflorescence is a terminal spike. (Eric Coombs, Oregon Department of Agriculture, Bugwood.org.)
D. The purple flowers are showy. (Norman E. Rees, USDA Agricultural Research Service, Bugwood.org.)

Seeds can germinate in any type of soil, acid to alkaline, rich or poor in nutrients. Soil
surface temperature of 59–68ºF (15–20°C), however, is critical. Established seedlings can
survive standing water up to 12–18 in. (30–45 cm) deep. As many as 1,000–2,000 seeds
can germinate in a single square foot (10,000–20,000 per m2), outcompeting native seed-
lings. As they grow, the plants crowd out or shade out native species. Germination can take
place in either spring or summer, but spring germinators have a better chance of survival.
Taproots grow rapidly when plants are seedlings. New plants can flower only 8–10 weeks
after germination. Plants that germinate in spring can produce a 12 in. (30 cm) inflorescence
the first year.
The rhizomes increase the size of stands by growing about 1 ft. (0.3 m) each year. Any
stem cutting or piece of root will sprout.
Impacts. Purple loosestrife invades both natural and disturbed wetlands, where it repla-
ces native plants and threatens the existence of rare and endangered species, such as
orchids. Its aggressive growth displaces grasses and sedges, which provide better nutrition
for wildlife. Redwing Blackbirds, however, have been noted to eat purple loosestrife seeds.
Species diversity is replaced with homogenous stands of purple loosestrife. Dense thickets
restrict access to open water and limit food and shelter for wildlife, particularly waterfowl,
because the thickets are unsuitable for either nesting or forage. Dense stands clog waterways,
irrigation ditches, and recreational water bodies, and can alter hydrology of moist meadows.
Debris builds up around the roots, enabling plants to invade deeper water and shade out
native emergent vegetation or eliminate floating vegetation by clogging waterways.
Management. Depending on objectives and available funds and personnel, purple loose-
strife can be prevented, contained, or even eradicated. It is difficult to impossible to totally
eradicate stands of three acres (1.2 ha) or more. Because the most effective herbicides are
nonselective and biological controls are still experimental, containment and minimizing
reproduction are more feasible goals than eradication.
Physical means such as hand-pulling, with as little disturbance to the soil as possible,
may be effective on small stands or at the edges of larger stands to prevent spread. Root dis-
turbance stimulates growth, and because any fragment will grow a new plant, all pieces must
be removed. Pulling or digging out plants before seed production also limits seed supply.
 SPOTTED KNAPWEED n 417

It is more difficult to control large populations. Frequent cutting to ground level over several
years may deplete root reserves and weaken the stand. All cuttings and fragments should be
burned so they cannot sprout. Burning plants without first cutting them down makes infes-
tations worse because desirable plants are also destroyed and purple loosestrife resprouts
from the root when aerial parts are damaged.
Systemic herbicides such as glyphosate and triclopyr offer effective chemical control, but
sites require frequent monitoring for regrowth. Treatment is best done late in the season
when plants are pulling nutrients into the root system in preparation for dormancy,
although midsummer applications can reduce seed production. Because these nonselective
herbicides also kill native vegetation, purple loosestrife can actually increase due to its abun-
dant seed production and new exposure of bare sites. If possible, spot treatment is best, on
stems cut down to 6 in. (15 cm). Follow-up is critical to prevent new growth, both during
the same growing season and for several years after treatment. Some plants survive, some
are missed, and new plants sprout from the seed bank. For large areas where spot treatment
is not feasible, spray with a broadleaf contact herbicide, such as 2,4-D. Most native wetland
plants are monocots and will not be harmed. Results are best when spraying is done in late
May or early June, but purple loosestrife may not be seen without its showy flowers.
Long term control of purple loosestrife requires biological methods. Three insect species
from Europe, approved by the U.S. Department of Agriculture in 1997, were released in
selected natural areas in northern states from Oregon to New York. They include a root-
mining weevil (Hylobius transversovittatus) and two leaf-feeding beetles (Galerucella calmar-
iensis, G. pusilla), which do not appear to substantially harm other plants. Experiments are
being conducted with flower-feeding beetles.
Some success has been achieved by replacing purple loosestrife with other plants. When
sown on newly emerged bare soil, Japanese millet (Echinochloa frumentaceae) can outcom-
pete purple loosestrife as long as its seeds have not begun to germinate. The Japanese millet
must be resown every year. Native curlytop knotweed (Polygonum lapathifolium) has pro-
vided similar results when sown into existing purple loosestrife stands.

Selected References
Bender, J. “Element Stewardship Abstract, Lythrum salicaria.” Global Invasive Species Team, Nature
Conservancy, 1987. http://www.invasive.org/gist/esadocs/documnts/lythsal.pdf.
“Purple loosestrife, Lythrum salicaria L.” Weed of the Week. U.S. Department of Agriculture, Forest
Service, Invasive Plants, n.d. http://www.na.fs.fed.us/fhp/invasive_plants.
Swearingen, Jil M. “Purple Loosestrife.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2005. http://www.nps.gov/plants/alien/fact/
lysa1.htm.
Zheng, Hao, Yun Wu, Jianquing Ding, Denise Binion, Weidong Fu, and Richard Reardon. “Lythrum
spp. Loosestrife.” Invasive Plants of Asian Origin Established in the US and Their Natural Enemies,
2004. http://www.invasives.org/weedcd/pdfs/asianv1/lythrum.pdf.

n Spotted Knapweed
Also known as: None
Scientific name: Centaurea stoebe
Synonyms: Centaurea biebersteinii, C. maculosa
Family: Sunflower (Asteraceae)
418 n FORBS

Native Range. Central Europe

to southwestern Russia, includ-
ing the Caucasus region
Distribution in the United
States. Widely distributed, with
the exception of Oklahoma,
Texas, Alaska, and Hawai’i.
Description. Spotted knap-
weed is a biennial or short-lived
perennial with a basal rosette of
leaves, erect branching stems,
and a deep taproot. Foliage is
gray-green and hairy. Plants usu-
ally live 3–9 years and resprout
from the root crown. During the
first year of growth, the plant
develops a basal rosette, sending
up flowering stems in sub-
sequent years. Rosette leaves are
long and narrow, 8 in. (20 cm)
by 2 in. (5 cm), wider above the
middle half and narrower
toward the base. They grow on
short stalks. Rosette leaf edges
are deeply lobed or wavy, and le-
af surfaces are rough. Flowering
stems, growing 1–4 ft. (0.3–
1.2 m) tall, are slender, multi-
branched, and either sparsely or
densely hairy. The pale-green
stem leaves are alternate and
As a contaminant in alfalfa seed, spotted knapweed rapidly became stalkless. Margins of the lower
widespread in the United States. (Native range adapted from USDA GRIN stem leaves are slightly lobed,
and selected references. Introduced range adapted from USDA PLANTS while the upper leaves are
Database, Invasive Plant Atlas of the United States, and selected references.)
linear and entire. Stem leaves are
1–3 in. (2.5–7.5 cm) long, decreasing in size toward the tip of the stem.
Plants bloom from June to October, with a shorter flowering time in more northern cli-
mates. Each plant produces 1–15 flowering stems. Flower heads, which may be solitary or
in groups of 2–3 at the ends of each branch, are 0.25 in. (6 mm) in diameter and 0.5 in.
(12.5 mm) long. Several, 25–35, pink or lavender (rarely white) tubular florets are clustered
on each head. Although seeds are shed when mature, the dry flower heads remain on the
plant. The yellow-green bracts at the base of the flower have obvious veins and are tipped
or spotted in black, giving the plant its name. Plants with white flowers usually lack the
black spots. Margins of the bracts are fringed with soft spines.
The pale-brown seeds are oval and tiny, less than 0.25 in. (6 mm) long, with a short
fringe of light-colored bristles or spines on one end.
Related or Similar Species. Several similar knapweeds and starthistles cause similar envi-
ronmental problems. Russian knapweed is a creeping perennial with black, scaly roots
 SPOTTED KNAPWEED n 419

A. Stems are freely branching. B. Rosette leaves are deeply lobed. C. Bracts at the base of the flowerhead are
distinctly spotted with black. (Steve Dewey, Utah State University, Bugwood.org.)

extending as deep as 8 ft. (2.5 m). Young plants in the spring are a rosette of silvery-green
hairy leaves with wavy edges, and flower stalks support pink or lavender cone-shaped flower
heads. Found on sites similar to where spotted knapweed invades, it can be distinguished by
its flower heads, which are smaller and lack the black spots. Russian knapweed is also alle-
leopathic, exuding toxins that prevent growth of competitors. It reproduces not only from
seed, but also vegetatively. Any piece of root will sprout a new plant, even from 3 ft. (1 m)
deep or under a plastic cover. Herbicide applications kill only the aerial portion of plants.
The plant is toxic to some animals, especially horses, which can die if they ingest too much.
Cattle and sheep are not affected.
Diffuse knapweed, a biennial growing 1–3 ft. (0.3–1 m) tall, has white, pink, or pale-purple
flowers, with 12–13 florets per head. Stem leaves are linear and entire. Straw-colored spines are
short, only 0.1 in. (3mm) long. The plant is self-fertile, producing about 13 dark-brown seeds
per flower head. Seeds are dispersed when the flower stems break and blow away in the wind.
Squarrose knapweed is a shorter perennial, 1.6 ft. (0.5 m) tall, with a woody base and a
stout taproot. Stem leaves are also linear and entire, although plants usually have no stem
leaves when they are in flower. It has 4–8 pink to pale-purple florets per head. The short,
0.01 in. (3 mm), central spines on the bracts are usually tilted downward. Each flower head
produces 1–4 pale brown seeds, and the head falls as a unit from the plant. This species is
more adaptable to drought and cold conditions.
Purple or red starthistle is an annual or perennial, growing as tall as 3 ft. (1 m). Its leaves
are resin-dotted, and the upper stem leaves are lobed or divided. New rosette leaves are
densely covered with gray hairs, and the rosette develops a circle of spines as it ages.
Flowers are distinctly reddish purple, and the thick, straw-colored spines on the bracts are
0.5–1 in. (1.3–2.5 cm) long. It grows a stout taproot and is frequently found on heavy, fer-
tile, or alluvial soils. The seeds are white, and the seed head falls as a unit from the plant.
Iberian knapweed is an annual, biennial, or short-lived perennial that resembles purple
starthistle except for its pale rose-pink or whitish flowers. Its leaves are resin-dotted, and
the stem leaves are pinnately divided. Spines are shorter than those of purple starthistle.
New leaves are covered with tiny bristly hairs. Its seeds are white. It has a stout taproot
and is common along water course banks and other moist sites.
Introduction History. Spotted knapweed was accidentally introduced to the United States
from eastern Europe in the late 1800s or early 1900s as a contaminant in alfalfa or clover
seed. Alternatively, the seeds may have been in soil used as ship ballast. First recorded in
British Columbia in 1883, the plant spread rapidly in seed shipments before it was noticed
to be an invasive problem.
420 n FORBS

Habitat. Spotted knapweed has wide environmental tolerances. It grows from sea level to
over 10,000 ft. (3,000 m) elevation and in regions with 8–80 in. (200–2,000 mm) of pre-
cipitation. An invader of both disturbed and natural land in many habitats, it grows pre-
dominantly in semiarid rangeland and pasture. It is also found in forest clearings, irrigated
areas, and agricultural fields. It grows best on well-drained soils with summer rain, and does
not compete well in moist areas. In seasonally dry areas, its deep taproot gives it an advan-
tage over other plants, enabling it to access deeper water sources. It is most invasive in dis-
turbed areas, such as gravel pits, vacant land, hayfields, and along highways, railroad
tracks, pipelines, utility lines, and waterways.
Reproduction and Dispersal. Spotted knapweed reproduces only by seed, 1,000 or more
per plant. Plants can produce 500–4,000 seeds per sq. ft. (5,000–40,000 per m2) of ground
each year, of which 90 percent are immediately viable the same or following season. The rest
of the seeds remain in the soil for 5–8 years and can re-infest a site after the initial plants
have been destroyed. Germination takes place during the entire growing season, from spring
to fall. Seedlings that sprout in the fall overwinter as rosettes, resuming growth to produce
flowering stems in subsequent years. Most seeds fall close to the plant and are dispersed
short distances by animals, in fur and in droppings, and by water. Because seeds have no
adaptations for long-distance dispersal, the plant is dependent on human influence to
spread to new territory. Seeds can be carried by livestock, on vehicles, or as contaminants
in crop seed or hay. Spotted knapweed spreads easily from gravel pits, equipment or
machine yards, grain elevators, fill dirt piles, logging yards, and railroad yards.
Although the root crowns may sprout stems, the plant does not reproduce vegetatively.
Impacts. Spotted knapweed infests several million acres in the United States. Because it is
alleleopathic, exuding a chemical that prevents other plants from growing nearby, it can cre-
ate a monoculture. It outcompetes native species, reducing biodiversity of both plants and
animals, and decreases forage potential of rangeland. Economic impact due to loss of forage
is estimated to be in the millions of dollars. The taproot replaces intertwined native grass
roots, reducing both the stability of the soil and its water-holding capacity. Dense infesta-
tions contribute to soil erosion and sedimentation caused by increased runoff. The plants
also lessen the visual attractiveness of recreational lands, and dry plants pose a fire hazard.
Management. The most effective means of control is to prevent the spread of seeds.
Livestock grazing should be limited when seeds are ripening. Clothing, all hiking gear,
vehicles, and equipment should be thoroughly cleaned. Transport of potentially contami-
nated hay should be prevented. Existing infestations are best controlled by using a combina-
tion of physical, chemical, and biological means. Competitive range plants, both grasses and
forbs, may help control spotted knapweed, but revegetation or reseeding is best done when
the spotted knapweed is stressed from prior treatments.
Control of small infestations may be attempted by physical means, such as hand-pulling
plants before seed sets. The entire root crown and taproot must be pulled out to prevent
resprouting. For best results, the area should be treated with a herbicide after hand removal.
Mowing stems before they flower can reduce seed production, and long-term grazing by
sheep and goats is sometimes successful in suppressing infestations. Livestock will eat it
when nothing else is available.
Chemical means of control have met with limited success. Herbicide applications that
control all plant stages, including mature plants, seedlings, and seeds in the soil, are best.
Because of the long-term viability of seeds in the soil, reduction in the seedbank should be
a goal. The herbicide 2,4-D will kill existing plants, but needs to be reapplied yearly to elimi-
nate seedlings that sprout from the soil seed bank. Picloram can control seedling growth for
 TOADFLAX n 421

2–5 years, but may contaminate groundwater in porous soils or in areas with a shallow water
table. Dicamba is less effective than picloram in preventing seed germination and also kills
other broadleaf plants. Although clopyralid is also effective in controlling spotted knapweed
and is a more selective herbicide; it will harm clovers, which are often desirable range plants.
The best times for herbicide application are on the rosettes in the fall and at the bud or
bloom stage in spring.
Although it may take decades to suppress or kill spotted knapweed on a large site, bio-
logical control, using several insects from the plant’s native range in Eurasia, is most efficient
for extensive infestations. Insect larvae from spotted knapweed’s natural enemies feed on
and damage the root, leaves, or flowers. From 1973 to 1992, 12 insects from Eurasia have
been released in North America. Insects selected for biological control are host-specific
and should not affect native plants. Although impact on natives has not been researched,
no incidences have been reported of nontarget plant species being attacked. Several species
are well established, with individual success related to location and climate. Especially effec-
tive are two seed head flies (Urophora affinis and U. quadrifasciata), which can reduce seed
production by 50 percent. Larvae of a seed head weevil (Larinus minutus) and a moth
(Metzneria paucipunctella) also damage growing seeds. Larvae of a root-boring weevil
(Cyphocleonus achates) burrow into roots and feed all winter, spring, and early summer,
depleting the plant’s resources. Insects weaken plants and reduce their competitive ability.
Economic benefits are not yet realized, but reductions in knapweed have been observed.
Research is ongoing, with no definitive results, but a combination of insects for biological
control appears to be more effective than just one species.

Selected References
“Centaurea Genus Part I.” Plant Pest and Health Prevention Services (PHHPS), California Department
of Food and Agriculture, n.d. www.cdfa.ca.gov/phpps/ipc/weedinfo/centaurea.htm.
Lym, Rodney G., and Richard K. Zollinger. “Spotted Knapweed (Centaurea maculosa Lam.).” North
Dakota State University Extension Service, Fargo, ND, 1992. http://www.ag.ndsu.edu/pubs/
plantsci/weeds/w842w.htm.
“Russian Knapweed.” Colorado State University Cooperative Extension Tri River Area, Grand Junction,
CO, 2004. http://www.coopext.colostate.edu/TRA/knapweed.html.
“Spotted Knapweed, Centaurea biebersteinii DC.” Chequamegon-Nicolet National Forest, U.S.
Department of Agriculture, Forest Service, 2005. http://www.fs.fed.us/r9/cnnf/natres/nnis/spotted
_knapweed.html.
Story, J. “Spotted Knapweed.” In: Biological Control of Invasive Plants in the Eastern United States, by R.
Van Driesche et al. U.S. Department of Agriculture, Forest Service Publication FHTET-2002-04,
Morgantown, WV, 2003. http://www.invasive.org/eastern/biocontrol/13Knapweed.html.

n Toadflax
Dalmatian Toadflax
Also known as: Broadleaf toadflax, broad-leaved toadflax, wild snapdragon
Scientific name: Linaria dalmatica ssp. dalmatica
Synonyms: Linaria genistifolia ssp. dalmatica
Family: Figwort (Scrophulariaceae)
422 n FORBS

Yellow Toadflax
Also known as: Common
toadflax, wild snapdragon, per-
ennial snapdragon, flaxweed,
butter and eggs, eggs and bacon,
Jacob’s ladder, rabbit-flower
Scientific name: Linaria vulgaris
Synonyms: None
Family: Figwort
(Scrophulariaceae)
Native Range. Dalmatian
toadflax is native to the Mediter-
ranean region of southeastern
Europe, extending into Turkey
and Afghanistan in Southwest
Asia. Yellow toadflax is native
to central and southern Europe
and to Turkey. It may also be
native to the British Isles,
Russia, Siberia, and China.
Distribution in the United
States. Dalmatian toadflax is com-
mon in the northern states, but is
problematic in the Rocky Mount-
ain and western desert regions. It
is not found in Alaska, Hawai’i,
or in the southeastern states.
With the exception of Hawai’i,
yellow toadflax is found through-
out the United States, including
southern Alaska. It is more com-
Dalmatian toadflax is found on coarse-textured, well-drained soils, pre-
dominantly in western states. (Native range adapted from USDA GRIN
mon in eastern North America
and selected references. Introduced range adapted from USDA PLANTS and is scattered in the West.
Database, Invasive Plant Atlas of the United States, and selected references.) Description. Toadflaxes are
short-lived herbaceous perenni-
als that grow 1–25 vertical flowering stems, which are thick-walled and fibrous. Although
leaves are alternate, crowding on the stem may make them appear to be opposite.
Plants have both a long taproot, usually 4 ft. (1.2 m) deep into the soil, and creeping hori-
zontal roots, at a depth of 2–12 in. (5–30 cm), which can grow as far as 12 ft. (3.6 m) from
the parent plant.
Flowers, which resemble snapdragons, develop from May to August. They are bright yellow,
with a distinctly two-lipped yellow corolla. A bulge in the lower lip, called the palate, which almost
closes the throat of the flower, is hairy and orange. Because flowers mature successively from the
lower stem to the top, several stages of flower and fruit can be on one plant or inflorescence.
Fruits are two-celled capsules, which grow upright on the stem. Seeds mature from July to October.
Dalmatian Toadflax: Dalmatian toadflax stems grow to 2–5 ft. (0.6–1.5 m) tall, and are
stiff and leafy. Plants are usually scattered rather than clumped, although they may be
 TOADFLAX n 423

connected beneath the surface.

Plants are robust, with waxy
stems that branch toward the
top of the plant. The stems
become rough and woody at
the base, while the upper
portions are smooth and more
herbaceous. Plants develop
prostrate stems in September,
which are tolerant to freezing
and remain green beneath the
snow in winter. These branches
give rise to floral stems the fol-
lowing summer. The lower
leaves, oval or heart-shaped
and pointed, are 1–2.5 in.
(2.5–6 cm) long and almost as
wide. They wrap around or
clasp the stem. Upper stem
leaves are smaller and oval to
lance-shaped. All leaves are blu-
ish green and waxy. Flowers
develop in long, loose racemes
at the ends of the stems, with
each flower supported by a
short pedicel growing from a
leaf axil. Flowers are 0.5–1 in.
(1.4–2.5 cm) long, with a
curved spur, 0.4–0.7 in. (1–
1.7 cm) long. Each egg-shaped
to round fruit capsule, approxi-
mately 0.2–0.4 in. (4–10 mm) Yellow toadflax, growing in moist sites, is common in the eastern states
in diameter, contains 60–300 and a frequent invader of cropland. (Native range approximated from
seeds. The small, 0.04–0.08 in. USDA GRIN and selected references. Introduced range adapted from
USDA PLANTS Database, Invasive Plant Atlas of the United States, and
(1–2 mm), black seeds are
selected references.)
sharply triangular, with a slight
papery wing on each edge. Dead flower stalks may remain standing for 2 years.
Yellow Toadflax: Yellow toadflax is a shorter plant, with smooth stems 1–3 ft. (0.3–1 m)
tall, growing in crowded patches or clumps. Stems are generally sparsely branched or not
at all. They are usually reddish at the base, and become more slender and green toward
the tips. The pale-green leaves are linear to lance-shaped, 1 in. (2.5 cm) long and less than
0.2 in. (0.5 cm) wide, pointed or tapering at both ends. They are sessile, do not clasp the
stem, and are not waxy. Yellow toadflax does not produce overwintering prostrate stems, and
all above-ground parts die back in winter. Flowers of yellow toadflax form in dense terminal
clusters of 6–30, which continue to grow and become longer as new flowers develop. Flowers
are approximately 1 in. (2.5 cm) long, including the short spur. Each stem may have as many
as 30 globular seed capsules, each containing as many as 250 seeds. Yellow, dark-brown, or
black seeds, 0.04–0.08 in. (1–2 mm), are flattened disks, with a distinctly winged edge.
424 n FORBS

A. Heart-shaped leaves on prostrate stems of Dalmatian toadflax wrap around the stem. (Bonnie Million, BLM,
Ely District, Bugwood.org.) B. Upper stem leaves of Dalmatian toadflax are lance-shaped. (Steve Dewey, Utah
State University, Bugwood.org.) C. Dalmatian toadflax flowers grow in loose racemes. (Joseph M. DiTomaso,
University of California–Davis, Bugwood.org.) D. Pedicels of Dalmatian toadflax flowers emerge from leaf axils.
(Joseph M. DiTomaso, University of California–Davis, Bugwood.org.) E. Yellow toadflax plants are shorter than
those of Dalmatian toadflax. (L. L. Berry, Bugwood.org.) F. Sessile leaves of yellow toadflax are linear and do
not clasp the stem. (Bonnie Million, BLM, Ely District, Bugwood.org.) G. Yellow toadflax flowers grow in dense
terminal clusters. (Steve Dewey, Utah State University, Bugwood.org.) H. Yellow toadflax flowers grow close to
the stem. (Richard Old, XID Services, Inc., Bugwood.org.)

Related or Similar Species. Because Dalmatian toadflax and yellow toadflax are highly
variable in shape, size, and color, hybrids may have developed between the two. A possibil-
ity exists that the narrow-leaf form of Dalmatian toadflax is a separate invasive from Europe,
broomleaf toadflax. Canada toadflax, also called blue toadflax, is native to the United States
and easily distinguished by its purple or blue flowers.
Introduction History. Both species of toadflax were introduced as ornamentals because of
their showy flowers. Yellow toadflax, a plant used in Germany for yellow dye, was brought
to Delaware from Wales in the mid-1800s. Immigrants, especially Mennonites, cultivated
it for dying yarn and cloth. Dalmatian toadflax was brought to the western United States in
the late 1800s. Widely available in the horticultural trade under several names, toadflaxes
are still used as garden plants.
Habitat. Toadflaxes are quick to invade bare areas, especially when perennial plants are
removed. Because the taproot efficiently taps deep moisture, toadflaxes are well adapted to
arid sites and are most competitive in dry-summer climates. Both species prefer sunny sites,
tolerate low temperatures, and thrive in open, disturbed habitats. Dalmatian toadflax does
best on well-drained, coarse-textured soils, such as road and railroad rights-of-way, gravel
pits, waste areas, abandoned lots and fields, clear cuts, and degraded rangelands. It also
grows in low-elevation coniferous forests and with oak and aspen at 5,000–6,500 ft.
(1,500–2,000 m) elevation. Although generally found on similar soils and sites, yellow
 TOADFLAX n 425

toadflax requires more fertile soils with more moisture, and can invade higher elevations,
such as upland meadows at 6,000–8,500 ft. (1,800–2,600 m) elevation. It is also a more
common weed in cropland.
Reproduction and Dispersal. Both species reproduce both sexually and vegetatively.
Infestations usually start by seed, but expand from the root system. Both species require
cross-pollination with another plant, usually done by large bees. Although most seeds drop
from the plant in the year they mature, some seed may remain on the dead stem. One
Dalmatian toadflax plant can produce as many as 500,000 seeds in a season, with 75 percent
germination. Yellow toadflax, is less prolific, as many as 30,000 seeds per plant, with a ger-
mination rate of 10 percent. Yellow toadflax capsules that mature later in the season, how-
ever, have a higher percentage of viable seeds. Wind is the primary dispersal agent for the
winged seeds, which can even be blown across encrusted snow. Broken stalks may be blown
as a unit. Seeds are also dispersed by livestock, deer, and other browsing animals.
Seeds, which may remain viable in the soil for as long as 10 years, germinate in both fall
and spring. Seedlings, with lance-shaped leaves in both species, grow rapidly. Dalmatian
toadflax may develop a 20 in. (50 cm) long taproot within eight weeks and 2–5 flowering
and prostrate stems in the first season. In the second and subsequent years of growth, a plant
can produce as many as 25 flowering stems and 40 prostrate stems, all of which grow in a
loose rosette at the base of the plant. Yellow toadflax seedlings grow a taproot in 2–3 weeks.
Regeneration in spring can be as early as February.
The horizontal root system has adventitious buds which produce new plants. Root buds
and lateral roots may begin to grow 2–9 weeks after the seed has germinated. The lateral root
system of one yellow toadflax plant can produce as many as 100 shoots in the first summer.
In one study, yellow toadflax increased its spread over 400 percent in one season. A patch
originally covering 1 acre (0.4 ha) increased to 85 acres (34 ha) in five years. Buds on sev-
ered or broken roots will grow, and fragments as short as 0.4 in. (1 cm) can produce new
plants. Root bud sprouts grow a separate root system and can become independent plants
in just one year. Individual plants and roots generally live for 3–5 years, but 13-year old
infestations may occur under good conditions.
Impacts. Both toadflax species are short-lived, but aggressive invaders. Although one
plant may do little damage to crops or to natural areas, the threat of toadflax is its potential
to spread. Crop yields may be significantly reduced and native communities stressed or dis-
placed. One study showed that a pasture plot without toadflax had 2.5 times more grass
than an infested plot. Once established, toadflax outcompetes crops, pasture grasses, and
native plants and becomes the dominant vegetation. The taproots and early spring growth
deplete the soil of water and nutrients needed by crops, forage, or native species.
Although both species contain a toxic glucoside harmful to cattle in large quantities, no
poisonings have been reported because the plants are not palatable to livestock. The toxins
do not affect sheep and goats.
Management. Eradication of toadflaxes is difficult because of their deep, robust root sys-
tems and abundant seed production. The genetic diversity of both species complicates con-
trol. Because toadflax seedlings are poor competitors against established perennials and
winter annual crops, maintenance of desirable grasses in pastures and rangeland will help
prevent infestations. The plants, however, can invade even small openings in quality range.
Once toadflax is established, the primary goal should be to decrease or eliminate seed pro-
duction. Seeds and broken root pieces may be spread by farm machinery and outdoor rec-
reational equipment that is not thoroughly cleaned. Livestock using infested areas should
be held 6–11 days before being transported elsewhere.
426 n FORBS

Seedlings are the most vulnerable to physical removal. Although small plants are fairly
easily pulled, the root system of large plants provides a solid anchor. Hand-pulling plants
before seeds mature, including all fragments of roots, every season for 5–6 years will deplete
the seed bank. Mowing plants will prevent seed development, but root buds will produce
new plants. Mowing for approximately 10 years, however, may deplete the seed bank.
Repeated cultivation, 7–10 times annually for two years, will disrupt new plants as they
sprout from broken roots. Prescribed fire is a poor choice. Although burning prevents seed
production, it fails to kill either the roots or the seed bank and opens more bare ground for
invasion. Grazing is not recommended because cattle avoid the plant and their hooves dis-
turb the ground, creating more potential habitat.
Toadflax cannot be controlled by chemical applications alone. Effectiveness varies due to
different climates, waxy leaves, and genetic variation in the two species. Spraying is most effec-
tive when done at the flowering stage, when root reserves are lowest. Because both picloram
and dicamba can have a residual effect on nontarget, broad-leaved perennials, glyphosate is
the best choice. Yellow toadflax is more resistant to chemicals than is Dalmatian toadflax.
Several insects, most of which attack both toadflax species, offer potential biological con-
trol. Three species were accidentally introduced to the United States on infected plants. A
flower-feeding beetle (Brachypterolus pulicarius) was first identified on yellow toadflax in
the 1920s in New York. Both adults and larvae feed on flower buds and pollen and can
decrease seed production by 74 percent. Two seed-feeding weevils (Rhinusa antirrhini and
R. neta) were discovered in Canada in the late 1950s. Adults feed on a variety of foliage parts,
while larvae eat the developing seeds. R. antirrhini feeds primarily on yellow toadflax and can
reduce seed production by 85–90 percent.
Five species have been intentionally released, a defoliating moth in the early 1960s, the
remainder in the 1990s. Larvae of the defoliating moth (Calophasia lunula) eat the foliage,
weakening the plant, but the insect does not adapt to cold temperatures. Larvae of two
root-boring moths (Eteobalea intermedia and E. serratella) feed on root tissue, also weaken-
ing the plant and reducing seed production. Adults of the root-galling weevil (Rhinusa
linariae) feed on foliage and stems, while the larvae eat the tissue inside the root gall,
depriving the roots of nutrients and water. Both adults and larvae of the stem-mining wee-
vil (Mecinus janthinus) feed on tissues inside the stems, reducing flowering, seed produc-
tion, and shoot biomass. Severe stem damage can kill the plant. No one insect, however,
completely destroys toadflax, and one species cannot adapt to the wide range of climate
where toadflax grows.

Selected References
Butler, M. D., and L. C. Burrill. “Yellow Toadflax and Dalmatian Toadflax.” Weeds. Pacific Northwest
Extension Publication 135, 1994. http://extension.oregonstate.edu/catalog/pdf/pnw/pnw135.pdf.
Carpenter, Alan T., and Thomas A. Murray. “Element Stewardship Abstract, Linaria genistifolia ssp. dal-
matica and Linaria vulgaris.” Global Invasive Species Team, Nature Conservancy, 1998. http://
www.invasive.org/gist/esadocs/documnts/linadal.pdf.
Erskine, Jennifer A., and Mark J. Renz. “Dalmatian Toadflax.” New Mexico State University Weed-
Factsheet, 2005. http://weeds.nmsu.edu/downloads/dalmatian_toadflax_factsheet_11-06-05.pdf.
Wilson, L. M., S. E. Sing, G. L. Piper, R. W. Hansen, R. De Clerck-Floate, D. K. MacKinnon, and
C. Randall. “Biology and Biological Control of Dalmatian and Yellow Toadflax.” U.S. Department
of Agriculture, Forest Service, FHTET-05-13, 2005. http://www.fs.fed.us/foresthealth/technology/
pdfs/Toadflax.pdf.
 YELLOW STARTHISTLE n 427

n Yellow Starthistle
Also known as: Golden starthistle, yellow cockspur, St. Barnaby’s thistle
Scientific name: Centaurea solstitialis
Synonyms: Leucantha solstitialis
Family: Sunflower family (Asteraceae)
Native Range. Mediterranean regions of southern Europe, northern Africa, and the
Balkan peninsula. Also native to the desert regions of Southwest Asia, From Turkey to as
far east as Pakistan.
Distribution in the United States. Most of the United States, except some southeastern
states, parts of New England, Alaska, and Hawai’i. Especially invasive in western states,
and still expanding its range.
Description. Yellow starthistle is usually an herbaceous winter annual, but can be bien-
nial in mild climates. Plant morphology is variable. Young plants first grow as rosettes,
where leaves are 2–6 in. (5–15 cm) long, with deep lobes reaching to the midrib of the leaf.
As the plant matures, several stiff flowering stems emerge from the basal rosette, generally
growing to 3.3 ft. (1 m) but sometimes as tall as 6.5 ft. (2 m). The basal leaves wither and
die as flowering stems grow. Stem leaves are short and linear, 0.5–1 in. (1.2–2.5 cm) long.
They may be smooth, toothed, or with wavy margins. Stem leaves are opposite and appear
gray-green or blue-green due to a dense covering of fine white hairs or cottony wool. Leaf
bases of the lower leaves extend down the stems, making them look winged. Plants have a
long taproot, normally 3.3 ft. (1 m) but occasionally as deep as 6 ft. (1.8 m).
Flowering occurs all summer, until plants are killed by frost. The 0.5 in. (1.25 cm) flower
heads support a cluster of 30–100 bright yellow tubular florets, 0.5–0.75 in. (1.3–2 cm) long,
in a terminal arrangement on the ends of the branched stems. Bracts at the base of the flowers
are spiny, with a stiff central spine usually 0.5–1 in. (1–2.5 cm) long, but sometimes as long as
2 in. (5 cm). The central spines are yellowish or straw-colored. Flowers require cross-
pollination by insects and are an important source of nectar for honey bees. Each seed head
has 35–80 tiny seeds, 0.08–0.15 in. (2–4 mm) long, which mature as the yellow flowers fade
in late summer. Most seeds (75–90%) are produced from flowers in the center of the flower
head and disperse when dried flowers become detached. Those seeds are smallest and have
slightly barbed tufts of short white hairs or bristles. Seeds from the periphery of the flower
head, slightly larger and hairless, remain attached to the edges of the flower head into fall
and winter. Plants dry in late summer or early fall, shedding the spines, but leaving dense,
fuzzy gray hairs on the flower heads. The old stems decay slowly and remain standing.
Related or Similar Species. Although two species with yellow flowers resemble yellow
starthistle, they have other distinguishing characteristics. Several are also listed as noxious
or invasive in various states. Leaves of the Malta starthistle are covered with stiff hairs, and
the spines surrounding the flower heads are slender and purplish. Grayish to tan seeds have
a fine pubescence. Dried flower heads retain the spines but shed the wooly hairs. It is not
known to cause chewing disease. Sicilian starthistle is not as invasive as other starthistles,
and populations remain small. Rosette leaves are not as deeply lobed, and stem leaves are
toothed. The central spines around the flower head are straw-colored at the top and black
or dark brown at the base. Seeds are glossy dark brown with dark brown or black bristles.
Both the central spines and the fuzzy gray wool are retained on the dried flower heads.
Taproots of both Malta starthistle and Sicilian starthistle do not extend as deeply as those
of yellow starthistle.
428 n FORBS

Two species have reddish-

purple or lavender flowers and
spines in the center of the
rosette leaves. Purple or red
starthistle has purple flowers,
The flower heads are sur-
rounded by long, thick white
or straw-colored spines, which
remain on the bracts at the end
of the season. Seeds are white
and lack hairs or bristles. Flow-
ering stems have a mound-like
form as tall as 4 ft. (1.2 m).
Iberian starthistle also has a
mounding habit, and the lower
leaves are longer, 4–8 in. (10–
20 cm) Flowers are rosy-
lavender and white, and spines
are straw-colored. It more often
colonizes moist sites such as
streambanks.
Russian knapweed is distin-
guished by its purplish thistle-
like but spineless flowers. This
perennial has woody stems up
to 3 ft. (1 m) tall, and its rhi-
zome system is invasive. Gray
hairs covering the leaves and
stems give it a gray-green
appearance. This species also
negatively affects horses.
Yellow starthistle is a major problem in rangeland and pasture in the Introduction History. Yellow
western United States. (Native range adapted from USDA GRIN and starthistle was introduced to
selected references. Introduced range adapted from USDA PLANTS California from Chile during
Database, Invasive Plant Atlas of the United States, and selected the Gold Rush in the mid-
references.) 1800s, probably as a seed con-
taminant in alfalfa. After 1903, starthistle was probably introduced several times in alfalfa
shipped directly from Europe or from Southwest Asia. Infestations remained localized
around the Sacramento River delta until alfalfa cultivation expanded between 1870 and
1905. Increased use of mechanization, such as tractors, contributed to its rapid spread in
the Pacific Northwest in the 1920s. Plants invaded the grazed rangeland of the California
foothills during the 1930s and 1940s. Three factors have contributed to its rapid spread
since the 1960s—road building, suburban development, and more ranching. It has since
spread from California to the Atlantic coast.
Habitat. Yellow starthistle grows best in deep, well-drained soils under full sun, and does
not do well in shady locations under trees or shrubs. It thrives in disturbed areas such as
rangeland, pasture, hayfields, orchards, vineyards, and along waterways and roadsides.
Although it can survive a wide range of precipitation, 10–60 in. (250–1,500 mm), yellow
 YELLOW STARTHISTLE n 429

A. Several flowering stems grow from the base of the plant. (Steve Dewey, Utah State University, Bugwood.org.)
B. Leaf bases appear to have “wings.” (Steve Dewey, Utah State University, Bugwood.org.) C. Rosette leaves are
deeply lobed. (Steve Dewey, Utah State University, Bugwood.org.) D. The bracts are very spiny, including a
long, stiff central spine. (Steve Dewey, Utah State University, Bugwood.org.) E. Flower heads grow at the ends
of branches. (Charles Turner, USDA Agricultural Research Service, Bugwood.org.)

starthistle does best in climates with summer drought and is less dense in summer-rain areas
east of the Rocky Mountains. Plants are found as high as 6,000 ft. (1,800 m) elevation.
Reproduction and Dispersal. Yellow starthistle reproduces only by seed, 95 percent of
which are viable. An average plant can produce 1,000 seeds in one season, and large plants
can produce as many as 75,000 seeds. Heavy infestations can produce 50–100 million seeds
per acre (125–250 million per ha). Seeds fall only 2 ft. (0.6 m) from the parent plant, and
natural spread is slow because the bristles are too coarse for wind transportation. Animals
and humans carry the bristled seeds short distances, trapped in fur or clothing. Human
activity, such as road maintenance equipment, other vehicles, and contaminated hay and
seed, is responsible for long distance dispersal. Seeds buried in the soil, as many as 2,900
per sq. ft. (29,000 per m2) are viable for 10 years, but viability of those on the surface is
3–5 years. Although most seeds germinate the year after they mature, they are capable of ger-
mination just one week after ripening.
Germination is best under warm, 50–68ºF (10–20ºC), moist, conditions with plenty of
light, usually in fall after the first rainfall in dry-summer climates. New plants emerge after
each rainfall event. Young plants quickly establish a taproot, which enables the plant to
access deeper water during the dry summer and fall. Fall seedlings overwinter as a rosette,
while roots continue to grow.
Impacts. Infesting 15–20 million ac. (6–8 million ha) of range and wildland in California
alone, yellow starthistle is a rapid colonizer that quickly develops into dense stands, displac-
ing native plants. Yellow starthistle is alleleopathic, exuding chemicals that prevent growth
of other plants nearby. The result is a reduction in biodiversity and less forage for both
native animals and livestock, which reduces rangeland values. Infestations also fragment
sensitive habitat or rare plant locations. Deep taproots deplete soil moisture, damaging both
annual and perennial grasslands. Water use by the plants is a serious problem in the arid
western states, especially those with summer dry seasons. The spiny thickets of yellow star-
thistle also lower the value of recreation land.
Grazing animals usually avoid areas infested with the spiny plants, putting more pressure
on other parts of pasture or range. Spines may also damage the eyes of grazing animals.
430 n FORBS

Yellow starthistle’s most significant impact is on horses, costing millions of dollars. Repeated
ingestion by horses (over 30–60 days) causes a disease, nigropalallidal encephalomalacia,
which results in a softening of a portion of the brain. It is commonly called “chewing dis-
ease” because it affects the animal’s ability to chew or swallow. The disease does not appear
to affect cattle, sheep, mules, or burros. It can be a serious problem when little other forage is
available or when yellow starthistle is a major contaminant in dried hay. Some horses, how-
ever, develop a taste and prefer starthistle over other feed, even when the plants are dry and
spiny. The lethal dose is 5–5.7 lb. (2.3–2.6 kg) of green plants per 220 lb. (100 kg) of body
weight per day. Russian knapweed is similar but more toxic. Symptoms, which can result
from either green or dried plants, appear abruptly. The toxin affects the nervous system, trig-
gering abnormal behavior, such as violent head tossing, lip twitches, tongue flicking, or
excessive yawning. The horse may hold lips open with tongue hanging out. Mouth muscles
become weak, and the horse loses the ability to chew or drink. Advanced stages include
brain lesions and mouth ulcers. Unable to eat or drink, the horse loses weight and dies from
starvation or dehydration, or requires euthanization. Parts of the brain die and do not regen-
erate. No treatment exists.
Management. Because of the difficulty of removing dense infestations of these spiny
plants, the best control is prevention. Inspection and removal of seeds from livestock,
clothes, and vehicles is effective but labor intensive. Once plants have become established,
any effective control must suppress seed production and requires more than one year of
effort. After eradication or suppression of yellow starthistle, areas should be revegetated with
perennial grasses or legumes. A combination of methods works best.
Physical removal of the plants or the seeds can be done in several ways. Plants in small
infestations may be pulled, but even immature flower heads must be disposed of carefully
because they might produce seeds. All of the stems must be removed because even a 2 in.
(5 cm) piece of stem attached to a root may regrow. Grazing or mowing reduces biomass
and limits seed production, but timing of those activities is critical. Grazing or mowing at
the wrong time may stimulate bolting to flowering stage. Mowing before seed sets, as long
as all stem tips and leaves are removed, may be effective, but mowing too early will cause
plants to resprout. Mowing or grazing too late may increase seed dispersal as seeds become
trapped in equipment or animal fur. Sheep, goats, and cattle will graze the plants in early
spring before the spines develop, and goats will also feed on spiny plants. Livestock may also
eat the more desirable plants, thus removing competition. Burning for three successive years
is effective and best done when the flowers appear. Yellow starthistle plants will still be
green, but associated grasses in dry-summer climates will be dry enough to carry the fire.
Fire does not affect the soil seed bank, but it reduces thatch and returns nutrients to the soil.
Repeated tillage may be effective along roadsides, but in pasture or rangeland, it disturbs and
exposes more soil to invasion.
Application of systemic herbicides in winter when yellow starthistle plants are in the
rosette stage of growth is an effective means of chemical control. Clopyralid controls yellow
starthistle for one season, while picloram has more soil residual and suppresses growth for
2–3 years. Although clopyralid does not harm grasses, it can damage some nontarget species
in the sunflower and legume families. Nonselective herbicides, such as glyphosate, are use-
ful for spot treatment on bolted plants. Eliminating or reducing the number of plants
encourages growth of native annuals.
Biological control may be accomplished by insects, which damage the seed head and
decrease the number of seeds. Six insects, three weevils, and three flies now have established
populations in North America. Larvae of the seed-head weevil (Bangasternus orientalis), hairy
 YELLOW STARTHISTLE n 431

weevil (Eustenopus villosus), flower weevil (Larinus curtus), seed-head fly (Urophora siruna-
seva), peacock fly (Chaetorellia australis), and false peacock fly (Chaetorellia succinea) attack
yellow starthistle flowers where the larvae feed on seed heads. The most successful agents
in California are the hairy weevil and the false peacock fly, which reduce seed production
by 43–76 percent. The Mediterranean rust fungus (Puccinia juncea var. solstitialis), which
attacks foliage and young plants, was introduced into California in 2003, but effects are
not yet conclusive. Tests showed that native plants were not susceptible to Mediterranean
rust. Other fungi are also being tested.

Selected References
“Centaurea Genus Part 2.” Plant Pest and Health Prevention Services (PHHPS), California Department
of Food and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/centaurea2.htm.
DiTomaso, Joe. “Element Stewardship Abstract, Centaurea solstitialis L.” Global Invasive Species Team,
Nature Conservancy, 2001. http://www.invasive.org/weedcd/pdfs/tncweeds/centsol.pdf.
Murphy, Alicia. “Yellow Starthistle.” Weeds Gone Wild: Alien Invasive Plant Invaders of Natural Areas.
Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/
fact/ceso1.htm.
“Yellow Starthistle.” Plumas-Sierra Noxious WEEDS Management Group. California Department of
Food and Agriculture, n.d. www.cdfa.ca.gov/wma.
n Graminoids
n Asiatic Sand Sedge
Also known as: Japanese sedge
Scientific name: Carex kobomugi
Synonyms: None
Family: Sedge (Cyperaceae)
Native Range. Coastal northeastern Asia, China, Korea, southern Japan, and Taiwan.
Distribution in the United States. Atlantic coastal regions from Massachusetts and Rhode
Island south to North Carolina, and in Oregon.
Description. Asiatic sand sedge is a perennial sedge growing 4–12 in. (10–30 cm) tall.
Characteristic of the sedge family, the stems are solid and triangular in cross-section. The
yellow-green, slightly arching leaves, about 0.1–0.25 in. (3–6 mm) wide, are coarse and some-
what stiff. Small marginal teeth, which can be seen with a 10x hand lens, give the leaves a rough
texture. The stem base is covered with brown scales. Older basal leaves are slightly wider,
darker green, and leathery. Leaf color becomes more yellow and brown in spring and fall.
The extensive rhizome system, extending many feet horizontally but just below the
surface of the sand, enables the plant to form extensive colonies. New shoots arise from
the rhizomes, which may be 0.25 in. (6 mm) in diameter and interconnected in the stand.
The tips of rhizomes are sharply pointed and can puncture skin. Long roots can extend
several feet deep into the sand.
Plants can be difficult to distinguish among other vegetation, but are more apparent when
flowering in spring, April through June. Asiatic sand sedge is paradioecious, which is slightly
different from dioecious. While male and female flowers occur on different plants, either gen-
der may develop on any plant, even on separate stems that are part of the same rhizome sys-
tem. The inflorescence, about 2 in. (5 cm) long, is a dense spike at the end of the flower
stalk, which is usually shorter than the leaves. Male flowers can be identified by white strands
of pollen. Female flowers have a seedhead, which is a triangular set of spikes and brown
scales. Each female flower is enclosed in a dark-brown, papery sac called a perigynium, which
develops a triangular achene, or hard fruit containing one seed, 0.15–0.25 in. (4–7 mm).
Related or Similar Species. Two native grasses may superficially resemble Asiatic sand
sedge, but because they are grasses, the stems are round in cross-section. Both American beach
grass and beach panic grass, also known as bitter panicum, have leaves that are two-ranked,
meaning they are on opposite sides of the stem. Their leaves also have a bluish-green cast.
The inflorescence of American beach grass is much larger, 4–15 in. (10–40 cm) long.
Large-headed sedge is native to coastal Russia, Japan, and western North America from
Oregon north to Alaska. A newcomer to the East Coast, it has recently been found on
dunes in coastal New Jersey. It can be distinguished by a more sparse growth pattern, a
maximum of 4–9 shoots per sq. ft. (40–90 per m2) compared with Asiatic sand sedge with
20–50 per sq. ft. (200–500 per m2). It has longer, greener leaves, larger seedheads, and darker
seeds with a sharp point that can penetrate flesh. Although the two species do not grow
together in their native ranges, possible hybridization is a concern because both are wind-
 ASIATIC SAND SEDGE n 433

pollinated. Hybrids are usually

more aggressive invaders, which
could pose a new problem to
coastal sand dune environments.
Other sedge species (Cyperus
spp.) flower in late summer to
early fall. They have smooth
leaves, without serrations. They
also are either weakly rhizoma-
tous or not at all.
Introduction History. Asiatic
sand sedge was first reported at
a beach park in New Jersey
in 1929. By the 1940s, it had
spread to the Delmarva Penin-
sula. Speculation that the sedge
was used as packing material for
Oriental ceramics is likely just a
story. Asiatic sand sedge would
not make good packing material
because it is not spongy for
cushioning and it has sharp
edges. More likely, seeds were
incorporated into sand, which
was used as ballast in empty
ships sailing from Japan. Plants
were intentionally distributed
along the East Coast as early as
the 1930s to bind sand dunes
for erosion control. It was delib-
erately propagated and planted
in the 1960s and 1970s. By Except for isolated sites in Oregon, Asiatic sand sedge is limited to Mid-
1977, the Cape May Plant Atlantic coastal regions. (Native range adapted from USDA GRIN and
Materials Center in New Jersey selected references. Introduced range adapted from USDA PLANTS
was distributing 20,000 plants Database, Invasive Plant Atlas of the United States, and selected references.)
per year. Because of increasing
realization of the plant’s invasive qualities, sales were stopped in the mid-1980s. Ironically,
Asiatic sand sedge was listed as an endangered species in the 1970s. Now, it is one of the
10 most unwanted plants in New Jersey.
Habitat. Asiatic sand sedge colonizes coastal beaches and sand dunes, specifically the
primary dunes, the first dunes landward from the coastline, but may also be found on
coastal sand pits. Storm disturbance allows Asiatic sand sedge to gain a foothold. The
species tolerates salt spray and high winds, but inundation by storm surges may kill
growing plants.
Reproduction and Dispersal. It is not clear how Asiatic sand sedge disperses to new loca-
tions. Some populations rarely bloom and set seed, the germination rate is low, and seed-
lings are rare. Because plants are paradioecious, both male and female flowers may not
occur in the same vicinity. Pollen, however, is easily transported over long distances by
434 n GRAMINOIDS

A. Infestations can cover the entire beach. (Louise Wootton, Georgian Court University, New Jersey.) B. New
plants sprout from the extensive rhizome system. (Louise Wootton, Georgian Court University, New Jersey.)
C. The stiff leaves slightly arch. (Louise Wootton, Georgian Court University, New Jersey.) D. Male flowers are
distinguished by long strands of pollen. (James Burkitt, Georgian Court University, New Jersey.) E. Female
flower seed heads have brown scales. (Louise Wootton, Georgian Court University, New Jersey.)

wind. Although no confirming evidence exists, seeds and plant pieces are probably saltwater
tolerant and carried by currents and storm surges to new locations.
Asiatic sand sedge’s major means of reproduction is vegetative, expanding its cover by its
extensive rhizome system.
Impacts. Large stands of Asiatic sand sedge form low dense mats, which degrade the eco-
systems of the sand dunes by altering habitat for beach dwellers, displacing native plant spe-
cies, reducing the biodiversity, and perhaps exposing the beach areas to more storm erosion.
Asiatic sand sedge outcompetes or crowds out native dune species, such as American beach
grass and sea oats, the two major sand-binding species. It also displaces, either by competition
or habitat changes, several rare or threatened species, such as seabeach amaranth, seaside knot-
weed, sea sandwort, slender seapurslane, seabeach evening primrose, wormwood, saltmeadow
cordgrass, coastal sand spurge, and sea-coast marsh elder. A study in New Jersey indicated that
Asiatic sand sedge reduced the abundance of native plant species by 50–70 percent. It also
affects the seaside goldenrod, which provides nectar for migrating monarch butterflies. With
a lower biodiversity, the ecosystem is less resilient and more prone to disruption.
By altering the beach habitat, it threatens many rare and endangered species, such as the
northeastern sea beach tiger beetle, which needs open sand. Birds that may be displaced
include Piping Plover, Least Tern, and Black Skimmer. Dense mats of sedge cover the open
sandy areas that the birds require for nesting sites.
By reducing the energy of storm waves, coastal dunes are the first line of protection against
erosion and flooding. Dunes protect the ecosystems and maintain habitat for native plants and
animals, as well as contribute to filtering rainwater. Although evidence is conflicting, Asiatic sand
sedge may change the dune profile. One theory is that because Asiatic sand sedge is shorter than
the native dune grasses, it traps less sand, resulting in lower dunes that provide less protection.
The lower dune is more vulnerable to shifting, blowouts from wind, and storm surges. Other evi-
dence indicates that the sedge stabilizes dune sand so well that dune blowouts, meaning that the
dune is breached and sand is blown inland, occur less frequently. Dune blowouts are responsible
for development of the bare sand habitat necessary for plovers and other high-beach species.
Management. Regardless of the method used to control Asiatic sand sedge, follow-up mon-
itoring and treatment are necessary. Early detection is critical because plants spread rapidly.
 BUFFELGRASS n 435

How Bad Can It Be?

I n a study at Island Beach State Park in New Jersey, the area occupied by Asiatic
sand sedge tripled from 1985 to 2003, and doubled from 2003 to 2008. At Sandy
Hook Unit of Gateway National Recreation Area in New Jersey, the situation was
worse. Asiatic sand sedge increased its presence 780 percent from 1985 to 2003.
From 2003 to 2008, it increased another 300 percent. As of 2008, 20 percent of the
dunes in both parks were dominated by the sedge, not by native plant species.
Source: Louise Wootton’s Pages, 2009.

Because the plant does stabilize the dunes, however, removal should be approached cau-
tiously. When the infestation is eradicated or controlled, revegetating the primary dunes with
American beach grass or sea oats will stabilize them and prevent recolonization by Asiatic sand
sedge. Stable dunes further from the beach can be revegetated with a number of additional
plants, including seaside goldenrod, beach panic grass, dune panic grass, and sea-rocket.
Physical removal is limited to small plants or small infestations, usually fewer than 200
shoots. All rhizomes must be excavated from the sand without breakage, because any buried
remains will sprout a new plant. Plants can be disposed of in an unsuitable habitat away
from the sandy beach or covered with black plastic to bake and die.
Larger stands are generally too labor intensive and costly for hand removal and should
undergo chemical treatment. One or two applications of glyphosate during the growing sea-
son, preferably in summer or fall, are effective, provided they are followed by spot treatment
on new shoots. Because the plant has narrow leaves that can easily be missed by a foliar
spray, a colorant will be helpful to ensure coverage.
No biological controls are known.

Selected References
“Asiatic Sand Sedge.” Invasive Alien Plant Species of Virginia. Department of Conservation and
Recreation. State of Virginia, n.d. http://www.dcr.virginia.gov/natural_heritage/documents/
fscako.pdf.
“Carex kobomugi.” Invasive Plant Atlas of New England (IPANE), 2004. http://nbii-nin
.ciesin.columbia.edu/ipane/icat/browse.do?specieId=121.
“Carex kobomugi.” Louise Wootton’s Pages, Georgian Court University, 2009. http://gcuonline
.georgian.edu/wootton/Carexkobomugi.htm.
“Carex macrocephala.” Louise Wootton’s Pages, Georgian Court University, 2010. http://gcuonline
.georgian.edu/wootton/carexmacrocephalanj.htm.
Lea, Chris, and Greg McLaughlin. “Asiatic Sand Sedge.” Weeds Gone Wild: Alien Plant Invaders of
Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2005. http://www.nps
.gov/plants/alien/fact/pdf/cako1.pdf.

n Buffelgrass
Also known as: African foxtail grass, anjan grass, pasto buffel, zacate buffel
Scientific name: Pennisetum ciliare
Synonyms: Cenchrus ciliaris, C. glaucus, Pennisetum conchroides, P. incomptum
Family: Grass (Poaceae)
436 n GRAMINOIDS

Native Range. Africa, Mada-

gascar, Arabia, and South-
west Asia east to western India.
Distribution in the United
States. Southern states from
California to Florida. Also in
New York and Hawai’i. Most
invasive in the deserts of
California, Arizona, New
Mexico, and Texas, and on the
Hawaiian Islands.
Description. Buffelgrass is a
perennial shrubby bunch grass,
usually 1.5–3.5 ft. (0.5–1 m)
tall and 3–4 ft. (1–1.2 m) in
diameter. It can, however, be
as small as 4 in. (10 cm) or as
large as 5 ft. (1.5 m) tall. The
erect stems are dense, all grow-
ing from the base of the plant
but branching freely at the
nodes, giving it a bushy appear-
ance. Plants form thick mats or
clumps. The previous season’s
growth turns gray and remains
in the clump, adding to the
bushy nature. Golden brown
during dry periods, it quickly
becomes green, growing leaves
and flower spikes after even a
light rainfall. The bluish-green
Deliberately introduced for livestock feed and soil stabilization, buffelgrass leaf blades are flat, 3–11 in.
has become invasive in desert areas in the southwestern United States. (7.5–30 cm) long, and narrow,
(Native range approximated from USDA GRIN and selected references. less than 0.4 in. (1 cm) wide.
Introduced range adapted from USDA PLANTS Database, Invasive Plant Plants feel rough because of
Atlas of the United States, and selected references.) small soft to stiff hairs on the
leaf blades. Ligules, where the
leaf blade separates from the leaf sheath on the stem, are noticeably hairy.
The root system is a long, dense network of stolons, usually extending 6–8 ft. (2–2.5 m)
deep, but sometimes reaching 10 ft. (3 m).
The inflorescences are cylindrical spikes resembling narrow bottle brushes, 0.75–5.5 in.
(2–14 cm) long and 0.4–1 in. (1–2.6 cm) wide. Each spike has 30–50 long bristles, usually
brown or purplish, but sometimes straw-colored. The spikelets, which have no awns, are
either solitary or in clusters. The rachis is rough after the seeds drop. Fruit are bristly burs,
and although the bristles are not hard or stiff, they catch on fur or clothing. Seeds cover the
flower spike so densely that individual seeds are difficult to distinguish.
Related or Similar Species. Buffelgrass is distinguished from similar looking native
grasses by details in morphology. Arizona cottontop is sparsely branched and less shrubby.
 BUFFELGRASS n 437

A. Buffelgrass clumps look bushy because all the stems emerge from the base of the plant. (Joseph M. DiTomaso,
University of California–Davis, Bugwood.org.) B. Ligules are hairy. (Tom Curry, Phoenix Weedwackers.) C. The
root system is dense with stolons. (Tom Curry, Phoenix Weedwackers.) D. The inflorescence is a bristly spike.
(Tom Curry, Phoenix Weedwackers.)

Bush muhly is a densely branched shrub with more delicate stems. Seeds of plains bristle-
grass do not densely cover the flower spike, and individual seeds can easily be distinguished.
Because buffelgrass was introduced for livestock forage, many cultivars with different prop-
erties have been developed.
Introduction History. Buffelgrass was deliberately introduced to Arizona for livestock feed.
Trial plantings were done in Tucson by the Soil Conservation Service from 1938 to 1952. It
was also used for erosion control and soil stabilization. Populations did not rapidly expand,
however, until about 1980. As an example, buffelgrass was rare in Organ Pipe Cactus
National Monument before 1984, but by 1994, plants occupied 20–25 sq. mi. (50–65 km2)
and its range was increasing. It is still promoted in some regions as a range or hay crop.
Habitat. Buffelgrass grows primarily in warm climates with frost-free winters and annual
rainfall of 6–24 in. (150–610 mm), most of which occurs in summer. It is very drought tol-
erant and grows best in dry climates with only 7–10 in. (180–250 mm) of rain. Plants are
most often found on disturbed land, such as roadsides, fields, vacant land, rangeland, or
grassland, but they also invade healthy desert communities and riparian habitats.
Buffelgrass also grows on rocky hillsides, usually east-facing and south-facing slopes. It is
abundant on talus or scree slopes and invades less steep sites from those locations.
Buffelgrass is tolerant of harsh environmental conditions, such as strong wind, aridity, ero-
sion, and poor soils. It grows on all types of soil but is most common on sandy soils, which
it prefers in its native range. Alkaline soils, with a pH of 5.5–8.2, are best. It does poorly on
clay soils with little calcium, and cannot survive extended periods of freezing or flooding.
Providing other conditions are met, it grows from sea level up to 6,550 ft. (2,000 m)
elevation.
Reproduction and Dispersal. Buffelgrass reproduces both sexually and vegetatively.
Plants begin growth in late winter and flower spring through fall. Although plants have both
male and female flowers, buffelgrass can produce seed either with or without fertilization.
Non-fertilized seeds are identical to the parent plant. Plants produce abundant seeds, which
are disseminated by wind, water, animal fur, and human clothing and footwear. Occasional
floodwaters also extend its range. Most seeds germinate at the start of the wet season, usually
spring, but can sprout any time it rains.
Plants also sprout from the rhizomes and stolons as the root system expands.
Impacts. Some of the same characteristics that make buffelgrass desirable for erosion con-
trol or livestock fodder contribute to its invasive nature. Seeds germinate rapidly, and plants
438 n GRAMINOIDS

do well on poor soils. Buffelgrass is resistant to fire, drought, heavy grazing, and trampling,
making it a good arid land forage. It has few pests or predators. The root system stores
carbohydrate reserves, which allow it to regenerate after a fire.
Buffelgrass outcompetes native species, and alters the landscape. The grass grows into
dense clumps that crowd out and kill many native plants. They compete for scarce water,
and the dense system of roots and branches shade the ground and prevent germination of
other plants. It also may have alleleopathic compounds that prevent seeds of other plants
from germinating. In Hawai’i, where it was introduced for erosion control, it is replacing
the native pili grass, also known as tanglehead. In southern Texas, it threatens federally
endangered species, such as Zapata bladderpod and Rio Grande ragweed, also known as
South Texas ambrosia. In the Sonoran Desert of southern Arizona and adjacent Mexico, it
outcompetes and excludes creosote bush, saltbush, bursage, and herbaceous grasses and
forbs, replacing the complex ecosystem with a monoculture grassland, and eventually leav-
ing it barren.
Dense stands of buffelgrass increase the fuel load, resulting in fires that burn hotter. It
burns readily even when green and resprouts after burning. Because fire is not an ecological
part of Sonoran Desert ecosystems, fires kill most native plants.
Evidence exists that after about 10 years, when it has depleted the soil of all nutrients,
buffelgrass dies, leaving behind a nutrient-poor wasteland.
In spite of its use for forage, buffelgrass may be detrimental to livestock. It may cause oxa-
late poisoning in sheep if the animals graze buffelgrass too heavily. Horses may develop a
condition called “bighead,” or hyperparathyroidism, due to calcium deficiency if they eat
too much buffelgrass.
Management. The best control is an integrated approach and early elimination of small
infestations. Because roadsides are corridors for linear spread and expansion into adjacent
fields, removal in those locations should be a priority. Removal is also made more difficult
if adjacent land is allowed to go unchecked, such as along the border between Mexico and
Organ Pipe Cactus National Monument. Any site should be monitored for a minimum of
3–5 years.
Although it is possible to remove extensive infestations, physical removal is best done
only for small stands because of the labor involved. All pieces of the root must be removed
or they will resprout. In Organ Pipe Cactus National Monument, buffelgrass was almost
eradicated from a 25 sq. mi. (65 km2) area when volunteer labor removed over 150 tons
of plant material. Follow-up the second year, which required pulling up new seedlings,
was much less intensive. Repeated tilling can be effective in destroying the plant, but cannot

Sonoran Desert Weedwackers

S everal volunteer groups in southern Arizona meet monthly for buffelgrass pulls.
Since 2000, when the Sonoran Desert Weedwackers was formed, approximately
120 tons of buffelgrass and nonnative fountain grasses have been dug out and
removed from Tucson Mountain Park. Much of the area now supports native
grasses and wildflowers. Weedwackers are also making progress in other areas,
such as Phoenix Mountain Preserves, Waterman’s-Ironwood Forest National
Monument, and Saguaro National Park.
 CHEATGRASS n 439

be done in a natural area. Cutting and mowing is not effective because even when plants are
mowed low to the ground, buffelgrass can produce seed. Mowing may also stimulate
growth, but can be used to decrease above-ground biomass prior to herbicide treatment.
Continued grazing may limit root depth and weaken plants. Burning is not an option
because fire damages native vegetation and buffelgrass will resprout from rhizomes.
Chemical applications are effective, particularly if the standing biomass is reduced first. A
combination of glyphosate and ammonium sulfate, if applied several times, will kill the root
mass. Tebuthiuron and hexazinone reduce growth in some plants. Seedlings can be con-
trolled by a variety of herbicides, including dicamba, 2,4-D, 3,6-dichlorpicolinic acid, triclo-
pyr, tebuthiuron, or hexazinone. Picloram is not effective.
Even though no biological controls, such as insects or pathogens, are yet known, it remains a
controversial concept because buffelgrass is still used as forage in many areas. A leaf blight, also
known as rice blast or buffelgrass blight, caused by a fungus (Pyricularia grisea) may cause sig-
nificant damage, but some cultivars are resistant. Evidence suggests that the fungal blight may
also harm commercial agricultural crops. Restoration of the native vegetation may help to reduce
infestations. Because buffelgrass does not compete well with dense vegetation, in well-managed
pastures, or in shade, planting native shrubs and trees may keep it under control.

Selected References
“Buffelgrass (Pennisetum ciliare).” Citizen Scientists Combat Invasive Species. Invaders of the Sonoran
Desert Region, Arizona-Sonoran Desert Museum. Tucson, AZ, n.d. http://www.desertmuseum.org/
invaders/invaders_buffelgrass.htm.
IUCN SSC Invasive Species Specialist Group (ISSG). “Cenchrus ciliaris (Grass).” ISSG Global Invasive
Species Database. 2006. http://www.issg.org/database/species/ecology.asp?si=846&fr=1&sts
=&lang=EN.
“Invasives, Sonoran Desert Weedwackers.” Arizona Native Plant Society, n.d. http://aznps.com/invasives/
weedwackers.html.
Tu, Mandy. “Element Stewardship Abstract, Pennisetum ciliare.” Global Invasive Species Team, Nature
Conservancy, 2002; modified 2009. http://wiki.bugwood.org/Pennisetum_ciliare.

n Cheatgrass
Also known as: Downy brome, downy chess, early chess, drooping brome, wild oats,
military grass, thatch bromegrass, broncograss, soft chess
Scientific name: Bromus tectorum
Synonyms: Anisantha tectorum
Family: Grass (Poaceae)
Native Range. Southern Europe, northern Africa, Arabia, and Southwest Asia, as far
north as Kazakhstan and southern Russia, and east to Pakistan and western China.
Distribution in the United States. Every state, including Alaska and Hawai’i. It is most
prominent in the grasslands and sagebrush range in the West.
Description. Cheatgrass is a winter annual grass with highly variable characteristics.
Three keys to identification are the drooping panicles, long awns, and color that changes
from green to purple to tan in early summer as plants mature and seeds ripen. Seedlings
are bright green with hairy leaf blades and sheaths. The narrow leaves, slightly twisted
initially, have a prominent midrib and soft hairs. Erect culms on mature plants, 2–35 in.
(5–90 cm) tall, grow in large tufts. The flat leaf blades, 1.5–6 in. (4–16 cm) long, are narrow,
440 n GRAMINOIDS

light green, and covered with

soft, white hairs. Ligules are gla-
brous. The lower sheaths are
also noticeably hairy, while the
upper sheaths may be glabrous.
The root system is finely di-
vided and fibrous, normally
reaching 12 in. (30 cm) deep,
with few main roots.
Flowering stems are erect,
either smooth or slightly hairy.
Inflorescences, densely grouped
in open panicles within the
leaves, are 2–8 in. (5–20 cm)
long, with several slender
branches that droop or sag to
one side from the top of the
inflorescences. The panicle
branches are pubescent, each
holding 3–8 spikelets. Spike-
lets, with 4–8 florets each, are
0.8–2 in. (2–5 cm) long, includ-
ing the awns, and also droop
from the stalk. Awns are slender
and straight, often longer than 1
in. (2.5 cm). Seeds are narrow,
about 0.5 in. (1.3 cm) long, light
and fluffy. Plants die after
flowering, usually by July.
Related or Similar Species.
Several introduced, invasive
Cheatgrass often excludes all other plants, forming a monoculture in bromes, with a variety of comm-
rangeland in western states. (Native range adapted from USDA GRIN and on names, resemble cheatgrass.
selected references. Introduced range adapted from USDA PLANTS Distinctions are in the details.
Database, Invasive Plant Atlas of the United States, and selected references.) Three annuals, Japanese brome,
native to southeastern Europe
and Asia, soft brome, native to southern Europe and northern Africa, and bald brome, native
to the Baltic region of Europe, are widespread in disturbed areas such as roadsides. Smooth
brome is a rhizomatous perennial native to Eurasia that has been widely planted for forage from
Texas to Alaska. No native bromes are similar to cheatgrass.
Introduction History. Accidentally introduced to North America sometime before 1860
and probably many times after that, cheatgrass was first identified in 1861 in New York and
Pennsylvania. It probably arrived as a seed contaminant in grain or in soil used as ballast for
ships. The plant became prominent along railroad right-of-ways, its dispersal probably facili-
tated by straw packing material in freight transported by railroads. It had become a serious
problem in crops by the 1920s and reached its present range throughout the United States
by 1930. The name “cheatgrass” was coined by farmers on the Great Plains, who believed they
had been cheated with impure seed when this grass sprouted in their wheat fields.
 CHEATGRASS n 441

A. In many places, cheatgrass has replaced the native shrub and grass community. (Chris Evans, River to River
CWMA, Bugwood.org.) B. The open panicles droop to one side of the stem. (Steve Dewey, Utah State University,
Bugwood.org.) C. The fine roots extend deep into the soil. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology
University of Connecticut, Bugwood.org.) D. Leaf blades and lower leaf sheaths are softly hairy. (Tom Heutte,
USDA Forest Service, Bugwood.org.)

Habitat. Cheatgrass grows under a wide variety of conditions. In the western states, it is
most commonly found in sagebrush, bunchgrass, and desert scrub communities, but it also
spreads into pinyon, juniper, and pine forests at higher elevations. Cheatgrass is a successful
invader of rangeland, roadsides, and abandoned croplands or fields. It is also common in
recently burned areas and a pest in winter crops. Although more prevalent on disturbed
ground, whether as major as overgrazed or cultivated or as minimal as rodent holes, cheat-
grass also invades undisturbed ground when seeds settle into and sprout in natural soil
cracks. Although capable of growing on almost any soil type, cheatgrass prefers coarse soil
and does not do well on heavy or saline soils or dry sites.
Although it occurs in many climates, with precipitation ranging from 6 to 27 in. (150–
685 mm) annually, it is most invasive in semiarid regions with 12–22 in. (300–560 mm)
of precipitation. Cheatgrass does not tolerate shade and will not survive in mature, shady
forests. It can be found growing to an elevation of 13,125 ft. (4,000 m).
Reproduction and Dispersal. Cheatgrass reproduces only by seed, but may resprout from
rhizomes if mowed or burned too soon. It is self-fertile and needs no cross-pollination.
Cheatgrass produces an extensive number of seeds, as many as 300 per plant, and maintains
a large soil seed bank. Seed production, however, depends on plant density and growing
conditions and is less in dense stands. The smallest seed producer may be only 1 in.
(2.5 cm) tall. Seeds are viable in the soil for only 2–5 years, but for as long as 11 years in a
dried state, such as in hay. At maturity, the spikelets break apart to disperse the seeds. The
majority of seeds fall near the parent plant. Wind, water, rodents, and ants disperse seeds
a short distance away. Seeds are carried longer distances by human activity, such as on
machinery, vehicles, or clothing, and in contaminated hay, grain, or straw. Dispersal is
common along transportation corridors such as roads or railroads. Both domestic and wild
animals and birds also transport seeds in fur, feathers, and feces.
Seeds do not need to contact bare soil, and litter promotes the germination and establish-
ment of seedlings. The germination rate is high, although dry conditions can induce dor-
mancy in seeds, as evidenced by seeds in dried hay. Most seeds germinate in the fall,
within 1 in. (2.5 cm) of the surface. Growth begins at 35–38ºF (2.0–3.5ºC) and ends when
temperatures are higher than 59ºF (15ºC). Plants overwinter as a small seedling, 0.8–2 in.
442 n GRAMINOIDS

(2–4 cm) tall, and grow a root system that is ready to outcompete native spring growers. The
shallow root system draws surface water early in spring, preventing other seeds and plants
from germinating or growing. The flower-to-seed cycle is quick. Inflorescences develop in
mid-spring, flowers follow within a week, and seeds mature in mid-to-late June. By complet-
ing its life cycle early, cheatgrass avoids dry summer conditions. By depleting soil moisture
early in spring, it deprives native plants of needed water.
Impacts. As an annual, cheatgrass produces leaves and inflorescences in a single season
and then dies in early summer. This accumulation of thatch is a major fire hazard, which
changes the fire regime from approximately once every 60–100 years to once every 3–
5 years. Fire is especially detrimental to desert communities that are not fire adapted.
Cheatgrass dominates over 100 million ac. (40.5 million ha) in the western states, perma-
nently changing the ecosystems. In many places, it has totally displaced native shrubs, such
as big sagebrush and antelope bitterbrush, and perennial grasses, such as bluebunch grass,
western wheatgrass, Sandburg bluegrass, needle-and-thread grass, and Thurber’s needle-
grass. Replacement generally takes several years, beginning with a disturbance, such as over-
grazing, that allows initial invasion. Frequent fires replace the shrubs and perennial grasses
with annual grasses, causing the soils to deteriorate. Eventually, the cheatgrass outcompetes
the remaining annuals and becomes a monoculture.
The loss in plant biodiversity affects native wildlife in several ways. Bird numbers and
species diminish because of less suitable habitat. Large animals, such as elk and mule deer,
no longer have native forage plants. The numbers and diversity of predators, such as coyo-
tes, gopher snakes, and golden eagles, decline as their prey, small mammals such as ground
squirrels and packrats, becomes scarce. Once the ecosystem is impoverished, cheatgrass-
dominated range is susceptible to other nonnative invaders, such as medusahead (see
Graminoids, Medusahead).
Cheatgrass is also a weed in winter wheat, alfalfa, and other crops. The estimated cost in
lost yields and control for the western United States and Canada is $350–370 million each
year. Depending on the density of cheatgrass infestation, wheat yields can be reduced by
40–92 percent.
The long, bristly awns injure livestock and other animals, sticking in eyes, mouths, and
throat if eaten when plants are dry in late spring or summer. Such injuries may prevent
the animal from eating, resulting in weight loss.
Management. For effective eradication, an integrated approach, which may involve fire,
herbicide, and reseeding, is necessary for at least 2–3 years to create a cumulative stress on
plants. Reduction of seed output is most important. Proper land management and a seed
source of native plants are essential to prevent re-invasion. Once roots of native perennials
reach more than 20 in. (0.5 m) deep, they can compete with cheatgrass because it uses water
at shallow depths. Total eradication, however, may not be the goal in areas where cheatgrass
is important early-season forage for livestock.
Physical control methods should minimize soil disturbance and seed dispersal and main-
tain the health of native plants. Hand-pulling is only appropriate for small areas or seedlings,
and all root pieces must be removed. It may take several years before the seed bank is
depleted. Live plants can be tilled into the soil in spring and fall before the plants turn pur-
ple, the process repeated whenever new plants appear. Seeds deposited deeper than 3 in.
(8 cm) do not sprout. Depending on the remnant native plant community and timing, live-
stock grazing can be effective. Mowing soon after flowering, every three weeks in spring and
early summer, prevents most seed formation. Fire will kill seeds if plants are burned after
they dry but before seeds are dropped. If burned or mowed too early, roots may resprout.
 COGONGRASS n 443

Benefits of Cheatgrass?

C heatgrass is a detrimental invasive weed, but it is also an early spring forage

crop. It is important forage for both livestock and wildlife when it is green in
spring, especially where native bunchgrasses have been eliminated. However, it is
not dependable because the season and amount of edible green foliage depends
on rainfall.
The Chukar Partridge, a pheasant introduced from Pakistan as a game bird, eats
cheatgrass, and is currently expanding its range, following the invasion of cheatgrass.

Chemical control is difficult because native plants or crops should be preserved. The
choice of herbicide depends on the remaining native vegetation or type of crop. Several
choices also exist for non-crop infestations or rangeland. Atrazine and imazapic are preemer-
gents used in late summer or early fall to prevent seeds from sprouting. Paraquat or glypho-
sate can be sprayed on emerging seedlings in early spring or while seeds are developing.
Biological control is limited, although rabbits, mice, and migratory grasshoppers
(Melanoplus sanquinipes) eat the seeds. Over 20 diseases have been reported, but research has
been limited. Cheatgrass is susceptible to a heat smut (Ustilagao bulleta), which, if severe, can
decrease seed yield. A pink snow mold (Fusarium nivale) is being investigated, but no conclu-
sions have been reached. Experiments are also being conducted on phytotoxins produced by
two rhizobacteria strains (Pseudomonas fluorescens and P. syringae), which inhibit germination.

Selected References
“Bromus tectorum, Cheatgrass.” NatureServe Explorer. An Online Encyclopedia of Life, 2009. http://
www.natureserve.org/explorer/servlet/NatureServe?searchName-Bromus%20tectorum.
Carpenter, Alan T., and Thomas A. Murray. “Element Stewardship Abstract, Bromus tectorum.” Global
Invasive Species Team, Nature Conservancy, 1999. http://wiki.bugwood.org/Bromus_tectorum.
“Cheatgrass.” Non-Native Plant Species of Alaska. Alaska Natural Heritage Program, Environment and
Natural Resources Institute, University of Alaska, Anchorage, 2005. http://akweeds.uaa.alaska.edu/
pdfs/species_bios_pdfs/Species_bios_BRTE.pdf.
“Downy Brome (Cheatgrass).” Colorado State Parks Best Management Practices Weed Profile, 2003;
revised 2005. http://parks.state.co.us/SiteCollectionImages/parks/Programs/ParksResourceSteward
ship/Downey%20Brome.pdf.
Pokorny, Monica. “Cheatgrass (Bromus tectorum).” 2007. http://www.ipm.montana.edu/cropweeds/
Extension/.

n Cogongrass
Also known as: Speargrass, cogon grass, satintail, blady grass, Japanese blood grass
Scientific name: Imperata cylindrica
Synonyms: Imperata cylindrica var. major, Imperata arundinaceae, Lagurus cylindricus
Family: Grass (Poaceae)
Native Range. The native range is not clearly known. Eastern Africa and Asia, north to
Kazakhstan and east through India, China, Korea, and Southeast Asia. Also Japan, the
Philippines, Indonesia, and New Guinea. Possibly native to southen Africa, northern
Africa, and Mediterranean Europe.
444 n GRAMINOIDS

Distribution in the United

States. Primarily in the south-
eastern United States, from Texas
east to Virginia. Also in Oregon.
Description. Cogongrass is a
tufted perennial grass that
grows in either loose or com-
pact bunches. Because it is so
widespread geographically, it is
variable in form. The bunch con-
sists of several lance-shaped
leaves rising from ground level,
directly from the rhizomes.
Leaves are 1–5 ft. (0.3–1.5 m) tall
and 0.5–1 in. (1.3–2.5 cm) wide,
with a prominent white midrib
that is slightly off center. The
sharply pointed leaf blades are
light green, turning orange-
brown as they age. Leaf margins
are finely toothed and embedded
with silica crystals. The upper
surface of the leaf blade is hairy
near the base, while the undersur-
face is glabrous. Where subjected
to freezing temperatures in win-
ter, leaves turn light brown.
Cogongrass has an extensive
system of hard, creeping rhi-
zomes that are sharply pointed
and scaly. Most occur in the
Cogongrass grows in many environments but does not tolerate cold top 6 in. (15 cm) of soil, but
weather. (Native range approximated from USDA GRIN and selected they can penetrate as deep as
references. Introduced range adapted from USDA PLANTS Database, 4 ft. (1.2 m). Where the plant
Invasive Plant Atlas of the United States, and selected references.) is firmly established, the rhi-
zome mass can weigh 3–16 tons
per acre (7.5–40 tons per ha).
Flowering season is both spring and fall. Silvery, plume-like flowers grow on a many-
branched cylindrical panicle, 3–11 in. (7.5–28 cm) long and 1.5 in. (4 cm) wide. The small spike-
lets, 0.1–0.2 in. (4–5 mm), are surrounded by several silky white hairs, which are 0.4–0.6 in.
(10–15 mm) long, making the spikelet seem larger. Spikelets are crowded on the stalks, enhanc-
ing the plume-like appearance. Flowers have three bracts within the glumes, and two stamens
per flower. Seeds, which develop in spring, are very small, with a plume of long white hairs.
Related or Similar Species. Japanese blood grass (I. cylindrica ‘Rubra’), a cultivar called
Red Baron, is distinguished by its brightly colored, blood-red leaf edges. This attractive vari-
ety, which is not on the U.S. federal noxious weed list, is frequently sold in nurseries or on the
Internet for ornamental plantings. Although it is described as noninvasive, no evidence exists
to support that claim. It is not known if this variety is a potential threat to natural areas, but
 COGONGRASS n 445

A. Cogongrass grows in dense stands. (Florida Division of Plant Industry Archive, Florida Department of Agriculture
and Consumer Services, Bugwood.org.) B. Rhizomes are sharply pointed. (Chris Evans, River to River CWMA,
Bugwood.org.) C. The upper surfaces of leaf blades are hairy. (Chris Evans, River to River CWMA, Bugwood.org.)
D. Leaf blades have a prominent white midrib. (Chris Evans, River to River CWMA, Bugwood.org.) E. Rhizome
buds produce many new sprouts. (Nancy Loewenstein, Auburn University, Bugwood.org.) F. Silvery plumed
flowers, with silky hairs, are showy. (John D. Byrd, Mississippi State University, Bugwood.org.)

it becomes invasive in horticultural settings. Seedlings exhibit aggressive characteristics similar

to those of the green form. The plant is very cold tolerant, and the red color is more intense in
colder climates. In southern regions, the plants revert to green. The variety tends to be more
invasive in regions where leaves lose their red color. Red Baron rarely blooms, and no records
exist of flowering or setting seed in cold climates, but the plant has the potential to produce
3,000 seeds per season. If Red Baron were to hybridize with the main variety of cogongrass
(I. cylindrica var. major) that grows in the southeastern United States, the hybrid would be both
cold tolerant and invasive.
Brazilian satintail, native to Central and South America, may sometimes be distinguished from
cogongrass by having two bracts within the glumes and one stamen per flower. These distinc-
tions, however, are variable. The two plants hybridize and may be the same species. California
satintail resembles Brazilian satintail, but is larger, with culms generally reaching 3.3–4.9 ft.
(1–1.5 m) tall. California satintail is genetically distinct from the nonnative Imperata species.
Introduction History. Introduction took place several times. Cogongrass was accidentally
introduced when it was brought to Mobile, Alabama, in 1912 as packing material in orange
crates. In 1921, the U.S. Department of Agriculture intentionally introduced the plant from the
Philippines for forage. In the 1930s and 1940s, it was brought to Florida for potential forage
and for soil stabilization. The finely serrated edges and silica crystals made it undesirable for for-
age. Cogongrass spread by illegal plantings and contamination in soil during road construction. It
was also sold as an ornamental due to its hardiness and attractive leaves. It has established in the
last 50 years in the southeastern United States, especially in Alabama, Mississippi, and Florida.
Habitat. Cogongrass can be found in a variety of ecosystems, from open fields to the edge of
standing water, including coastland, riparian, swamplands, savannas, forests, and desert dunes.
It is quick to overtake disturbed sites, such as ditch banks, pastures, roadsides, mine sites, and
forestry plantations. It forms a monoculture on hundreds of acres of old phosphate mines, and
is also a weed on open cultivated ground, wastelands, and in parks and golf courses. Although
446 n GRAMINOIDS

it prefers acidity, pH of 4.7, cogongrass grows on any soil type that has enough moisture, includ-
ing fine sand to heavy clay and soils with low fertility. Cogongrass is tolerant of many adverse
environmental conditions, including shade, high salinity, and drought. It normally does not tol-
erate cold weather, but some stands have survived temperatures of 7ºF (−14ºC).
Reproduction and Dispersal. Plants may produce several thousand small, wind-
disseminated seeds, which may also be carried over short distances by animals. Plants are not
self-fertile and must be cross pollinated to produce viable seed. Less than half of the spikelets,
however, will develop seeds. Seeds have no dormancy requirement, and 90 percent of the seeds
produced are viable. Seedling survival is poor, with fewer than 20 percent living to be one year
old. Cogongrass spreads primarily by rhizomes. A fragment as small as 0.08 in. (2 mm) will
grow. Both seeds and fragments of rhizomes are inadvertently carried long distances on equip-
ment, including roadside mowers, and in fill dirt during road construction. Rhizomes, which
can grow 5–10 ft. (1.5–3 m) per year, can be dormant for a long time before sprouting. They
have numerous buds ready to sprout, and are resistant to fire and deep burial.
Impacts. Cogongrass forms dense mats of thatch and leaves that native plants cannot pen-
etrate. The sheer mass of rhizomes enables cogongrass to dominate, but the plant may also
exude alleleopathic substances. Eventually, cogongrass excludes all other plants, halting suc-
cession. Monospecific grass thickets displace native plants used by animals, including mam-
mals, birds, and insects, for forage, shelter, and host plants. Ground-nesting species are
especially at a disadvantage because of the dense cover of cogongrass. The grasses provide poor
habitat for wildlife and poor forage for grazers, either wild or livestock. New growth, however,
is not coarse and has some limited use as forage. The tall grasses shade out native species, espe-
cially short herbs. It invades and displaces longleaf pine flatwood communities. In Florida,
cogongrass destroys the habitats of the rare gopher tortoise and the rare indigo snake.
The biomass of thatch alters fire regimes, causing fires to be both more frequent and
intense. Fires may injure or destroy native vegetation outright, or reduce their ability to
flower and set seed. Fires especially impact small mammals, herpetofauna, and invertebrates
that cannot move fast enough to escape.
Management. The density and mass of rhizomes makes cogongrass difficult to eradicate.
When used independently, methods such as burning, cultivation, cover crops, and herbi-
cides have only limited success. An integrated management system, which includes
mechanical, cultural, and chemical means, is necessary to eradicate or control cogongrass.
The goal is to eliminate all rhizomes.
Physical control of cogongrass can involve burning or mowing, but both actions stimulate
growth from rhizomes. Plowing or disking, at least 6 in. (15 cm) deep, several times during a
dry season will cause rhizomes to become desiccated and lose their carbohydrate reserves.
This method may keep cultivated fields free of cogongrass. Mowing, burning, or fertilization,
however, also stimulates seedhead formation. Mowing the grass in late spring will remove
thatch and old leaves, but the area should be deeply plowed or disked 6–8 weeks later, when
grass has resprouted to 6–12 in. (15–30 cm) high. Regrowth in delicate ecosystems should
be followed with application of a systemic herbicide, preferably in early fall before the first frost.
While hundreds of possibilities have been tested, only a few herbicides provide effective
chemical control. Soil sterilants are effective, but kill everything for 6–12 months, exposing
bare ground to potential erosion. If the area is to be planted to prevent erosion, bahia grass or
bermuda grass provides good control in nonnatural areas. Choice of plants in natural areas is
more difficult and depends on the ecosystem. Glyphosate or imazapyr applied to growing
plants will reduce infestations, but several applications may be necessary. Because it leaves no
soil residual, glyphosate should be used if the area will be replanted soon. If revegetation of
 COMMON REED n 447

Cogongrass in the World

O ne of the world’s 10 worst weeds, cogongrass is invasive in tropical and sub-

tropical regions throughout the world that receive 30–200 in. (750–5,000 mm)
annual rainfall. It grows best with daytime temperatures of 84ºF (29ºC) and nights
of 73ºF (23ºC), temperatures that are typical for the wet tropics. It is a pest in 73
countries and 35 crops, including teak, cocoa, kola, coffee, cashew, oil palm, coco-
nut, and rubber. Annual yields may be reduced by 80–100 percent, especially signifi-
cant where farmers are dependent on subsistence agriculture. Found from sea level
to 6,550 ft. (2,000 m) elevation, cogongrass infests approximately 500 million ac.
(20.25 million ha) of plantation and agricultural land worldwide. It thrives in dis-
turbed sites, such as burned, overgrazed, or intensely cultivated land, and invades
abandoned shifting agricultural land. Benefits of cogongrass are limited. It is used
locally for thatch, forage, erosion control, paper making, and bedding for livestock.

the area is not planned, imazapyr is a better choice. Fluazifop, which is selective to grasses, is
moderately effective. Sites should be monitored and spot treated as necessary.
The search for natural pests for biological control of cogongrass has met with limited suc-
cess. The gall midge (Orseolia javanica) is host-specific but still has limited effectiveness. Its
larvae enters the plant and penetrates the shoot apical meristem, forming a gall where it de-
velops and pupates. Few eggs survive, however, due to predation by the midge’s many
natural enemies. A nematode (Heterodera sinensis), a mite (Aceria imperata), and two uniden-
tified stem borers may have potential. A host-specific fungus (Colletotrichum caudatum),
which was tested in Malaysia, did some damage, but failed to kill whole plants. Some patho-
gens are possibilities, but none provide effective control.

Selected References
“Cogongrass, Imperata cylindrica.” Excerpted from Invasive Species Management Plans for Florida,
Circular 1529, by G. MacDonald et al. IFAS Extension, University of Florida, 2009. http://
plants.ifas.ufl.edu/node/199.
Johnson, Eric R. R. L., and Donn G. Shilling. “Cogon Grass.” Weeds Gone Wild: Alien Plant Invaders of
Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps
.gov/plants/alien/fact/imcy1.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Imperata cylindrica (Grass).” ISSG Global Invasive Species Database. 2009. http://
www.issg.org/database/species/ecology.asp?fr=1&si=16.
Van Loan, A. N., J. R. Meeker, and M. C. Minno. “Cogon Grass.” In Biological Control of Invasive Plants in
the Eastern United States, by R. Van Driesche et al. U.S. Department of Agriculture, Forest Service
Publication FHTET-2002-04, Morgantown, WV, 2002. http://www.invasive.org/eastern/biocontrol/
28CogonGrass.html.

n Common Reed
Also known as: Giant reed, phragmites, giant reedgrass, Roseau, Roseau cane, yellow cane, cane
Scientific name: Phragmites australis ssp. australis
Synonyms: Phragmites communis, P. communis var. berlandieri, P. phragmites
Family: Grass (Poaceae)
448 n GRAMINOIDS

Native Range. Because

common reed is widespread
throughout the world, its native
origin is not clear. It is believed
to have originated in Eurasia.
The subspecies americanus is
native to contiguous United
States and Canada. Believed to
be native to Puerto Rico.
Distribution in the United
States. Every state except Alaska.
It is a problem primarily from the
Great Lakes states to the East
Coast and south to Georgia, and
in Colorado.
Description. Common reed
is a warm-season, sod-forming,
perennial grass that forms large
clones. The erect culms, which
are rigid, smooth, and hollow,
are almost 1 in. (2.5 cm) in
diameter and usually 6–13 ft.
(2–4 m) tall. Lance-shaped
leaves, 10–20 in. (25–50 cm)
long, and 0.4–2 in. (1–5 cm)
wide, do not grow from the
base of the plant but are alter-
nate on the culms. The leafy
stems do not branch. Shoots
and leaves are stiff and sharp
because of high cellulose and
Common reed, genetically distinctive from the native subspecies, was silica content, and leaf margins
inadvertently dispersed along transportation corridors. (Native range can be somewhat rough.
approximated from USDA GRIN and selected references. Introduced Leaves are deciduous in winter.
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the While the roots grow to about
United States, and selected references.) 3.3 ft. (1 m) deep, the depth of
the dense rhizome system
depends on soil conditions. Rhizomes may be anywhere from 4 in. (10 cm) to 6.5 ft. (2 m) deep.
They are thick and scaly and can extend 70 ft. (20 m) away from the parent plant. In soil, they are
long, thick, and unbranched. In water, rhizomes are shorter, more slender, and branching.
During times of low water, plants may grow stolons at the surface up to 43 ft. (13 m) long.
From July through September, purple or golden flowers appear at the tips of the tall
stems. The inflorescence is 1–1.25 ft. (0.3–0.4 m) long, consisting of a bushy panicle with
many feathery branches or spikelets. Each spikelet, which has many silky hairs, has 3–7
florets, each 0.5 in. (1.25 cm) long, with the larger florets at the base of the panicle. Lower
florets are male or sterile, without awns, contrasting with the awned female upper florets.
The panicles have a fluffy look when seeds mature because of the hairs, giving the plume a
gray sheen. Seeds are small, less than 0.1 in. (2.5 mm) long.
 COMMON REED n 449

A. Plants can be more than 10 ft. (3 m) tall. (James H. Miller, USDA Forest Service, Bugwood.org.) B. Leaves are
alternate on the smooth stems. (Steve Dewey, Utah State University, Bugwood.org.) C. Stiff leaves are as wide as 2
in. (5 cm). (Richard Old, XID Services, Inc., Bugwood.org.) D. The inflorescence is a long, bushy panicle.
(Richard Old, XID Services, Inc., Bugwood.org.)

Clones of common reed can live more than 1,000 years, but individual portions live only
about eight years.
Related or Similar Species. The species Phragmites australis is complex. Of the 27 genetic
strains of Phragmites worldwide, 11 are native to the United States. Although all strains are var-
iable, making identification difficult, the introduced invasive plant appears to be a distinctive
genetic type. The native common reed, P. australis ssp. americanus, and the introduced
common reed, P. australis ssp. australis, are different genotypes that can be distinguished by
details. Leaf sheaths on native plants loosely adhere to the stems and drop off as the plant dies.
Leaf sheaths on introduced plants are tight to the stem and are not shed. Ligules on the native
variety are slightly longer than those on the introduced common reed. The glumes are also
longer. No evidence exists for natural hybrids between the native and introduced forms.
The native subspecies of common reed is often found intermixed with other plants, such
as bulrush, cattail, arrowhead, northern redgrass, and sedge (Carex spp.), in undisturbed
freshwater marshes. Introduced plants most frequently form a monoculture in disturbed
or undisturbed brackish water along the Atlantic coast or along roadsides.
A third subspecies (Phragmites australis ssp. berlandieri), growing in the southern states
from Florida to California, is possibly a Gulf coast native. It is not easily distinguished, and
its status remains unresolved. The closest relative of Phragmites australis is giant reed
(Arundo donax), another invasive large grass (see Graminoids, Giant Reed).
Introduction History. The aggressive genotype of common reed is grown commercially in
Europe for thatch, fodder, and cellulous production. It was accidentally introduced to the
United States in the late 1700s or early 1800s, probably at several Atlantic ports in ship bal-
last. Not noticed at first because of its similarity to native Phragmites, it became established
on the Atlantic coast and spread west in the 1900s along roads, railroads, and waterways.
In addition to genetic differences from native Phragmites, the fact that wildlife fail to use
the invasive species and that no herbivores specialize in consuming it is further evidence that
the plant was introduced. Common reed is still sold in nurseries.
Habitat. As its wide geographic distribution suggests, common reed has broad environ-
mental tolerances. It is found in climate zones ranging from desert to continental to sub-
tropical, wherever moisture is available. It grows from sea level to 7,000 ft. (2,100 m)
elevation. Leaves and stems are killed by the first frost in autumn, but stems remain erect
450 n GRAMINOIDS

all winter. It prefers sunny wetlands that may be either fresh or brackish, but high salinity
limits growth. In shady locations, plants will be short and stems less stiff. It has no specific
soil requirements and grows in soils with a variety of nutrient levels, organic matter, acidity,
and texture. It may even grow as a floating mat, not rooted in soil. Although also found in
undisturbed sites, common reed more easily outcompetes native plants in areas undergoing
some type of environmental stress, such as altered hydrology, dredging, restricted tidal
flooding, storm drainage, and road salt or water pollution. It can be found growing in tidal
or nontidal marshes, river edges, lake and pond shores, springs, irrigation ditches and other
waterways, and in disturbed areas, such as roadsides and reclaimed strip mines. It does best
in slow or stagnant water with silty bottoms.
Reproduction and Dispersal. Common reed reproduces both by seed and vegetatively.
Plants are wind-pollinated, self-fertile, and can also produce seeds without fertilization.
Each plant produces thousands of seeds each year. Viability of seeds is affected by site or
environmental factors, and dispersal is by wind and water. Seeds germinate in spring, under
alternating night and day temperatures of 60ºF (15.5ºC) to 85ºF (29.5ºC). They need
exposed moist soils, and seedlings cannot emerge from depths below 2 in. (5 cm). A seed
bank in marshy soils may re-infest sites after eradication. Dispersal to new sites is often
accomplished by seeds, even though mortality rate of seedlings is high, due to flooding,
drought, salt, or freezing.
Rhizome fragments carried along waterways and shorelines grow into new plants, giving
rise to new stands. Rhizome pieces caught in machinery are responsible for spreading the
plant along roadsides. Sprouts from rhizomes are less sensitive to environmental conditions.
Each rhizome lives for 2–3 years.
Impacts. The nonnative aggressive strain of Phragmites has invaded native Phragmites hab-
itat and displaced it on the East Coast, where most stands are now the introduced European
genetic strain. The native genotype is still common in the Southwest and Pacific Northwest
except along roads and waterways and in urban areas. Common reed changes hydrology,
alters wildlife habitat, and increases the fire potential because of the high biomass. While the
native common reed is valuable to waterfowl, such as ducks, herons, and egrets, and provides
cover for deer, the introduced plant crowds out native plants and reduces biodiversity. It
degrades the quality of wetland habitat, especially for migrating waterfowl, and provides little
food or shelter for wildlife. Some rare and threatened bird species in Connecticut that are
present in native, short-grass marshland habitats are absent from Phragmites monocultures.
Limited research indicates that bird diversity rebounds after control or eradication.
Common reed forms dense, impenetrable stands of both living and dead stems, as many
as 20 per sq. ft. (200 per m2). The nonnative variety sprouts earlier in spring than the native
Phragmites, producing a greater biomass sooner. Plants grow quickly, and colonies can cover
hundreds of acres. Stems can grow 1.5 in. (3.8 cm) in one day, and rhizome runners can
grow more than 10 ft. (3 m) in one growing season. The rapid growth rate of rhizomes
and shoots starves competing native plants of nutrients. The dense growth blocks light,
and the root system occupies all the soil space beneath the surface.
Management. Any attempts to control common reed must involve a long-term plan
because colonies are long-lived. Infestation of new water bodies can be prevented if all
equipment on boats and trailers is thoroughly inspected when leaving and entering a water
area. Deliberate planting for erosion control or ornamental purposes also should be avoided.
Physical elimination may include several years of cutting, pulling, or mowing. Prescribed
burns after flowering will reduce dead biomass, both standing and ground litter, and
decrease the hazard of wildfire. Reduction in biomass may also allow other plants to sprout
 COMMON REED n 451

the following spring. Burning in spring and summer before plants flower, however, may
stimulate growth because rhizomes are not affected by fire. Repeated mowing slows the
spread of common reed by depriving it of its photosynthesis ability but will not kill the
plants. The increase in temperature under a cover of plastic, however, may kill rhizomes.
Sediments may be excavated to remove the rhizomes, but any pieces left in the soil will
regenerate. Because common reed will die if water is too salty, manipulation of tidal flow
may increase salinity above its tolerance levels.
Foliar application of glyphosate is the most effective chemical treatment. Applications
work better when done before the rhizome network becomes established. Best applied in
late summer or early fall after flowering, the herbicide can be either sprayed onto foliage
or painted on cut stem bases. Treatments are necessary for several years to kill regrowth from
rhizomes.
No biological control for common reed is available in the United States, but research is
being conducted on several nonnative insects. Larvae of three shoot flies (Lipara spp.) pro-
duce galls, which prevent the formation of inflorescences. A gall midge (Lasioptera hungar-
ica), a legless red mealybug (Chaetococcus phragmitis), a shoot-boring moth (Archanara
geminipuncta), and a fly (Platycephala planifrons) all weaken stem tissue. Four moths
(Rhizedra lutosa, Phragmataecia castaneae, Chilo phragmitella, and Schoenobius gigantella) feed
on roots and rhizomes. Although none of these insects have been deliberately introduced
to North America, some are found in the United States. No evaluations have yet been made
on their effectiveness.

Selected References
Blossey, B., M. Schwarzlander, P. Hafliger, R. Casagrande, and L. Tewksbury. “Common Reed.” In
Biological Control of Invasive Plants in the Eastern United States, by R. Van Driesche et al. U.S.
Department of Agriculture, Forest Service Publication FHTET-2002-04, 2002. http://
www.invasive.org/eastern/biocontrol/9CommonReed.html.
“Common/Giant Reed.” Aquatic Invasive Species. Indiana Department of Natural Resources, 2005.
http://www.in.gov/dnr/files/PHRAGMITES2.pdf.
Gucker, Corey L. “Phragmites australis.” In Fire Effects Information System, U.S. Department of
Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, 2008.
http://www.fs.fed.us/database/feis/.
Saltonstall, Kristin. “Common Reed.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/
phau1.htm.

History of Common Reed

C ommon reed has been part of the native flora in the southwestern United
States for at least 40,000 years, as indicated by remains found in ground sloth
dung. Preserved remains have also been found in coastal salt marsh sediments
dated 3,000–4,000 years ago. Native Americans used common reed for arrow
shafts, ceremonial objects, musical instruments, and cigarettes. Leaves and stems
were used by the Anasazi in southwestern Colorado about 1,000 years ago for con-
struction of mats.
452 n GRAMINOIDS

n Cordgrasses and Their Hybrids

Also known as: Smooth cordgrass, Atlantic cordgrass (Spartina alterniflora); dense-flowered
cordgrass (Spartina densiflora); salt meadow cordgrass, salt marsh hay (Spartina patens);
common cordgrass (Spartina anglica); smooth cordgrass hybrid (Spartina
alterniflora x foliosa)
Synonyms for smooth cordgrass: Spartina alterniflora var. glabra, S. alterniflora var. pilosa
Synonyms for salt meadow cordgrass: Spartina patens var. juncea, S. patens var. monogyna
Synonym for common cordgrass: Spartina maritima x alterniflora
Family: Grass (Poaceae)
Native Range. Smooth cordgrass and salt meadow cordgrass are native to the Atlantic
and Gulf coasts, from Texas to
Newfoundland. Smooth cord-
grass is also native to the east
coat of South America and the
islands of Trinidad and Guade-
loupe in the Lesser Antilles, but
not to Central America. Salt
meadow cordgrass is also native
to many Caribbean Islands in
both the Greater and Lesser
Antilles. Dense-flowered cord-
grass is native to southern
South America, including Brazil,
Uruguay, Argentina, and Chile.
Common cordgrass is a hybrid
beween smooth cordgrass and
small cordgrass (S. maritima),
which is native to England.
Distribution in the United
States. Distribution of alien
cordgrasses in disjunct. They
are found in various bays and
coastal locations from Washing-
ton south to the San Francisco
Bay region in California. Dense-
flowered cordgrass is also found
in isolated parts of Texas.
Description. Smooth Cord-
grass: Smooth cordgrass is a
perennial emergent grass that
usually grows in dense stands.
Although its size varies accord-
Although native to the Atlantic and Gulf coasts of North America, as well ing to conditions, its stiff, erect
as to the Atlantic coast of South America, smooth cordgrass is invasive on stems, 1 in. (2.5 cm) wide at
the Pacific coast. (Adapted from USGS Nonindigenous Aquatic Species the base, are often 1 ft. (0.3 m)
Database and selected references.) tall in spring and may reach a
 CORDGRASSES AND THEIR HYBRIDS n 453

A. Smooth cordgrass and its hybrids colonize mudflats, changing the habitat to meadow. (California Coastal
Conservancy, San Francisco Estuary Invasive Spartina Project.) B. The extensive root system, with rhizomes
spreading just beneath the surface, traps sediment. (California Coastal Conservancy, San Francisco Estuary
Invasive Spartina Project.) C. Stems are round, and both leaves and leaf sheaths are hairless. (Joseph M.
DiTomaso, University of California–Davis, Bugwood.org.) D. The inflorescence is a narrow spike. (Joseph M.
DiTomaso, University of California–Davis, Bugwood.org.)

height of 6–8 ft. (1.8–2.5 m) by fall. Stems are round, spongy, and hollow, decreasing to less
than 0.3 in. (8 mm) in diameter toward the top. Gray-green leaves, 8–22 in. (20–55 cm)
long and 1–2 in. (2.5–5 cm) wide, are flat when fresh and sharply pointed at the tips. The
green color of healthy young shoots and leaves is often streaked with red or purple just
below the sediment surface. Both upper and lower leaf surfaces, as well as the leaf sheaths,
are hairless, but the upper leaf surface is ribbed. Ligules are hairy.
The tough, whitish rhizomes are 0.15–0.25 in. (4–7 mm) wide and fleshy. Although the
rhizomes may extend deeply into the substrate, approximately 75 percent of the root system
is in the top 6 in. (15 cm), and more than 40 percent grow in the upper 2 in. (5 cm) of soil.
Smooth cordgrass blooms from July to November and has colorless flowers. The narrow
inflorescence is a spike-like panicle, 4–18 in. (10–45 cm) long and less than 1 in. (2.5 cm)
wide. It has 3–30 spikes or branches, each as long as 5 in. (12.5 cm). The branches, loosely
overlapping and loosely pressed against the main axis of the inflorescence, each have 5–35
spikelets, or flowers. The glumes may be glabrous or hairy.
Dense-Flowered Cordgrass: Dense-flowered cordgrass has a compact form, growing in erect,
tufted clumps. The slender stems are 1–5 ft. (0.3–15 m) tall. It has grayish foliage and narrow
leaf blades (approximately 0.25 in. (7 mm) wide. Leaf blades are 4.5–17 in. (12–43 cm) long
and inrolled when fresh. Its rhizomes are short and stout, about 0.4 in (1 cm) wide, and the
root mass is fairly shallow. Plants flower from April through July. The inflorescence is 2.5–12
in. (6–30 cm) long and less than 0.5 in. (1.3 cm) wide. Its 2–20 spikes, approximately 4 in
(11 cm) long, are overlapping and closely pressed to the axis. The glumes are short with sharp
bristles. Until 1984, this plant was misidentified as the native California cordgrass.
Salt Meadow Cordgrass: The decumbent and matted stems of salt meadow cordgrass form
a dense turf or sod. When not in bloom, it is distinguished by its thin, wiry stems which
reach 1–4 ft. (0.3–1.2 m) tall and its tightly furled, or inrolled, leaf blades. Both stems and
leaves are narrow at the base, less than 0.2 in. (5 mm) wide. Leaf blades are green, 4–20
in. (10–50 cm) long and very narrow, less than 3 mm wide. Plants are reddish purple at
the base. The plant flowers in late summer. The inflorescence, 1–9 in. (5–22 cm) long and
0.8–4 in. (2–9 cm) wide, is an open panicle. Several spikes, usually 1–4, but as many as
454 n GRAMINOIDS

13, branch at a 60° angle from

the stem axis. Each spike or
branch is 0.4–3 in. (1–8 cm)
long. Rhizomes are long and
slender, approximately 0.2 in.
(3 mm) wide.
Common Cordgrass: Com-
mon cordgrass is a fertile hybrid
between smooth cordgrass and
small cordgrass, a species native
to England. The cross, which
originated in England, initially
produced a sterile hybrid,
Spartina x townsedii, but back-
crossing produced the fertile
S. anglica. Although common
cordgrass may have many of
the same characteristics as
smooth cordgarss, its morphol-
ogy is highly variable, probably
due to its extensive hybridiza-
tion. It spreads to cover ex-
tensive meadows with large
monospecific stands. Height of
the stout stems is variable, usu-
ally 1–4.3 ft. (0.3–1.3 m), but
plants can be as short as 2 in.
(5 cm). Ligules have a hairy
fringe. Leaves may be green or
grayish-green, flat or inrolled,
persistent or falling. Blades are
Dense-flowered cordgrass is native to southern South America. (Native 14–18 in. (35–45 cm) long and
range adapted from USDA GRIN and selected references. Introduced 0.2–0.5 in. (5–12 mm) wide.
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the Flowering season is variable,
United States, and selected references.) usually July to November, but
sometimes as late as February.
The inflorescence, 4.5–15.5 in. (12–40 cm) long, is an erect and dense panicle with 2–12
slightly-spreading spikes. Spikes are 5.5–8.5 in. (14–21 cm) long.
Smooth Cordgrass x California Cordgrass Hybrid: Smooth cordgrass has hybridized with
native California cordgrass to the extent that few to no pure smooth cordgrass stands remain
in some regions. Hybridization has resulted in many different morphologies. Early hybrids
are usually taller than either parent, have larger flowers, and produce more pollen. Later
hybrids do not resemble either smooth cordgrass or California cordgrass, and may have
spiky leaves, an odd orange color, or other variations. Later, back-crossed generations
become increasingly more difficult to identify in the field on the basis of morphology.
Related or Similar Species. California cordgrass, native from Bodega Bay, north of San
Francisco, south to Baja California, is shorter, less than 4.5 ft. (1.4 m) tall by autumn. It usually
has no red pigment when healthy, and its leaf blades are 6–18 in. (15–45 cm) long. This native
 CORDGRASSES AND THEIR HYBRIDS n 455

species blooms from June to

September. The inflorescence,
3.5–10 in. (9–25 cm) long, is a
dense cylinder, with spikes that
are closely overlapping and
pressed against the main axis.
The native cosmopolitan bul-
rush, also called saltmarsh club-
rush or alkali bulrush, has grass-
like leaves with reddish-purple
bases. As a sedge, its stems are
triangular in cross-section. The
unbranched inflorescence is a
short open umbel, 4–8 in. (10–
25 cm) long, with a cluster of
2–15 spikelets. Each brown
flower is oval or cylindrical,
0.5–1 in. (1.3–2.5 cm) long.
Sea-side arrowgrass, also a
native plant, grows 8–30 in.
(20–76 cm) tall with very slen-
der stems. The plants bloom
from summer to fall, and the
inflorescence is an unbranched
spikelike raceme, 4–16 in.
(10–40 cm) tall. The six-parted
green or brown flowers are sho-
wy and grow on short pedicels.
Introduction History. Al-
though the date is disputed,
smooth cordgrass may have been
introduced to Willapa Bay, Native to the Gulf and Atlantic coasts of North America and to the
Washington, in 1894, mixed Caribbean, salt meadow cordgrass displaces native coastal grasses in
with packing material for eastern Pacific coast states. (Native range adapted from USDA GRIN and selected
oysters shipped into the state for references. Introduced range adapted from USDA PLANTS Database,
propagation. It also may have Invasive Plant Atlas of the United States, and selected references.)
arrived as ballast in ships in the
early 1900s. Because the plant is important for sedimentation and stabilization in its native
range, it was intentionally introduced into Puget Sound in the 1940s for shoreline stabilization
and salt-marsh restoration. A small amount was planted in the Siuslaw River estuary, Oregon,
in the 1970s for the same reasons, but when the plot rapidly increased to an acre in size by
the 1990s, the grass was recognized as invasive and eradicated. Smooth cordgrass was also
deliberately planted in San Francisco Bay in the early 1970s for stabilization and restoration.
Dense-flowered cordgrass was probably inadvertently brought to Humboldt Bay in 1850
as ballast on ships carrying lumber back to Chile. It was recognized as an alien pest in the
1980s, and is the alien species which dominates the salt marshes in Humboldt Bay. Dense-
flowered cordgrass was introduced into Marin County, California, for a landscape restoration
project, and some plants occur in San Francisco Bay.
456 n GRAMINOIDS

A small patch of salt

meadow cordgrass occurs at
Southampton Marsh in the
San Francisco Estuary. The
plant is also in Siuslaw Estuary
in Oregon and also on the west
side of Puget Sound in
Washington.
Small patches of common
cordgrass are visible on a 1939
aerial photo of the Siuslaw River
in Oregon, and two plants were
found in San Francisco Bay in
1970. Thought at the time to be
the sterile hybrid, common cord-
grass was introduced into Puget
A hybrid originating in England, common cordgrass is invasive to Pacific Sound in the early 1960s, as for-
coastal zones. (Native range adapted from USDA GRIN and selected age for cattle. It was also inten-
references. Introduced range adapted from USDA PLANTS Database, tionally taken from Puget Sound
Invasive Plant Atlas of the United States, and selected references.) to San Francisco Bay in 1977.
Habitat. Although each spe-
cies or hybrid has a preferable habitat, cordgrasses generally grow in saline or brackish water
on muddy banks, salt marshes, and beaches in the intertidal zone, where they are protected
from strong wave action. Plants may tolerate as much as 12 hours a day of flooding in water
with a pH of 4.5–8.5 and salinity of 10–60 ppt. It especially invades bare mudflats and sand-
flats in shallow estuaries, but is found on a variety of substrates, including clay, gravel, and cob-
ble. The hybrid (S. alterniflora x foliosa) of smooth cordgrass with the native California cordgrass
grows both lower and higher in the tidal zone in San Francisco Bay than does the native
California cordgrass, which has a more narrowly defined range. Cordgrasses do not grow in
freshwater, and are generally excluded from terrestrial sites by competition from other plants.
Reproduction and Dispersal. Cordgrasses reproduce and spread both by seed and by rhi-
zomes and root fragments. Plants may flower and set seed in their first year, but most do so after
2–3 years of growth. Due to low soil temperatures, plants may not flower and set seed in all loca-
tions where the species has been introduced. Although plants develop many inflorescences, little
seed is produced because cross-pollination (by wind) is required, a rare occurrence in isolated
patches or clones. Seeds ripen from October to January and, if deposited in mud, germinate from
February through May. Seeds are carried by tidal action and currents and may adhere to animals
and birds. No seed bank exists because seeds are viable for only one year and cannot survive des-
iccation. Most seedlings fail to survive winter storms, but the remainder grow quickly in spring.
Erosion, waves, and currents break off pieces of established plants, which are then dis-
persed by currents and tides. A fragment must have roots or rhizomes in order to grow.
Initial colonization consists of isolated clones, which grow outward in circular patches.
Spreading quickly by growth of rhizomes, frequently more than 3.3 ft. (1 m) per year, these
circular patches coalesce to become continuous meadows.
From April through September, plants grow rapidly. All the flowering stems die back in cold
weather and are usually washed away with tides, but young, green shoots remain all winter.
Impacts. Smooth cordgrass grows fast to create monotypic, impenetrable stands, altering
the habitat and negatively affecting plants, birds, and other animals. It threatens the native
 CORDGRASSES AND THEIR HYBRIDS n 457

California cordgrass by producing more seeds, which also have a better germination rate,
and grows 6–7 times faster. By hybridizing with California cordgrass, smooth cordgrass
endangers the genetic distinction of the native species. Both smooth cordgrass and its
hybrids are more vigorous than the native species.
By overgrowing and shading, it displaces native pickleweed, seaside arrowgrass, fleshy jau-
mea, and algae. Its propensity to colonize bare mudflats converts those bare sites and channels
to marshland, changing the ecosystem, which formerly provided prey-rich nurseries for juvenile
chum salmon and English sole. It reduces forage habitat for shorebirds and waterfowl, both
residential and migratory, including the endangered California Clapper Rail in San Francisco
Bay. Smooth cordgrass thickets also alter the habitat for the endangered salt marsh harvest
mouse, as well as for other animals. The loss of mudflat habitat also negatively affects
Dungeness crab, clams, and other marine life, and threatens commercial oyster beds. At its worst,
almost 30 percent of the mudflats in Willapa Bay, Washington, were dominated by smooth
cordgrass.
Smooth cordgrass stands trap sediment, which not only changes the invertebrate popula-
tion, but also clogs and fills in natural and flood control channels. Flood patterns are altered,
permanently changing the hydrological conditions where it invades.
Most alien Spartina species present the same problems, perhaps in slightly different eco-
systems. Although dense-flowered cordgrass has displaced native pickleweed, saltgrass,
and seaside arrowgrass in the lower- and mid-elevation salt marshes in Humboldt Bay, it
especially threatens the high-elevation marsh, which has the most biodiversity and two rare
annuals, Humboldt Bay owl’s clover and Pt. Reyes bird’s beak. It has also begun to colonize
tidal mud flats, where its stem and root density alters marsh habitat for rodents, birds, inver-
tebrates, crustaceans, and gastropods. A large amount of dead matter accumulates at the
upper tidal limit of the upper marsh.
Management. A clonal patch of smooth cordgrass may live for more than 100 years, and
given its coastal location, it is likely that plants will continue to spread along the West Coast.
Many cordgrass stands grow in deep mud that is impossible to approach on foot or by boat,
a complication to management strategies.
Small populations, covering fewer than 5 ac. (2 ha), may be controlled by a number of
physical methods. Plants rooted in mud, especially small plants or seedling, are easily
extracted, but a shovel may be needed for harder substrates. Larger plants of dense-
flowered cordgrass may be pulled out because that species grows in clones that are more
clumped, with a shallower root system. Plants that are removed should be deposited above
the high-tide line, where they will dry and die. Small infestations, 3.3–33 ft. (1–10 m) in
diameter, may be solarized, buried under a 100 percent shadecloth or black plastic.
Mowing plants first may facilitate covering, which is best done in spring. Although plants
may die within four months, it is best to keep the area covered and monitored for two years.
Tarping, of course, will kill everything beneath the cover. In climates where they go dor-
mant, plants may be mowed to mud level in late fall. In areas of year-round growth, mowing
eight or more times a year may be effective in reducing shoot density and seed production.
Chemical applications are preferred when digging would be too damaging to the environ-
ment. Glyphosate failed to eliminate cordgrasses, and imazapyr is now the only herbicide
used for effective control. It is the most economical herbicide that is also safe for the workers
and the natural environment. Spring applications are best. Significant progress has been
made toward eradicating cordgrasses in both Washington and in California.
Although snow geese like to eat the roots of smooth cordgrass in Florida, few biological con-
trol agents have been approved because they may also attack the native and endemic California
458 n GRAMINOIDS

Eradication Success

I n only three years, an invasion of smooth cordgrass in Humboldt Bay, California, increased
from 100 sq. ft. (9.3 m2) to almost 5,000 sq. ft. (465 m2). By isolating the area with dikes,
mowing the plants, applying excess salt, and covering the area with black plastic, the
California Department of Fish and Game succeeded in eradicating the stand, preserving the
local mudflat ecosystem.
Two years of repeatedly mowing a small experimental plot in the nearby Mad River Slough
killed all the dense-flowered cordgrass plants, allowing native plants to recolonize. No seed
was recruited from adjacent areas, and the area remained clear of dense-flowered cordgrass.
When similar treatment was applied to a larger, 15 ac. (6 ha) area, mowing also killed the cord-
grass, but seeds were carried in by waves and currents. Although young cordgrass plants were
selectively torched in spring, the growth of native species complicated the eradication efforts.
This experiment indicated that eradication efforts must have a regional approach to avoid seed
recruitment from elsewhere. (Humboldt Bay National Wildlife Refuge Complex, 2010.)

cordgrass. A planthopper (Prokeleisa marginata), which successfully stunts or kills smooth cord-
grass in greenhouse experiments, has been approved for release into Willapa Bay in Washington
State. Smooth cordgrass in other locations, however, proved to be tolerant of the planthopper.
The marsh periwinkle (Littoraria irrorata), native to the Atlantic and Gulf coasts, is a sea snail
that grazes on and causes a fungus to develop, which weakens but does not kill plants.

Selected References
Daehler, Chris. “Spartina alterniflora.” In Invasive Plants of California’s Wildlands, edited by Carla C. Bossard,
John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000. http://www
.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=75&surveynumber=182.php.
Howard, V. “Spartina alterniflora.” U.S. Geological Survey Nonindigenous Aquatic Species Database,
Gainesville, FL, 2010. http://nas.er.usgs.gov/queries/FactSheet.aspx?speciesID=1125.
“Invasive Spartina in Humboldt County.” Humboldt Bay National Wildlife Refuge Complex, 2010.
http://www.fws.gov/humboldtbay/spartina.html.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Spartina alterniflora (Grass).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=792.
San Francisco Estuary Invasive Spartina Project. Coastal Conservancy. n.d. http://www.spartina.org/
species.htm.

n Crimson Fountain Grass

Also known as: African fountain grass, crimson fountain grass, tender fountain grass
Scientific name: Pennisetum setaceum
Synonyms: Pennisetum ruppelii, P. macrostachyon, P. rueppelianum, Phalaris setaceum,
Cenchrus setaceus
Family: Grass (Poaceae)
Native Range. North Africa, from Morocco east to Egiypt and south to the Sudan and
Ethiopia. Also on the Arabian Peninsula. Possibly native to eastern Africa as far south as
Zimbabwe.
 CRIMSON FOUNTAIN GRASS n 459

Distribution in the United

States. From Oregon and
California on the West Coast, east
to Colorado and Texas, and in the
southern states of Arkansas,
Louisiana, Kentucky, Tennessee,
and Florida. Also in Hawai’i.
Description. Fountain grass
is a coarse perennial grass that
forms dense clumps of erect
stems as tall as 1.5–5 ft. (.5–
1.5 m). The rigid leaf blades
are 8–26 in. (20–65 cm) long
and only 0.08–0.14 in. (2–
3.5 mm) wide. Leaf sheaths are
1.5–3 in. (4–8 cm) long, and
either smooth or hairy. The leaf
color depends on water avail-
ability. It is greener with more
water, but changes to purplish
and then tan as plants dry. Leaf
margins often have white hairs.
The fibrous root system extends
as deep as 12 in. (30 cm) into
the soil, and has no rhizomes.
Depending on climatic
region, crimson fountain grass
may have a long flowering
season, from January to Novem-
ber. Inflorescences are a long
panicle-like spike, with small,
densely grouped pink or purple Crimson fountain grass continues to spread throughout the Southwest,
flowers and many bristles. The displacing desert scrub. (Native range approximated from USDA GRIN
prominent, feathery flowerheads, and selected references. Introduced range adapted from USDA PLANTS
3–15 in. (7.5–38 cm) long, Database, Invasive Plant Atlas of the United States, and selected references.)
resemble bottlebrushes. Spikelets are usually 0.18–0.27 in. (4.5–7 mm) long and very narrow.
Of the two florets on each spikelet, the lower is usually sterile. The many short, slender bristles
remain on the spikelet when it detaches from the rachis. Seeds are yellowish brown and smooth.
Related or Similar Species. Several Pennisetum species, introduced primarily as ornamen-
tal garden plants, have become invasive. Most are tall plants with showy panicles. They often
have more than one common name and may be difficult to distinguish. Missiongrass, also
called feathery pennisetum, is native to tropical Africa and India. It is now distributed on
many Pacific islands, including Hawai’i, as well as in Florida and Puerto Rico. It is a tufted
annual or perennial with few simple or branched culms 1.6–10 ft. (0.5–3 m) tall. The nar-
row leaf blades are 2–17.5 in. (5–45 cm) long, either smooth or hairy. The inflorescences
are dense yellow-brown spikes, 2–10 in. (5–25 cm) long. Spikelets, with two florets on each,
are purplish or yellow brown and surrounded by many bristles. It is sometimes confused
with kyasuma grass or with elephant grass. Missiongrass predominantly invades dry
460 n GRAMINOIDS

A. Plants are often used as attractive ornamentals. B. Leaf sheaths and leaf margins may be hairy. C. The
inflorescence is a showy plume. (Joseph M. DiTomaso, University of California–Davis, Bugwood.org.)

rangeland, grasslands, and disturbed sites. Adapted to low fertility and sandy soils, it grows
quickly and is somewhat shade tolerant. It has been deliberately introduced as a pasture
grass in some areas and also spreads by vehicles. It outcompetes native species, preventing
regeneration of native trees and shrubs. Its dense tussocks alter the normal fire regime,
and it spreads quickly after a fire. It can also reproduce from cuttings. Management is similar
to fountain grass.
Feathertop, a showy tufted perennial grass native to northeastern Africa, is scattered in
several states, including California, Texas, Colorado, Michigan, Georgia, and Hawai’i. It is
distinguished by the presence of rhizomes and its white flowers. Growing to 2.6 ft. (0.8 m)
tall, it invades disturbed sites such as roadsides. Leaf blades are 2–15.5 in.
(5–40 cm) long, and sheaths are either glabrous or slightly hairy. Rhizomes are short, wiry,
and branched, reaching a depth of about 8 in. (20 cm). The dense fibrous root system may
be 24 in. (60 cm) deep. Greenish-white to straw-colored flowers grow on short, spike-like,
panicles, 1.5–4 in. (4–10 cm) long. The many bristles are straw color or white and can be 2
in. (5 cm) long, while the lower parts appear feathery. Seeds are yellowish brown, often with
a purple tinge. This species tolerates saline conditions and frequent mowing, and recovers
after a fire. Because it also invades coastal scrub, it has the potential to become noxious.
Elephant grass is an annual or perennial grass found in California, Texas, Florida,
Hawai’i, Puerto Rico, and the Virgin Islands, where it invades forests, grassland, riparian
sites, and coastal beaches. This stout plant has a vigorous root system of creeping rhizomes
6–10 in. (15–25 cm) long. Leaf blades, 1–3 ft. (.3–.9 m) long and 0.4–1.2 in. (1–3 cm) wide,
terminate in a fine point. The smooth culms, 6.5–16.5 ft. (2–5 m) tall, branch toward the
top. Yellow-brown inflorescences are compact, with bristly spikes 3–12 in. (8–30 cm) long.
Spikelets, with two florets each, are surrounded by several short bristles, and one bristle as
long as 1.5 in. (4 cm). Elephant grass grows in a wide range of habitats but is tallest in moist,
rich soils where it can create dense reedy thickets to 10 ft. (3 m) tall. Seeds are rarely pro-
duced. This species reproduces primarily vegetatively and resprouts vigorously when cut.
Like other Pennisetum species, it displaces native plants. Digging out the plants is an ineffec-
tive method of control because of the extensive rhizome system. It is best to treat it with a
herbicide.
African feathergrass occurs in California, Texas, and Hawai’i. A tussock-forming peren-
nial grass that reaches 3.2–6.5 ft. (1–2 m) tall, it resembles pampas grass (see Graminoids,
Jubata Grass and Pampas Grass), but is native to southern Africa. Leaf blades, which grow
from both the base of the plant and from the stems, are either light or dark green, sometimes
with bluish-purple edges. They reach 4 ft. (1.2 m) in length, have pronounced ribbing on
 CRIMSON FOUNTAIN GRASS n 461

the upper surfaces, and are slightly curved rather than flat in cross-section. They have ser-
rated edges and hairy sheaths. The root system is extensive. Stout rhizomes can be 6.5 ft.
(2 m) long, and both rhizomes and fibrous roots extend 3.2 ft. (1 m) deep into the soil.
Inflorescences are 3–12 in. (7.5–30 cm) long, with prominent bristles, about 0.4 in.
(1 cm) long. Yellow-brown seeds mature in late summer and fall and are dispersed by tiny
barbed hairs that catch in fur and clothing. Like many other feathergrasses, it was imported
for use as an ornamental, often for garden ponds, and now invades disturbed areas and
roadsides. African feathergrass prefers damp riparian habitats in full sun, where its
unchecked growth can block waterways. Young plants need moisture, but become drought
tolerant as they mature. Dry plants pose a fire hazard. It reproduces both by rhizomes and
abundant seed production, spreading quickly and crowding out native species. Physical
control is made worse by the difficulty of digging out the extensive root system.
Kyasuma grass, currently found in Florida, is native to northern tropical Africa and India.
It is a summer annual that propagates solely by seed and grows to 3.2 ft. (1 m) tall. Leaf
blades, 6–10 in. (15–25 cm) long, are flat and glabrous. Culms are freely branching, both
from the base and higher on the stem. The attractive pinkish-brown flowerheads grow on
a panicle-like spike, 2–6 in. (5–15 cm) long. Many bristles, each 0.2–0.4 in. (5–10 mm)
long, surround the spikelets, giving it a wooly look, but one bristle is conspicuously longer,
0.6–1 in. (16–24 mm). Although kyasuma grass is useful for soil stabilization and provides
good forage, hay, and silage because it can be cut several times a year, it can become an inva-
sive plant in disturbed sites, roadsides, fallow fields, or cultivated crops.
Several cultivars of fountain grass have been developed, with green, reddish, or purple
foliage. Although cultivars are supposedly sterile, some do set seed and may become a
problem.
Introduction History. Crimson fountain grass has been introduced to both Hawai’i and
mainland United States several times as an attractive ornamental grass in landscape plant-
ings. The first records of the plant are dated 1914 in Hawai’i and 1917 in California. It has
now become naturalized in the southwestern states and is spreading in southern California
and in the deserts of the southwestern United States and Mexico, where it is converting des-
ert scrub to grassland. Unsuitable as a pasture grass, it has become a troublesome weed.
Habitat. Tolerant of many conditions, fountain grass can be found growing on several
habitats, from lava flows to rangeland, in a wide elevation range. Plants grow up to 2,000 ft.
(600 m) elevation in desert areas of California and Arizona, and in Hawai’i, from sea level
to the subalpine zone at 9,185 ft. (2,800 m). It is very drought tolerant and grows where
annual precipitation is less than 50 in. (1,270 mm). It is a common desert plant in grasslands,
washes, canyons, and roadsides. In California, fountain grass is found on disturbed coastal
dunes, coastal sage scrub, and inland desert canyons and shrubland. In spite of its coastal
locations, it does not tolerate saline soils. Roadsides are an especially common habitat near
urban centers where ornamental plants have escaped cultivation. Seeds also germinate and
grow in rock crevices and broken pavement.
Reproduction and Dispersal. Fountain grass can produce viable seeds with or without
fertilization. Pollination is not necessary because the female plants can produce seeds asex-
ually, from other plant cells, not just from egg cells. However, even when flowers are polli-
nated, few seeds develop. Seeds are wind dispersed and remain viable in the soil at least
seven years. Long-distance dispersal takes place by water, vehicles, livestock, and human
activity.
Plants grow quickly. In just five years, a plant can become 4 ft. (1.2 m) tall, with a basal
diameter of 12 in. (30 cm). Individual plants may live for 20 years.
462 n GRAMINOIDS

Impacts. In addition to replacing native plant species, the most serious impact of fountain
grass is how it changes fire regimes. Plants accumulate dead biomass, providing more fuel,
which results in increased intensity and spread of fires. It is an aggressive, fire-adapted
invader that quickly colonizes burned areas and outcompetes native plants, eventually creat-
ing single-species stands. As a major threat to Hawaiian ecosystems, it replaces native species
not adapted to fire. Fountain grass can cause forests in Hawai’i to convert to grassland, and it
can replace coastal sage in California.
Management. Because seeds are long lived, continued monitoring of a site is necessary.
Control should begin at the edges of infestations, to prevent it from spreading further.
Once fountain grass is successfully removed, it is important to reseed with native species.
Physical removal by hand-pulling is only feasible in small stands or with small plants. It is
difficult to break up plants that have a base larger than 6 in. (15 cm) in diameter. Hand-
pulling should be done several times a year to prevent new sprouts from becoming estab-
lished. The inflorescence must be properly disposed of, such as placing in plastic bags, to
prevent seed dispersal. Because fountain grass is fire adapted, burning is a very poor option.
Chemical control may be effective when used in combination with physical methods.
Preemergents, such as hexazinone, used after plants have been removed will prevent seeds
from sprouting for 9–12 months. A systemic, such as glyphosate, applied to leaves does
not provide consistent results.
Nothing has been proposed for biological control of fountain grass. Any insects would
most likely also affect related forage and crops, such as kikuyugrass and pearl millet.
Although kikuyu grass (see Graminoids, Kikuyugrass) is also an invasive pest in some
regions, it is an important forage crop for cattle in Hawai’i.

Selected References
Benton, Nancy. “Fountain Grass.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/archive/plants/
alien/fact/pese1.htm.
IUCN SSC Invasive Species Specialist Group (ISSG). “Cenchrus setaceus (Grass).” ISSG Global Invasive
Species Database. 2010. http://www.issg.org/database/species/ecology.asp?si=309.
Lovich, Jeffrey. “Pennisetum setaceum.” In Invasive Plants of California’s Wildland, edited by Carla C.
Bossard, John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000.
http://www.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=66&survey
number=182.php.

n Giant Reed
Also known as: Arundo, Spanish reed, wild cane, carrizo
Scientific name: Arundo donax
Synonyms: Arundo versicolor
Family: Grass (Poaceae)
Native Range. Eastern Asia, northern India, and Nepal. Introduced to and cultivated in
other regions of Asia, southern Europe, North Africa, and the Middle East for thousands of
years.
Distribution in the United States. Southern half of the United States, from Illinois south
to the Gulf of Mexico and from the Atlantic Ocean to the Pacific Ocean. Primarily problem-
atic along waterways in the desert states.
 GIANT REED n 463

Description. One of the larg-

est herbaceous species in the
grass family, giant reed is a per-
ennial that grows 6.5–30 ft. (2–
9 m) tall. Smooth stalks are
0.25–2 in. (0.6–5 cm) thick
and hollow between the nodes.
After the first year of growth,
stems branch sparingly from
the nodes. Continually cutting
the culms causes the plant to
branch profusely. Pale, green or
blue-green leaves, which may
be either flat or folded, grow
alternately all along the stalk
and are two-ranked, meaning
that they grow on opposite sides
of the stem, not all around it.
The leaf blades are long, 1–2 ft.
(0.3–0.6 m), and narrow, 1–2.5
in. (2.5–6 cm) wide, gradually
tapering to a point. The bases of
the leaves where they join the
stalk are heart-shaped with a
hairy tuft. Leaf margins are
rough to the touch.
Roots are tough and fibrous,
penetrating deeply into the soil.
Creeping rhizomes are thick
and fleshy.
Long, 2–3 ft. (0.6–1 m), plu-
mes of cream or light-brown Giant reed colonizes moist areas along ditches and riverbanks. (Native
flowers, borne in a panicle at range approximated from USDA GRIN and selected references.
the top of the stalks, appear as Introduced range adapted from USDA PLANTS Database, Invasive Plant
early as March, but mostly in Atlas of the United States, and selected references.)
August and September. Many
spikelets, 0.5 in. (1.2 cm) long with several florets, branch from the flower stalk. Flowers
are densely packed on the inflorescence, and spikelet branches are upright, not drooping.
Related or Similar Species. Giant reed resembles other tall members of the grass family,
including bamboo, cultivated corn, and common reed (see Graminoids, Common Reed).
Common reed also has smooth, hollow stems, but the leafy stems do not branch.
Inflorescences are purple or golden, in a bushy panicle usually 12 in. (30 cm) long.
Introduction History. Because it has useful properties, giant reed was intentionally intro-
duced to North America, probably by the Spanish, and has also been introduced to most
subtropical and warm temperate areas of the world. It was purposely planted on stream-
banks to control erosion and as an ornamental in warmer areas of the United States. Stalks
and leaves were also used for fencing, thatching, and other building materials. It was first
known in the Los Angeles area in the early 1800s.
464 n GRAMINOIDS

A. Plants can be as tall as 30 ft. (9 m). (Chuck Bargeron, University of Georgia, Bugwood.org.) B. Leaves are two-
ranked on the stem. (James H. Miller, USDA Forest Service, Bugwood.org.) C. The large inflorescence is a feath-
ery panicle. (David J. Moorhead, University of Georgia, Bugwood.org.) D. The heart-shaped ligules are hairy.
(Amy Ferriter, State of Idaho, Bugwood.org.) E. Culms grow in dense stands. (Bonnie Million, BLM, Ely District,
Bugwood.org.) F. Buds on the rhizomes sprout many new plants. (Joseph M. DiTomaso, University of
California–Davis, Bugwood.org.)

Habitat. Giant cane grows best in moist conditions, where water is available but soil is
well drained. Riparian environments along ditches, streams, and riverbanks are ideal.
Plants also tolerate both a high water table and dry upland areas along roads. Any type of soil
is suitable, from clay to sand, and plants are tolerant of saline conditions. Although young
plants are susceptible to drought, stands older than 2–3 years can withstand dry conditions
because roots are not only drought resistant, but extend deep into the soil to extract mois-
ture. Plants can survive low temperatures when dormant but not when growing. They are
damaged by spring frosts.
Reproduction and Dispersal. Not much is known about the viability of seeds or germina-
tion requirements, but studies have shown that seeds produced from plants in California are
not viable. Commercial stands are planted by use of rhizomes, and natural reproduction is
almost exclusively vegetative. Giant reed spreads when stem and root fragments float down-
stream. Floods break off pieces of the rhizomes or stalks, and any that contain nodes will
readily sprout a new plant.
Impacts. Giant reed causes a number of problems, including flooding, erosion, habitat
loss, and increased fire danger. The plant forms dense thickets that clog stream channels
and ditches. One clump can cover several acres. Root masses can be more than 3.3 ft.
(1 m) thick, stabilizing banks and altering stream flow. The interconnecting roots form mats
that block water flow under bridges and into culverts, where eventually the weight of the
mat and trapped debris can damage structures. In contrast to bank stabilization, giant reed
may increase bank erosion. During flood events, masses of vegetation can break off from
the bank, with the entrwined roots and rhizomes carrying the soil as well.
Because the plant grows very fast, up to 2 in. (5 cm) a day, giant reed outcompetes and
totally displaces native plants, which reduces habitat for wildlife, including federal endangered
species such as Least Bell’s Vireo and Willow Flycatcher in California. Native stands of willows
and cottonwoods depend on periodic flooding and redeposition of silt, which is required for
the evolution of the flood plain community. The resultant diversity in the plant community
 GIANT REED n 465

is necessary as nesting sites for several species of birds. Giant reed provides no food or nesting
sites for natives. Leaves of giant reed contain noxious chemicals such as alkaloids, which give
the plant natural protection from insects and grazers, another reason why the plant does not
support wildlife. Stands of giant reed provide less shade than do native riparian plants, result-
ing in higher water temperature that limits aquatic diversity, including fish species.
Because giant reed produces abundant plant material that is highly flammable, especially
when dry, fire potential in normally moist riparian habitats is increased. In wild stands of
giant reed, the amount of dried plant material per acre can be as much as 8.3 tons (20 tons
per ha). Whereas the moist native vegetation serves as a firebreak, the flammable giant reed
does not. Its height also leads flames into crown fires in nearby vegetation. Buried rhizomes
quickly send up new shoots after a fire, outcompeting native plants.
Because of its large size and mass, giant reed stands consume a considerable amount of
water in arid regions.
Management. A watershed approach must be used to control or eradicate giant reed.
Eradication of all plants and rhizomes in the upper reaches of streams will prevent reestab-
lishment of the plant downstream. Physical removal must include total removal of all plant
debris to prevent fragments of stems or rhizomes from re-infesting the site. However, cut
and dried chips will not sprout. Hand removal, although useful for small sites because it is
selective, is labor intensive and expensive. Roots and rhizomes must be dug out.
Mechanical removal by cutting or mowing is not selective and appropriate only for pure
stands of giant reed where no native vegetation remains. Repeated mowing to deplete root
reserves may be effective. If only one mowing can be accomplished, it is best done just
before flowering to prevent production of any seeds. Prescribed burning, whether done
selectively or not, should be done after flowering. Although fire does not destroy the roots,
it may destroy the seed bank in the soil. Grazing is not a preferred method of reducing plant
mass. Cattle find giant cane unpalatable, but will eat it during the dry season. Angora goats,
and possibly sheep, are better options.
Any physical removal should be followed by chemical application, with a herbicide that is
appropriate for use in wetlands or riparian habitats. Systemic herbicides, such as glyphosate,
should be applied on the whole plant after flowering but before dormancy, in mid-August to
early November, when plants are actively translocating nutrients to the roots. Herbicides can
also be applied on cut stalks or new growth. Areas may be reseeded with natives after the
cane density has been reduced.

Beneficial Uses of Giant Reed

G iant reed has been used by civilizations for thousands of years. Use for thatch-
ing and building materials is widespread. Ancient Egyptians lined grain stor-
age pits with leaves and sometimes wrapped mummies in the leaves. It has been
used in construction of musical instruments, such as pan pipes and bagpipes.
Because no good substitute exists, giant reed is still grown commercially, especially
in Argentina, China, Australia, Spain, Italy, and France, as a source of reeds for use
in musical instruments. Rhizomes are beneficial in medicine, as a sudorific, diuretic,
and antilactant. Additional uses include basketry, fishing rods, trellises, poultry
pens, and fodder.
466 n GRAMINOIDS

Little is known about insect predators or pathogens that can be used for biological con-
trol. The green bug (Schizaphiz graminum) feeds on giant cane in winter. Phothedes dulcis cat-
erpillars in France may eat it, as will Zyginidia guyumi in Pakistan. A moth borer (Diatraea
saccharalis) may be effective in Barbados. In the Mediterranean region, some damage occurs
from corn borers, aphids, and spider mites. In the United States, plants are somewhat sus-
ceptible to root rot, lesions, crown rust, and stem speckle, but all result in little damage.

Selected References
Bell, Gary P. Ecology and Management of Arundo donax, and Approaches to Riparian Habitat Restoration in
Southern California. Santa Fe: Nature Conservancy of New Mexico, 2002. http://www.invasive.org/
gist/moredocs/arudon01.rtf.
Benton, Nancy, Gary Bell, and Jil Swearingen. “Giant Reed.” Weeds Gone Wild: Alien Plant Invaders of
Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://
www.nps.gov/plants/ALIEN/fact/ardo1.htm.
Dudley, Tom L. “Arundo donax.” In Invasive Plants of California’s Wildlands, edited by Carla C. Bossard,
John M. Randall, and Marc C. Hoshovksy. Berkeley: University of California Press, 2000.
http://www.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=8&survey
number=182.php.
Hoshovsky, Marc. “Element Stewardship Abstract for Arundo donax.” Global Invasive Species Team,
Nature Conservancy, 1996. http://www.invasive.org/gist/esadocs/documnts/arundon.pdf.

n Japanese Stilt Grass

Also known as: Chinese packing grass, Nepalese browntop, eulalia
Scientific name: Microstegium vimineum
Synonyms: Eulalia viminea, Andropogon vimineum, Microstegium imberbe
Family: Grass (Poaceae)
Native Range. Southeastern part of Asia, including southern China, Korea, Japan,
Taiwan, Southeast Asia, and the Philippines.
Distribution in the United States. Most states east of the Mississippi River and along the
Gulf coast, north to southern New England, and continuing to spread northward.
Description. Growing in dense mats, Japanese stilt grass is a sprawling annual grass that
can be 1–3.5 ft. (0.3–1 m) tall. The usually decumbent stems are as long as 3.3 ft. (1 m). The
stems are weak, usually branched, smooth, and glabrous. The pale, lime-green leaves, alter-
nate on the stem, are lance shaped, 1–3 in. (2.5–8 cm) long and as wide as 0.5 in. (1.25 cm).
Leaves are distinctly tapered at each end, causing the plant to resemble a small bamboo. Both
leaf surfaces are sparsely pubescent, and the leaf’s midrib on the upper surface has a pale-
silver stripe of reflective hairs that make it shiny. The ligules are membranous and fringed
with short hairs. Plants take on a slight purple tinge in fall before dying. Roots are shallow,
and stems root where nodes touch the ground.
Appearing late summer to early fall, the inflorescence is a spike-like panicle with 1–6
racemes, each 1–3 in. (2.5–7.5 cm) long. Plants have multiple inflorescences, both from
the end of the stem and from axils where the stems branch. The spikelets occur in pairs
and are coarsely hairy. Glumes are 0.2 in. (5 mm) long, with no awns. The lemmas either
have no awn or a slender awn, 0.1–0.3 in. (4–8 mm) long. Florets on the inflorescence at
the end of the stem are chasmogamous (cross-pollinated), while those on the inflorescences
in the axils are cleistogamous (self-pollinated). Small yellowish reddish seeds ripen from
September to November.
 JAPANESE STILT GRASS n 467

Related or Similar Species.

Virginia cutgrass, also called
white grass, which often grows
with and resembles Japanese
stilt grass, can be distinguished
by its glabrous leaf sheaths,
spikelets with only one flower,
and lemmas that sometimes
have a hairy fringe. Jointed
grass, an invasive alien species,
has short, broad leaves, 2.5 in.
(6.3 cm) long and 0.75 in.
(2 cm) wide, with hairy leaf
bases that encircle the stem.
Introduction History. Jap-
anese stilt grass was first col-
lected in Tennessee in 1919. It
spread rapidly, and, by 1960,
was present in several midwest-
ern, southeastern, and Atlantic
states. It appears to be expand-
ing its range and was recently
found along the Hudson River
in New York and in Connec-
ticut. Its use as packing material
to protect porcelain shipments
from Asia is the probable cause
of its initial introduction, and it
most likely was dispersed in
transported hay or soil.
Habitat. Adapted to low
light conditions, the plant pre- Tolerant of both light and shade, Japanese stilt grass may form mats in
fers shady, moist locations and both open areas and woodlands in southeastern states. (Native range
is commonly found in wood- adapted from USDA GRIN and selected references. Introduced range
lands near streams, forested adapted from USDA PLANTS Database, Invasive Plant Atlas of the
wetlands, floodplains, and United States, and selected references.)
swamps. Also able to grow in
high light and moderately dry soils, it grows in uplands, such as fields, clearings, paths,
seldom-used roadbeds, and road and utility right-of-ways. Japanese stiltgrass does not toler-
ate wet soil or standing water and does not grow in clay soils. It invades primarily acidic to
neutral soils with high nitrogen content, but can also be found on neutral soils overlying
limestone and marble. It easily invades areas that are regularly mowed, tilled, subject to foot
traffic, or affected by any action that causes a soil disturbance. Naturally disturbed areas,
such as scoured stream beds or banks caused by flooding, may also be invaded. A disturb-
ance to a forest that creates a canopy opening will also stimulate the plant to spread.
Upland forests in Pennsylvania were colonized after defoliation by gypsy moth. The coldest
winter temperature where populations of Japanese stilt grass are considered to be invasive is
approximately −7ºF (−22ºC).
468 n GRAMINOIDS

A. This groundcover is a monotypic stand of Japanese stilt grass. (Chris Evans, River to River CWMA,
Bugwood.org.) B. Decumbent stems grow in a dense mat. (Chuck Bargeron, University of Georgia,
Bugwood.org.) C. Florets grow close to the stem. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology
University of Connecticut, Bugwood.org.) D. Lance-shaped leaves taper to a point at both ends. (David J.
Moorhead, University of Georgia, Bugwood.org.)

Reproduction and Dispersal. Japanese stilt grass reproduces both sexually and asexually.
Seeds are most likely responsible for populations that develop in new locations, but the
plant’s ability to root at stem nodes explains its ability to colonize large areas. Although
one plant is capable of producing more than 1,000 seeds, 100 is the norm. Seeds have no
obvious mode of dispersal, and most fall close to the parent. They may be dispersed by water
flow during heavy rains and are frequent contaminants in hay, soil, or potted plants. Seeds
may also adhere to vehicles and shoes. Although the grass is not eaten by deer and other
browsers or grazers, those animals may aid in seed dispersal. Seeds remain viable in the soil
for seven or more years and can survive 10 weeks of submergence in water. Seeds germinate
in spring, and seedlings grow slowly during the summer.
Impacts. Japanese stilt grass can spread extensively to create dense, monotypic stands. In
disturbed areas, the grass can shade out and replace herbaceous ground cover in 3–5 years.
It grows more slowly in undisturbed sites. It crowds out and displaces native herbaceous
species in wetlands and forest floors. By changing the character of forest understory habitat,
it negatively affects ground-nesting birds. It replaces the nesting habitat of native Bobwhite
Quail and other wildlife, but provides good habitat for predators of ground-nesting birds,
especially cotton rats. The annual dieback of the grass results in a thick layer of thatch on
the ground. Although the leaves decompose quickly, the stems do not. Plants can alter soil
chemistry. Litter and organic horizons become thinner, and the pH is higher.
It is possible that Japanese stilt grass could negatively impact populations of the northern
pearly eye butterfly along the Potomac River in the Washington, D.C., area. The butterfly has
been observed laying eggs on Japanese stilt grass as an alternative host plant for its caterpil-
lars, but the relationship is not yet clear. White-tailed deer selectively feed on native grasses,
reducing competition and encouraging Japanese stilt grass to spread.
Management. Management priority should be to prevent Japanese stilt grass introduction
into noninfested areas. Because it is an annual with a shallow root system, physical removal
is feasible. Hand-pulling is best accomplished in mid-to-late summer when plants are larger,
and it is too late in the season for more seeds to germinate a new crop. Removing plants
before early July triggers sprouting from the soil seed bank. Pulled plants may be left to
dry on the surface, but plants in the fruiting stage should be bagged and disposed offsite.
Repetition of hand removal methods for several years will exhaust the seed bank. Mowing
 JOHNSONGRASS n 469

the grass in late summer, August and September, while the plants are in flower, but before
seed sets, will ensure that plants do not resprout the same season. Grazing is not an option
because livestock, even goats, do not eat it. Flooding for more than three months during
the growing season will kill mature plants but will not affect the seed bank. Prescribed burn-
ing in spring stimulates germination from the seed bank, but burning in late fall may be ben-
eficial in that it reduces the thatch layer. Burning the litter layer before application of
herbicides will allow the herbicides to reach more actively growing plants.
Chemical applications of systemics, such as glyphosate, are practical for large infesta-
tions. The preemergent imazapic has been effective in controlling Japanese stilt grass in some
regions, with the advantage that it can be sprayed throughout the summer.
No biological controls are known.

Selected References
“Invasive Plant, Japanese Stilt Grass (Microstegium vimineum).” The Nature Conservancy in
Connecticut, 2010. http://www.nature.org/wherewework/northamerica/states/connecticut/science/
art24048.html.
“Japanese Stilt Grass.” New Invasive Plants of the Midwest Fact Sheet. Midwest Invasive Plant Network,
n.d. http://mipn.org/Midwest%20Invasives%20Fact%20Sheets/PDF/Japstilt.pdf.
Swearingen, Jil M. and Sheherezade Adams. “Japanese Stiltgrass.” Weeds Gone Wild: Alien Plant
Invaders of Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://
www.nps.gov/plants/alien/fact/mivi1.htm.
Tu, Mandy, ed., and John Randall. “Element Stewardship Abstract, Microstegium vimineum.” Global
Invasive Species Team, Nature Conservancy, 2000. http://wiki.bugwood.org/Microstegium
_vimineum.

n Johnsongrass
Also known as: Aleppo grass, Johnson grass
Scientific name: Sorghum halepense
Synonyms: Sorghum miliaceum, Holcus halapensis
Family: Grass (Poaceae)
Native Range. Mediterranean region of northeastern Africa and Southwest Asia. Possibly
native to the desert areas as far east as Pakistan.
Distribution in the United States. Every state except Maine, Minnesota, and Alaska.
Description. Johnsongrass is a coarse, tall perennial grass that grows as high as 3–8 ft.
(1–2.4 m). It can form dense clumps, with stems branching from the base, or nearly solid
stands from the extensive growth of rhizomes. Stems are pink or reddish near the base. Leaf
sheaths are glabrous, but the membranous ligules have a white fringe. The smooth leaves,
alternate on the stem, are 0.6–2 ft. (0.2–0.6 m) long and 0.4–1.2 in. (1–3 cm) wide, with
rough margins. Leaves are distinguished by a prominent white midrib on the upper surface.
The fleshy rhizomes, as much as 0.4 in. (1 cm) in diameter and 6.5 ft. (2 m) long, are
scaly and sharply pointed. Most are in the top 8 in. (20 cm) of soil, but the rhizome mass
is deeper in light-textured soils and closer to the surface in heavy clay soils. Rhizomes attain
a much greater biomass than stems and leaves and may be as much as 90 percent of the total
weight of the plant.
Flowering occurs throughout the growing season, usually April to November. Flowers
occur in loosely branched panicles, 6–20 in. (15–50 cm) long. The numerous panicle
470 n GRAMINOIDS

branches grow upright and

whorled around the stem. The
inflorescence is purplish and
hairy, with short spikelets at
the ends of the branches.
Spikelets, 0.2 in. (5 mm) long,
grow in pairs, or occasionally
in groups of three, each with a
conspicuous awn. Although
each spikelet produces one
seed, plants produce many
reddish-brown seeds, 0.1 in.
(3 mm) long and with a con-
spicuous awn. Seeds ripen from
May through March.
Related or Similar Species.
Sorghum, also known as shat-
tercane, is originally from
northern Africa. The species
and its cultivars, either annual
or perennial, are cultivated for
the edible grain. Although inva-
sive in several states, this plant
lacks creeping rhizomes. Sudan-
grass, a subspecies of sor-
ghum, is an annual. Although
available for sale as an orna-
mental, it is also invasive in
some areas. Bulb panicgrass,
frequently mistaken for John-
songrass, can be distinguished
Johnsongrass, a weed in both crops and natural ecosystems, is especially by swollen tissue at the base of
problematic along the Gulf coast. (Native range adapted from USDA the culms and its much shorter
GRIN and selected references. Introduced range adapted from USDA rhizomes.
PLANTS Database, Invasive Plant Atlas of the United States, and selected Introduction History. Widely
references.) used in the world as a forage
crop, Johnsongrass was brought
to Carolina from Turkey in the early 1800s. It is called Johnsongrass because a farmer named
Johnson took the plant from South Carolina to Alabama in 1840. By the end of the nineteenth
century, Johnsongrass had spread to most of the United States.
Because it can be a nutritious fodder under the right growing conditions, it has been pro-
moted as a forage plant. Its spreading rhizomes make it useful for erosion control.
Habitat. Originally preferring Mediterranean or tropical climates, Johnsongrass has
evolved new ecotypes that enable it to grow in many environments and continue to expand
its range. Most ecotypes are frost sensitive, but some can overwinter in warmer climate
zones. The aerial parts of the plant die back in colder winter regions, but the rhizomes
may persist. Cold-tolerant ecotypes have been discovered in the northern United States
and in Canada.
 JOHNSONGRASS n 471

A. Johnsongrass grows in dense clumps that can be 8 ft. (1.2 m) tall. (Chris Evans, River to River CWMA,
Bugwood.org.) B. Leaf blades have a prominent white midrib. (Chris Evans, River to River CWMA,
Bugwood.org.) C. Flowers are open panicles. (James H. Miller, USDA Forest Service, Bugwood.org.)
D. Rhizomes are pointed and scaly. (Steve Dewey, Utah State University, Bugwood.org.) E. Seeds are plentiful.
(Steve Dewey, Utah State University, Bugwood.org.)

Johnsongrass is most common in agricultural land, such as crops, pasture, and aban-
doned fields, and in disturbed areas, such as right-of-ways, irrigated ditches, and the
trampled land around springs and stock tanks. It also grows at forest edges and in wetlands,
but requires open areas and is not found under closed canopies. It thrives in disturbed open
lowlands with rich soils, particularly cultivated fields. It grows best on moist, porous soils
and less vigorously on poorly drained clays. The rhizomes are intolerant of hot, dry condi-
tions and may be killed by exposure.
Reproduction and Dispersal. Johnsongrass produces both sexually and vegetatively. Plants
are self-fertile, which accounts for most of the seed production. It is a short-day plant; more
seeds are produced when daylight hours are fewer than 12. Seed production also depends on
the ecotype, varying 40–350 seeds per panicle. Seeds are spread by water and wind, and strong
winds in thunderstorms can carry seed as far as 0.6 mi. (1 km) distant. The twisted awns are
frequently found on farm equipment or caught in the coats of livestock. Seeds are able to sur-
vive the intestinal tract of birds or livestock, and are also a contaminant in hay and commercial
seed. Most seeds germinate the year after they are produced. Although dormancy varies with
the ecotype, some seeds enter dormancy immediately after ripening, with 50 percent capable
of germination after five years. After six years, viability drops to 2 percent. Germination is best
at 77–86ºF (25–30ºC), in light soils from a depth less than 3 in. (7 cm).
Rhizomes, which root at nodes, begin to grow 30–45 days after seeds germinate. They
grow more rapidly than shoots, and one plant can produce a total rhizome length of 200–
300 ft. (60–90 m) in one month. Shoots can emerge from rhizomes 4 ft. (1.2 m) deep in
the soil, and any small piece is capable of sprouting a new plant. Rhizomes may remain dor-
mant until conditions cause the buds to sprout.
Impacts. Considered by some to be one of the 10 worst weeds in the world, Johnsongrass
causes problems in most of the populated regions of the world, between 55º N and 45º S lat-
itudes, and is particularly troublesome along the Gulf coast of the United States. Its impacts
on both natural ecosystems and agriculture are many and varied.
It forms dense colonies, which displace native plant species and restrict the establishment
of tree seedlings, affecting natural succession. As a weed in cultivated fields, it shades out
crops and deprives them of needed moisture and nutrients. It is also alleleopathic to several
472 n GRAMINOIDS

crop species, including corn, sugar cane, grain sorghum, soybeans, sunflowers, wheat, bar-
ley, mustard, citrus, and cotton. It is an alternate host for insect, fungal, bacterial, and nem-
atode pathogens that affect crops. Its flowers cross-pollinate with S. bicolor, reducing yields
of commercial sorghum crops.
Ingestion of Johnsongrass by livestock is very toxic, especially when plants are stressed, and
can be fatal. Glycosides in the plant are converted into free cyanide in the digestive system of live-
stock. Certain feeds, such as alfalfa hay and linseed cake, provided along with Johnsongrass may
retard the development of free cyanide. Cyanide is a rapidly acting poison, and symptoms can
manifest just five minutes after the plant is ingested. Death can occur in 15 minutes or after sev-
eral hours. Symptoms may begin with salivation and labored breathing, then progress to muscle
tremors and incoordination. Venous blood becomes bright red, and convulsions and death may
follow. Plants can also accumulate toxic amounts of nitrate, which causes poisoning in sheep
and cattle. Animals can be treated for both cyanide and nitrate poisoning.
Rarely, Johnsongrass can cause sorghum cystitis in horses, a disorder that permanently
damages the spinal cord. The animal then loses control of its rear legs and bladder. Few
horses that eat the plant are affected, and the cause is not known. No treatment exists, and
the animal must be euthanized.
The excessive amount of dried plant matter can be a fire hazard in summer, and the pol-
len is a known allergen.
Management. Control of Johnsongrass is difficult, and eradication almost impossible
because of the plant’s extensive rhizome system, its abundance of seeds, and ability to grow
in various climates or environments. The best management is prevention, by maintaining land
in an undisturbed state. Any physical control should avoid soil disturbance. Light infestations
may be eliminated by hand-pulling. Plants should be pulled in June and all parts removed.
Heavy infestations may be mowed or tilled. Frequent summer mowing may deplete rhizome
reserves and prevent seed production. Mowing or grazing that keeps plants shorter than
12 in. (30 cm) tall may deplete the rhizomes stores and kill plants within 2–3 years. Winter cul-
tivation will expose rhizomes to the fatal effects of cold temperatures. Cultivation early in the
growing season, however, will spread rhizomes and make the infestation worse. Burning
encourages regrowth. Solarization may provide some success. Covering infested areas with
black plastic in hot climates will increase temperatures to a level where even the rhizomes die.
Chemical applications of herbicides such as glyphosate for several years may be effective.
No biological controls are known.

Selected References
IUCN SSC Invasive Species Specialist Group (ISSG). “Sorghum halepense (Grass).” ISSG Global Invasive
Species Database, 2005. http://www.issg.org/database/species/ecology.asp?si=213&fr=1&sts=sss.
“Johnsongrass.” Toxic Plants of Texas. Texas AgriLife Extension Service, n.d. http://essmextension
.tamu.edu/plants/toxics/detail.aspx?plantID=79.
Newman, Dara. “Element Stewardship Abstract, Sorghum halepense.” Global Invasive Species Team,
Nature Conservancy, 1993; updated 2009. http://wiki.bugwood.org/Sorghum_halepense.

n Jubata Grass
Also known as: Andes grass, Andean pampas grass, selloa, cortaderia,
pink pampas grass, purple pampas grass
Scientific name: Cortaderia jubata
Synonyms: Cortaderia atacamensis, Gynerium jubatum
 JUBATA GRASS AND PAMPAS GRASS n 473

n Pampas Grass
Also known as: Uruguayan pampas grass, silver pampas grass
Scientific name: Cortaderia selloana
Synonyms: Arundo selloana, C. argentea, C. dioica, Gynerium argentium
Family: Grass (Poaceae)
Native Range. Jubata grass is native to Northern Argentina and the Andes of Bolivia,
Chile, Peru, and Ecuador, from sea level to over 11,000 ft. (3,400 m) elevation. Pampas
grass is native to Argentina, Brazil, and Uruguay, in damp river margin soils.
Distribution in the United States. Jubata grass grows in coastal California, Oregon,
Washington and in Texas and
Hawai’i. Pampas grass is found
in the Pacific coast states of Cali-
fornia, Oregon, and Washington;
in the desert and mountain states
of Arizona, New Mexico, Utah,
and Colorado; and in the
southeastern states from Texas to
Virginia. Also in Hawai’i.
Description. Jubata grass and
its close relative pampas grass are
often confused. Both are large
tufted perennial grasses with
leaves rising from a tussock and
have a long feather-like, plumed
panicle extending above the
leaves. Leaves are glabrous on
both upper and lower surfaces,
and leaf margins are sharply ser-
rated. The two species, however,
are distinguishable by many
characteristics, including stem
height, details of leaves and flow-
ers, and method of reproduction.
Jubata Grass: The leaves of
jubata grass are more spreading
or drooping than erect, result-
ing in a shorter and broader
plant than pampas grass. The
leaf sheath of the bright-green
to deep-green leaves is densely
hairy, and the leaf tip is not
bristled or curled. Leaf blades,
Although a more aggressive invader than pampas grass, jubata grass is
which are 3–5 ft. (1–1.5 m) limited to frost-free areas in western states. (Native range adapted from
long and 0.8–4 in. (2–10 cm) USDA GRIN and selected references. Introduced range adapted from
wide, may be flat or slightly V- USDA PLANTS Database, Invasive Plant Atlas of the United States, and
shaped in cross-section. selected references.)
474 n GRAMINOIDS

Although roots are fibrous,

clustered around a crown not
far beneath the surface, they
may be either fine or thick.
Plants flower from July to
September. The inflorescence
stem typically rises 3–6 ft. (1–
2 m) above the tussock, and
the panicle itself is 1–3 ft.
(0.3–1 m) long. The flower
stalk is generally 2–4 times the
height of the tussock. Jubata
grass produces only female
flowers, which are initially deep
violet or purple, fading to pink-
ish or tan when mature. Spike-
lets are 0.6 in. (15 mm) long,
with 3–5 florets each. Florets
have hairs, 0.2–0.4 in. (6–
10 mm) long, on the base of
the lemma, and the short awns,
less than 0.04 in. (less than
1 mm), extend only slightly
beyond the hairs. The stigma
extends very little beyond the
awns, and the glumes are pur-
ple. The ripened seed is easily
separated from the rachilla.
Pampas Grass: Leaves of
pampas grass are more erect,
resulting in a taller plant,
Pampas grass is widely used as an ornamental and is still expanding its 6–13 ft. (2–4 m), not including
range. (Native range adapted from USDA GRIN and selected references. the inflorescences. Leaves, as
Introduced range adapted from USDA PLANTS Database, Invasive Plant long as 6 ft. (2 m) and 1–3 in.
Atlas of the United States, and selected references.) (3–8 cm) wide, are a bluish
green, with a tapering tip that is
bristly and curled. They are somewhat folded at the midrib, giving them a V-shape profile.
The sheath can be glabrous or variably hairy. Tussocks can be 9.8–16.4 ft. (3–5 m) in
diameter.
Roots of each established plant can fill about 1,100 sq. ft. (103 m2) of soil. Lateral roots
spread as far as 13 ft. (4 m) from the parent plant, and taproots can extend 11.5 ft. (3.5 m) deep.
Plants flower during August and September, with male and female flowers on separate
plants. Although flower stalks may be no higher than the height of the tussock leaves, they
are usually taller. Generally, the length of the flower stalk is equal to the height of the tus-
sock. The flower stalks in female plants may reach 3 ft. (1 m) high above the tussock, and
the male flower stems can be larger, as high as 6 ft. (2 m) above the tussock. The plume color
of female plants is light violet to silvery or creamy white, while the plumes of male plants are
a slightly darker violet. The inflorescence bears many spikelets, each 0.6 in. (15 mm) long.
 JUBATA GRASS AND PAMPAS GRASS n 475

A. The drooping nature of the stems of jubata grass results in a wide plant. (Joseph M. DiTomaso, University of
California–Davis, Bugwood.org.) B. Jubata grass leaf sheaths are hairy. (Joseph M. DiTomaso, University of
California–Davis, Bugwood.org.) C. Flowers on the panicle of jubata grass are all female. (Joseph M.
DiTomaso, University of California–Davis, Bugwood.org.) D. Culms of pampas grass grow more erect. (Forest
and Kim Starr.) E. Pampas grass leaf sheaths are usually glabrous. (Joseph M. DiTomaso, University of
California–Davis, Bugwood.org.) F. The female panicle (left) of pampas grass is larger and more showy than
the male inflorescence (right). (Joseph M. DiTomaso, University of California–Davis, Bugwood.org.)

Female plants have 5–7 florets per spikelet, which have dense hairs, 0.04–0.4 in. (1–10 mm)
long, at the base of the lemmas. The 3–5 florets on male plants have few to no hairs. Awns,
0.2–0.5 in. (5–12 mm), on female flowers are twice the length of the hairs. The stigma,
about twice the length of the awns, extends beyond them. Glumes on the spikelet are white.
Mature seed is not easily separated from the rachilla. Plants seldom produce seed because
most ornamental plants are female.
Related or Similar Species. Toe toe, also called Richard’s pampas grass, is native to New
Zealand and widely available on the Internet. A clump-forming perennial, it grows to 9 ft.
(2.7 m) tall and 6.5 ft. (2 m) wide. The arching leaves are stiff, leathery, and glabrous. The
inflorescence is a plume of creamy flowers that can be almost 10 ft. (3 m) tall.
Introduction History. Jubata grass was first used as an ornamental in France and Ireland,
and probably reached California in horticultural trade from France at an unknown date.
Although first recognized as a weed in California in the 1950s when it escaped cultivation,
it was not until 1966 that it was first reported as a problem in logged redwood forests of
northern California. Nurseries often sell jubata grass as “pampas grass.” The first confirmed
record of jubata grass in Hawai’i was in 1991.
476 n GRAMINOIDS

Pampas grass was taken as an ornamental to Santa Barbara, California, in 1848. By 1895,
Santa Barbara nurseries were primary producers that shipped plants worldwide. In 1946,
the U.S. Soil Conservation Service planted pampas grass in Ventura County and Los
Angeles County for dryland forage and erosion control. Selected for the showy female
plumes, plants were propagated vegetatively by division. More recently, plants have been
produced by seed, and because the sex of the plant cannot be determined until it blooms,
both male and female plants are currently sold as ornamentals. Because more males have
now been planted, pampas grass is producing more seed. Although pampas grass was intro-
duced to Hawai’i in 1925 as an ornamental, jubata grass was the only Cortaderia species
identified in Hawai’i until 1998.
Habitat. In California, jubata grass is found only in the coastal fog belt below 2,600 ft.
(800 m). It cannot tolerate frost, hot summers, or intense sun or drought. It prefers bare
or disturbed sites, such as roadcuts, forest clearcuts, landslides, cliffs, eroded soil, or burned
areas. Although it does best on sandy soils, it is capable of growing on a wide range of sub-
strate, including serpentine, with either low or high moisture. It grows in full sun to dense
shade, but cannot survive in redwood forest shade. It cannot colonize healthy native grass-
lands. On Mau’i, jubata grass grows from 2,000–7,000 ft. (660–2150 m) elevation, with as
little as 19.5 in. (500 mm) of annual rainfall.
Pampas grass is most commonly found below 1,000 ft. (330 m) elevation along the
California coast. Although tolerant of a wide variety of soils, it prefers deep sandy soils,
moist but with good drainage. It also grows in disturbed areas, roads and trails, and right-
of-ways. Although currently less extensive than jubata grass, it is still expanding its range.
Because pampas grass tolerates winter frost, warmer summers, more intense sun, and mod-
erate drought, it is used as an ornamental in the dry Central Valley of California, where it is
not invasive.
Reproduction and Dispersal. Jubata grass is apomictic, meaning that the female flowers
produce seeds without pollination. Year-old plants can flower, and some plants can flower
a second time in one season. Although it produces abundant viable seed, jubata grass has
no genetic diversity because no pollen transfer takes place. One inflorescence can produce
100,000 tiny seeds, and large clumps can produce a million or more. Wind-blown seeds
can be carried as far as 20 mi. (32 km). Plumes from mature plants are frequently used as
decoration, a human activity that spreads seeds. Seeds can also be dispersed by water and
machinery. No evidence exists of seed longevity or a soil seed bank. Germination occurs in
spring, ideally on sandy soil with plenty of moisture and light. Plants can germinate when
temperatures are 55–70ºF (13–21ºC). Seedlings grow best on bare soil, but need cool, foggy,
moist conditions. Survival is low under shady conditions or in competition with grasses.
Growth rate increases after young plants become established. New plants may also grow
from tiller pieces, sometimes caught in machinery, which root in moist soil.
Plants of pampas grass produce flowers 2–3 years after germination, and only produce
seed when male and female plants are found together. Seeds germinate in the spring under
the same requirements as jubata grass, and have a similar survival rate. Like jubata grass,
each inflorescence may have 100,000 seeds on each inflorescence. Tiny seeds are dispersed
primarily by wind, up to 20 mi. (32 km) distant.
Plants of both species live for about 15 years.
Impacts. The more aggressive of the two species, jubata grass threatens coastal ecosys-
tems of California, particularly sensitive dune areas, by displacing native vegetation. In cut-
over redwood forests in northern California, it has become a weed problem where its
presence interferes with establishment and growth of conifer seedlings. The buildup of dry
 JUBATA GRASS AND PAMPAS GRASS n 477

plant matter, leaves and flower stalks, is a fire hazard, and large tussocks can block access
and impede fire suppression efforts. The sawtoothed leaves cause injury to hikers, degrading
recreational experiences.
Pampas grass is a problem on the central and southern California coast, and may become
more so as more seeds are produced and disseminated. Because it cross-pollinates, it is more
genetically diverse than jubata grass, enabling it to adapt to a wider variety of environmental
conditions and possibly expand its range. Impacts of pampas grass are similar to those of
jubata grass, but not as severe.
Both jubata grass and pampas grass are currently limited to small stands on Mau’i. Bare
ground and cinder fields in Haleakala National Park, however, offer prime habitat. It is not
known if jubata grass can consistently withstand the freezing temperatures encountered
above 7,000 ft. (2,150 m) elevation, but seedlings and young plants are occasionally found,
indicating a potential for invasion of montane and subalpine regions. Infestations of jubata
grass have been located in pristine rainforest, the first known anywhere in the world.
These sites on open, sunny ridgetops receive as much as 197 in. (5,000 mm) of rainfall.
Management. Control or eradication is the same for both species. In order to preserve the
native ecosystems, options for control of either grass are limited in sensitive coastal
California sites or in Hawai’i rainforests and national parks. Seeding disturbed sites with
desirable vegetation may help to prevent Cortaderia seeds from germinating. Mulches, such
as layers of straw, on disturbed sites may also reduce germination.
Physical removal is only possible with small plants or in areas where little native vegeta-
tion remains to be saved. Small seedlings may be hand-pulled, while larger plants may
require a shovel or other tool. To prevent resprouting, the entire root crown must be
removed. Pulled plants left lying on the surface can root if the soil is moist. For large
plants, it may be necessary to cut or remove the leaves to expose the crown, which then
can be dug out. Inflorescences must be removed from the site or burned to destroy the
seeds, even if seeds are not yet mature. In severe, large-scale infestations, bulldozers and
backhoes may be necessary to remove plants. Prescribed burns are ineffective because
the growing points are protected by the deep thatch and leaves, and plants will resprout.
Fire, however, can be used to remove the foliage and expose the base for other treatments.
On very small scales, tarps placed over cut plants will shade out sprouts and also prevent
seeds from germinating.
Chemical treatment is most effective, although the amount of foliage requires a lot of con-
tact herbicide. It is more efficient to cut the foliage and spray the new sprouts. Spot treat-
ment with glyphosate, fluazifop, or imazapyr will kill the plant, although applications
must be repeated to control resprouting. Herbicides are most effective if applied before the
seed matures. An alternative to herbicides is to pour diesel oil on the root crown after the
leaves have been removed. However, this method leaves an oily residue in the soil.
No biological controls are known.

Selected References
Chimera, Charles, Forest Starr, Kim Martz, and Lloyd Loope. “Pampas grass (Cortaderia jubata and C.
selloana): An Alien Plant Report.” U.S. Geological Survey Biological Resources Division, American
Water Works Association Research Foundation, and Maui County Board of Water Supply, 1999.
http://www.hear.org/species/reports/corspp_fskm_awwa_report.pdf.
DiTomaso, Joseph. “Cortaderia jubata.” In Invasive Plants of California’s Wildlands, edited by Carla C.
Bossard, John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000.
478 n GRAMINOIDS

Almost Eradicated

A former residential property on Mau’i at 3,160 ft. (960 m) elevation supported

100 jubata grass clumps—landscape plants that got out of control. Tussocks
were 6.5 ft. (2 m) wide and interconnected, forming barriers with razor-sharp
leaves. Repeated physical control and herbicide applications over several years
reduced the population. When the efforts stopped due to financial and labor lim-
itations, however, the population rebounded.

http://www.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=33&survey
number=182.php.
DiTomaso, Joseph. “Cortaderia selloana.” In Invasive Plants of California’s Wildlands, edited by Carla C.
Bossard, John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000.
http://www.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=35&survey
number=182.php.
DiTomaso, Joseph M., Evelyn Healy, Carl E. Bell, Jenifer Drewitz, and Alison Stanton. “Pampasgrass
and Jubatagrass Threaten California Coastal Habitats.” Leaflet #99-1. Cooperative Extension,
Weed Research and Information Center, University of California, Davis. 1999. http://
wric.ucdavis.edu/PDFs/pampasgrass%20and%20jubatagrass%20WRIC%20leaflet%2099-1.pdf.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Cortaderia jubata (Grass).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?si=375&fr=1&sts=&lang=EN.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Cortaderia selloana (Grass).” ISSG Global Invasive Species Database, 2006. http://
www.issg.org/database/species/ecology.asp?fr=1&si=373.

n Kikuyugrass
Also known as: West African pennisetum
Scientific name: Pennisetum clandestinum
Synonyms: Pennisetum inclusum, P. longstylum, P. longstylum var. clandestinum
Family: Grass (Poaceae)
Native Range. Highlands of tropical East Africa, from Kenya to Tanzania and the eastern
Congo. Named for the Kikuyu tribe in Kenya.
Distribution in the United States. California and Arizona in the western states. Also in
Hawai’I and Puerto Rico.
Description. Kikuyugrass is usually a low-growing perennial grass, although some culms
may reach up to 1.6 ft. (0.5 m) tall. Several cultivars account for variation in size. Its pros-
trate stems root at the nodes, and the plant expands its coverage as a creeping, sod-
forming mat. Leaf blades are short, 0.5–2 in. (1.25–5 cm) and narrow. Leaf margins are
rough, from tiny serrations, and the tips can be either blunt or pointed. When leaf blades
first emerge, they are folded or rolled, subsequently opening to lie flat on the ground.
Leaves, either glabrous or covered with sparse soft hairs, have pronounced midveins on
the underside. Culms have overlapping leaf sheaths, and ligules are a fringe of white hairs.
Leaf sheaths, 0.4–4 in. (1–10 cm) long, are glabrous or slightly hairy.
 KIKUYUGRASS n 479

Rhizomes are branched and

thick, usually forming an exten-
sive network in the top 4 in.
(10 cm) of soil, but they can
reach 12–15.5 in. (30–40 cm)
deep. The multi-branched sto-
lons are coarse and flattened,
with many short internodes.
The nodes are swollen. Many
vertical leafy branches sprout
from the stolons and rhizomes.
Plants bloom from April to
October. Flowers, if present at
all, grow not on tall stems like
many other Pennisetum species,
but on leafy, vegetative side
shoots. The small, white, or
tawny-colored panicles, with
2–4 spikelets each, are incon-
spicuous, almost entirely hid-
den within the leaf sheaths,
with only the stamens barely
visible. The spikelets are 0.4–
0.8 in. (10–20 mm) long, gray-
green to straw colored with few
bristles. The dark-brown seeds
form inside the leaf sheaths.
Long, flexible bristles, attached
to the stem below each spikelet,
are dispersed with the spikelets
when seeds mature.
Related or Similar Species. Kikuyugrass grows best in moist sites and is especially invasive in coastal
Bermuda grass also grows close areas. (Native range adapted from USDA GRIN and selected references.
to the ground and propagates Introduced range adapted from USDA PLANTS Database, Invasive Plant
by rapidly growing stolons as Atlas of the United States, and selected references.)
long as 65.5 ft. (20 m). Leaves
are longer, 1–8 in. (2.5–20 cm)
and may be flat or folded. They have no distinct midvein but have rough edges. Flowering
stems can be as short as 6 in. (15 cm) or as tall as 3.3 ft. (1 m). Flower spikes, 1–4 in.
(2.5–10 cm) long, are prominent at the top of stems, where 2–12 are arranged in a star-
like shape. Spikelets are tiny and lack awns. Seeds can develop all year in warm climates,
but only in summer in cooler regions.
St. Augustine grass is used as a lawn grass in much of the moist tropics. Usually 6–12 in.
(15–30 cm) tall, it spreads by stolons and often forms a dense mat. Leaf blades, which are
folded when young, are up to 6 in. (15 cm) long, with a blunt apex. Inflorescences are short,
2–4.5 in. (5–12 cm) tall, with 1–3 spikelets.
Introduction History. Kikuyugrass was introduced to Hawai’i in 1925 as a forage crop. It
was planted in California in the 1930s as a ground cover to control erosion.
480 n GRAMINOIDS

A. Although it is usually low-growing, Kikuyugrass can be more than 1.5 ft. (0.5 m) tall. (Forest and Kim Starr,
U.S. Geological Survey, Bugwood.org.) B. Ligules are fringed with hairs. (Joseph M. DiTomaso, University of
California–Davis, Bugwood.org.) C. Plants form a thick sod. (Steve Dewey, Utah State University,
Bugwood.org.) D. Leaf blades are short. (Steve Dewey, Utah State University, Bugwood.org.) E. Creeping rhi-
zomes root at the nodes. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.)

Habitat. Kikuyugrass grows in a wide variety of habitats, including rangeland, grasslands,

orchards, disturbed sites, urban areas, wetlands, plantation crops, and forest edges.
Although it invades both dry and mesic habitats, the grass grows best under moist and
humid conditions, but it needs good drainage. It is especially invasive in coastal sites. It
grows in areas with mild winters and some summer moisture. It requires more than 35 in.
(900 mm) of annual rainfall, but established plants can also survive long periods of drought.
Because it does not tolerate dense shade, it is not found in healthy forests, but will invade
wet forests after a disturbance. In Hawai’i, it is often found on volcanic soils or red tropical
or subtropical soils, but tolerates most soil types. Found from 35º N to 37º S and up to
9,850 ft. (3,000 m) elevation, it can survive light frost but not sustained cold. It grows most
quickly at temperatures between 70ºF and 104ºF (21–40ºC), and slowly during winter,
becoming brown and dormant with frequent night frosts. Frequent defoliation by grazing
or mowing causes no negative effects.
Reproduction and Dispersal. Kikuyugrass reproduces vegetatively, spreading via an
extensive network of rhizomes and stolons. Pieces of rhizomes or stolons are easily trans-
ported long distances on machinery or on vehicles. Although rarely produced, seeds are dis-
persed by wind and are also found in cow or sheep droppings. The best soil temperatures for
germination are 68–86ºF (20–30ºC), and seedlings can emerge from soil depths up to 2.4
in. (6 cm).
Impacts. Kikuyugrass is an aggressive invader of pasture, crops, and natural ecosystems.
Its thick mats smother native vegetation, and its alleleopathic compounds prevent other
plants from growing. It also invades manicured turf, such as lawns and golf courses. It can
overrun agricultural areas and roadsides, and can climb over shrubs and small trees. Plants
can accumulate high levels of oxalates and nitrates, which are toxic to grazers, both wildlife
and livestock. The basal layer of dead leaves is a ground fire hazard.
In spite of its invasive tendencies, kikuyugrass has widespread use in the world as pasture
for dairy and beef cattle. It is also used for soil stabilization and erosion control and as a lawn
grass.
Management. Prevention of accidental spread of kikuyugrass requires that all agricultural
equipment be thoroughly cleaned of all plant pieces. Although small plants can be removed
 MEDUSAHEAD n 481

Kikuyugrass in Peru

T his African grass is sometimes used as a lawn grass on the terraces of Incan ruins,
such as Machu picchu. Stolons from the intentional plantings, however, have
overrun some ancient ruins, sprouting in cracks and breaking stones, causing
irreparable damage to priceless archaeological structures.

by hand when first identified, physical control is limited because it is almost impossible to
dig out all the rhizomes and stolons. Removing rhizomes at the edges of stands, however,
will prevent its spread in the immediate area. Disking is discouraged because it will break
and spread rhizomes and stolons. Covering the infested area with a plastic sheet for
4–12 weeks may be effective in killing the plants.
Chemical control can be achieved with the use of systemics, such as glyphosate. Mowing
before application reduces the standing biomass and amount of herbicide required.
Potential for biological control includes fungus diseases and insects. The rust fungus
(Phakopsora apoda), now established in South Africa, reduces the plant’s ability to photosyn-
thesize but does not kill the plant. A fungus disease caused by Pyricularia grisea kills seed-
lings. Two insects (Sphenophorus entus vestitus and Herpetogramma licarsicalis) damage
plants in Hawai’i. Plants are also susceptible to sugarcane aphids (Sipha spp.).

Selected References
“Bermuda Grass.” Blue Planet Biomes, 2002. http://www.blueplanetbiomes.org/bermuda_grass.htm.
Fukumoto, Glen K., and Chin N. Lee. “Kikuyugrass for Forage.” Livestock Management, LM-5.
Cooperative Extension Service, College of Tropical Agriculture and Human Resources, University
of Hawai’i at Manoa, 2003. http://www.ctahr.hawaii.edu/oc/freepubs/pdf/LM-5.pdf.
“Kikuyugrass (Pennisetum clandestinum).” Plant Pest and Health Prevention Services (PHHPS),
California Department of Food and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/
pennisetum.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Pennisetum clandestinum (Grass).” ISSG Global Invasive Species Database. 2005. http://
www.issg.org/database/species/ecology.asp?si=183.
“Stenotaphrum secundatum (St. Augustine Grass).” ZipcodeZoo, n.d. http://zipcodezoo.com/Plants/S/
Stenotaphrum_secundatum/.

n Medusahead
Also known as: Medusahead wildrye
Scientific name: Taeniatherum caput-medusae
Synonyms: Taeniatherum asperum, T. crinitum, Elymus caput-medusae, Hordeum caput-
medusae
Family: Grass (Poaceae)
Native Range. Western part of the Mediterranean Basin, southern Europe and northern
Africa, from Portugal and Morocco east to Turkey and Syria. Possibly native to Southwest
Asia as far east as Pakistan.
482 n GRAMINOIDS

Distribution in the United

States. West Coast, from
California north to Washington,
east to Idaho, Montana, Neva-
da, Utah, and Arizona. Also in
Nebraska and northeastern
states of New York, Connec-
ticut, and Pennsylvania.
Description. Medusahead is
a winter annual grass with slen-
der stems and leaves. The
stems, which branch from the
base of the plant, can be either
decumbent or erect and reach
8–24 in. (20–60 cm) tall. Each
stem has 2–4 short and narrow
leaf blades, 1.2–2.5 in. (3–6 cm)
long and less than 0.2 in. (5 mm)
wide. Leaf margins are some-
what in-rolled and sometimes
fringed with hairs. Leaf sheaths
are slightly inflated. Both sheaths
and leaves may be either gla-
brous or covered with a downy
pubescence of minute hairs.
The fibrous roots continue to
grow during the cool season,
using the upper soil moisture first.
Later in the season, they draw
moisture from deeper soil layers.
If the primary root becomes des-
After accidental introduction into Oregon, medusahead spread iccated, seedlings will survive
throughout dry western rangeland and shrub communities. (Native because they are able to grow
range adapted from USDA GRIN and selected references. Introduced new roots when moisture again
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the becomes available. Plants form a
United States, and selected references.) dense mat of stems and roots
1.5–5.0 in. (4–12.5 cm) thick.
The inflorescence, which develops in June, is a dense, bristly spike, 0.6–2 in. (1.5–5 cm)
long, not including the awns. Spikelets occur in pairs, with two florets each, only one of
which is fertile. The lemma of the fertile floret has three veins. Long awns, 1–3 in. (2.5–
7.5 cm), extend from the tips of the lemmas. They are straight or slightly curved when green,
but twist and spread erratically as they dry, giving the grass the appearance of the mythical
Medusa. Glumes are also awn-like, 0.4–1.6 in. (1–4 cm) long. Both awns and glumes are flat
and stiff, with tiny pointed barbs along the margins. When seeds are ripe, spikelets separate
from the axis above the glumes, while the axis stays intact. Old flower spikes with the awn-
like glumes remain on the plant.
Because medusahead matures 2–4 weeks later than perennial native grasses, it is conspic-
uously yellowish green in pastures of dried native plants.
 MEDUSAHEAD n 483

A. Medusahead stems branch from the base of the plant. B. When green, the long awns are straight. C. Awns
become curved or twisted as they dry. D. Seeds have long awns. (Steve Dewey, Utah State University,
Bugwood.org.)

Related or Similar Species. Barley and wildrye are distinguished by having three florets
per spikelet and five veins on the lemmas. The flower spike axis breaks apart when seed is
mature. Arizona wheatgrass is a closely related California native. Because of the long awns,
medusahead may also be confused with foxtail, but the flower spike on that grass also disar-
ticulates when mature.
Introduction History. Medusahead was introduced to the United States in the late 1880s,
possibly as a seed contaminant. The first collected specimen was near Roseburg, Oregon, in
1887. The plant spread rapidly in the 1930s.
Habitat. Medusahead is found in disturbed pasture, sagebrush, chaparral, oak wood-
lands, agricultural fields, and along dry roadsides. It threatens sparsely populated commun-
ities of rangeland, especially those originally dominated by big sagebrush, bluebunch
wheatgrass, and Sandberg bluegrass. More complex communities that have been degraded
due to overgrazing, fire, or cultivation are also susceptible to medusahead. The grass favors
soils with well-developed profiles that have some type of clay horizon. It is likely that a clay
layer is necessary to retain water longer, allowing medusahead to mature later in the
summer. The species is also found in swales that accumulate runoff. Unlike most perennial
grasses, medusahead is adapted to vertisols, which are dry soils that expand and contract
when moistened and dried. It is usually not found on well-drained, coarse-textured sub-
strate, or on soils with little profile development. Those areas remain dominated by cheat-
grass (see Graminoids, Cheatgrass).
Medusahead grows in areas with mild to cold winters and hot summers, but not in places
that experience long cold periods. It is found where annual precipitation, falling primarily in
autumn, winter, and spring, is 10–40 in. (250–1,000 mm), but most infestations are in
sagebrush-bunchgrass communities that receive 10–20 inches (250–500 mm) annually.
Reproduction and Dispersal. Flowers are self-pollinated, and reproduction is solely by
seed, with 8–15 on each flower spike. Seeds are dispersed locally by wind and water, and
carried in coats of livestock, especially sheep, for long distances. New infestations are com-
monly seen along livestock trails. Seeds remain viable for at least one year. Most seeds germi-
nate in the fall after the first rain, under optimal temperatures of 50–59ºF (10–15ºC), but
some germination also takes place in winter and spring. Germination rate is as high as
98 percent, with seeds sprouting within 8–10 hours of being moistened. They do not need
484 n GRAMINOIDS

direct contact with soil and germinate best in dense litter with low moisture. They may also
emerge from a depth of approximately 3 in. (8 cm). Seedlings grow all winter, and roots
grow 7–8 in. (18–20 cm) deep before the plant branches. Roots can be 40 in. (100 cm) deep
by February. Plant density can be 100–200 per sq. ft. (1,000–2,000 per m2).
Impacts. Medusahead alters the natural plant community and adversely affects the live-
stock industry. It displaces native grasses and degrades rangeland. The plant changes the
shrub and perennial grass community into an annual grass community with less plant diver-
sity for wildlife. Medusahead germinates sooner than bluebunch grass, and plants take water
from the same soil depth. In some areas of southwestern Idaho, the grazing capacity of ran-
gelands has been reduced by as much as 80 percent. Because plants contain a lot of silica
and are not palatable, medusahead is eaten by livestock only in its early, green stage of
growth or when nothing else is available. When dry, the long, sharp awns may injure ani-
mals’ mouths and eyes. At any stage of growth, it has little feed value and is not important
as wildlife forage, although rabbits may graze the leaf blades. Seeds are indigestible to game-
birds, even to the introduced chukar, which eats cheatgrass seeds.
In California and Oregon, where the range and habitat of medusahead overlaps with two
introduced bromes, soft brome and cheatgrass, medusahead is displacing cheatgrass on
mesic sites. Seeds germinate sooner and faster than cheatgrass and bluebunch wheatgrass.
The roots of medusahead seedlings grow faster than those of cheatgrass, tapping soil mois-
ture first, but plants mature 2–3 weeks later than cheatgrass.
The thick mat of stems decomposes slowly, tying up nutrients and changing the temper-
ature and moisture regimes in the soil. Although the mat reduces germination of other
grasses, such as cheatgrass, it favors germination of medusahead seeds.
The thick stem mats increase the fuel load and pose a fire hazard.
Management. The best management is prevention, by maintaining healthy native peren-
nial vegetation. Eradicating other invasive plants, especially cheatgrass, can open the ground
to medusahead. Most attempts at restoration by reseeding with perennial grasses, such as
bluebunch wheatgrass, will not be successful unless medusagrass is first eradicated from
the site. A possible first step to reestablishment of native sagebrush-wheatgrass-bluegrass
communities, however, is to overseed with squirreltail, a native perennial bunchgrass,
which may outcompete medusahead. The self-pollinating squirreltail germinates and
matures quickly. It has better root reserves to compete with annuals, is fire tolerant, and
has good seed dispersal. Depending on the area, other competitive perennial grasses include
intermediate wheatgrass, Thurber’s needlegrass, needle and thread grass, Indian ricegrass,
Sandberg bluegrass, and sheep fescue.
Physical control targets seed production. Controlled burns conducted in early June,
after native grasses have matured but before medusahead seeds set, can almost eliminate
medusagrass for several years. A slow, hot burn kills developing seeds, and fires also
serve to reduce thatch. Disking or plowing before seed set also reduces seed production.
Tillage breaks the thatch and buries seeds too deeply to sprout but also destroys all other
plants and alters the soil moisture regime by removing microbiotic crusts. Mowing is also
nonselective and fails to crop plants close enough to prevent resprouting. Any mowing after
seed set, especially along roadsides, spreads the seeds. Grazing by sheep in the spring when
the plant is still in the green stage can control seed production, but animals must be moved
before seeds develop to avoid transportation of seeds.
Chemical control is limited. Applications of glyphosate and paraquat have provided var-
iable results. Because they are nonselective, they cannot be used where medusahead is
mixed with desirable species. Many herbicides used on rangeland target broadleaf weeds
 QUACKGRASS n 485

Does Grazing Cause Medusahead Invasions?

I n an attempt to assess the impact that grazing has on medusahead invasions, an

area of rangeland in Modoc County, northern California, was fenced off from
grazing in 1967. The enclosure, which did not have medusahead, was susceptible
to invasion because the plant occurred in the surrounding area. Thirty years later,
it was discovered that medusahead had successfully invaded and was the dominant
herb in both areas. Shrub cover almost doubled in the grazed areas, which logically
had fewer perennial grasses, and the non-grazed enclosure had more perennial
grasses. However, the only sites in the grazed areas that were totally converted to
medusahead were deep vertisol clay soils. A major conclusion was that ecological
relations are more complicated and that grazing pressure, or lack of it, is not the
only factor than influences medusahead invasion.
Source: Wagner, Delmas, and Young, 2001.

and may encourage the growth of annual grasses like medusahead by removing competition.
Fall application of atrazine, which targets annual grasses, controls medusahead but may also
harm native perennial grasses, such as bluebunch wheatgrass and Columbia needlegrass.
Little research has been conducted on biological control. Five soil fungi native to the
western United States have been considered, but they are not host-specific and present a
danger to winter wheat crops and crested wheatgrass. The near future holds no possibilities.

Selected References
Archer, Amy J. “Taeniatherum caput-medusae.” In Fire Effects Information System. U.S. Department of
Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, 2001.
http://www.fs.fed.us/database/feis/plants/graminoid/taecap/all.html.
Maurer, Teresa, Mary J. Russo, and Audrey Godell. “Element Stewardship Abstract, Taeniatherum
caput-medusae.” Global Invasive Species Team, Nature Conservancy, 1988; modified 2009. http://
wiki.bugwood.org/Taeniatherum_caput-medusae.
“Medusahead (Taeniatherum caput-medusae).” Plant Pest and Health Prevention Services (PHHPS),
California Department of Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/
taeniatherum-caput-medusae.htm.
Wagner, Joseph A., Richard E. Delmas, and James A. Young. “Thirty Years of Medusahead: Return to
Fly Blown-Flat.” Rangelands 23(3): 6–9, 2001. http://uvalde.tamu.edu/rangel/jun01/wagner.pdf.

n Quackgrass
Also known as: Couch grass, dog grass, twitchgrass, creeping wild rye, witchgrass,
devils-grass, wiregrass, and others
Scientific name: Elymus repens
Synonyms: Agropyron repens, Elytrigia repens, E. vaillantiana, Triticum repens,
T. vaillantianum, and others
Family: Grass (Poaceae)
Native Range. Europe, Asia, and Mediterranean North Africa, from the British Isles east
to northern India and western China.
486 n GRAMINOIDS

Distribution in the United

States. Throughout the country
except the extreme south-
eastern states. In Alaska, but
not in Hawai’i.
Description. Quackgrass is a
cool-season perennial with
many regional genotypes that
give it variable morphology.
Erect stems are usually 1–4 ft.
(0.3–1.2 m) tall, but decum-
bent stems, hugging the ground
only 0.2–0.8 in. (0.5–2.0 cm)
above the soil surface, are more
common. The glaucous culms
are green to whitish. Leaf
blades, 1.5–12 in. (4–30 cm)
long and less than 0.5 in.
(1.3 cm) wide, are lax and usu-
ally droop from the stem.
Although becoming flat as they
mature, the leaf blades are
inrolled when in bud. Leaves
are pointed, with a slight con-
striction at the tip. Small
auricles, ear-shaped appen-
dages that occur at the junction
of the leaf blade and sheath,
are whitish to violet-tinged and
clasp the stem. The upper sur-
face of leaf blades is sparsely
Quackgrass is found in a wide variety of habitats, but grows best in the hairy, but the lower surface is
northern states, with cool climates with long summer days. (Native glabrous. The open sheaths may
range adapted from USDA GRIN and selected references. Introduced be either glabrous or softly pubes-
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the cent. Hairiness of leaf blades and
United States, and selected references.) sheaths is extremely variable,
and hairs are usually more notice-
able on new spring growth. Ligules are membranous, sometimes with a small fringe.
The root system consists of long, somewhat fleshy, slender rhizomes, approximately 0.1
in. (3 mm) in diameter. The many branches of the rhizomes, 4–8 in. (10–20 cm) below
the surface, create a tough mat-like network and a loose sod. Rhizomes are pale yellow or
straw-colored, with a tough brown sheath at each node that makes the rhizomes appear
scaly. Fibrous roots extend from each node. The sharply pointed tips of the rhizomes are
able to penetrate hard soils.
Quackgrass flowers from May to September. The inflorescence is an erect spike, 2–8 in.
(5–20 cm) long, with spikelets in two long rows, alternate on the rachis. Each node on the
spike has a single, sessile spikelet, with the spikelets overlapping and flattened toward the
stem. Each spikelet has 3–8 florets, each approximately 0.4 in. (1 cm) long. Both the glumes
 QUACKGRASS n 487

A. Stems may be 1–4 ft. (0.3–1.2 m) tall. (Steve Dewey, Utah State University, Bugwood.org.) B. Stems are
glaucous and smooth. (Ohio State Weed Lab Archive, The Ohio State University, Bugwood.org.) C. Auricles
clasp the stem where the leaf blade joins the sheath. (Ohio State Weed Lab Archive, The Ohio State University,
Bugwood.org.) D. Seeds develop on a narrow spike. (Ohio State Weed Lab Archive, The Ohio State University,
Bugwood.org.) E. New plants sprout from rhizome nodes. (Steve Dewey, Utah State University, Bugwood.org.)

and the lemmas have either short awns or none. The pale, yellow to brown seeds, which
mature from July to September, are elliptical, less than 0.4 in. (1 cm) long.
Related or Similar Species. Quackgrass is distinguished from several similar grasses by its
solitary spikelets and extensive scaly rhizome system. Blades of rush wheatgrass, hairy wheat-
grass, and Russian wheatgrass are stiff and strongly ribbed. Tall wheatgrass and ryegrasses
have no rhizomes. Beardless wheatgrass, tall wheatgrass, and crested wheatgrass are all tuft-
forming grasses without rhizomes. Northern wheatgrass has longer internodes on the spikes.
Western wheatgrass leaf blades are strongly ridged and furrowed. Tick quackgrass has pithy
culms at anthesis (when anthers and ovules are mature), while quackgrass culms are hollow.
Introduction History. First reported in North American in 1672, quackgrass probably
arrived as a contaminant in hay or straw. It has since been crossed with other grass species
in the development of many hybrids planted for livestock consumption. Its mat of rhizomes
makes the plant useful for stabilization of sandy areas or steep slopes, and it can be used for
revegetation of mine tailings.
Habitat. Quackgrass, found in several types of low-lying moist areas of grassland com-
munities, can colonize both disturbed sites and undisturbed areas. As an early successional
plant, it is common in waste areas, along ditches, stream banks, and roadsides, and in culti-
vated and abandoned fields. It also invades pastures, mixed-grass prairies, open woodlands,
and mountain meadows, as high as 9,000 ft. (2,750 m) elevation in Rocky Mountain
National Park. Although the grass does best in neutral to slightly alkaline soils (pH
6.5–8.0) that are fertile and loamy, it can grow in a variety of soils, including saline. It does
not grow on very acid soils or on rocky outcrops. Quackgrass requires only 90 frost-free
days, and grows best when temperatures are 68–77ºF (20–25ºC). Rhizomes grow best in
cool temperatures, 50ºF (10ºC), and long, 18-hour days. Rhizomes can survive tempera-
tures as low as 1.4ºF (-17ºC), but surface shoots are killed by frost. Quackgrass is drought
tolerant, but does not grow in shade.
Reproduction and Dispersal. Quackgrass reproduces both by seed and vegetatively.
Although each flower stem may produce as many as 400 seeds, 25–40 is typical. Plants are
able to flower and set seed more than once in a season. The wind-pollinated plants require
cross-fertilization. Although seeds have no special adaptations for dispersal and most fall
488 n GRAMINOIDS

near the parent plant, they are frequently transported in crop seed, straw, or manure. Seeds
in the soil remain viable for 2–4 years, and seeds can germinate after passing through an
animal’s digestive tract. Seeds germinate in both fall and spring, ideally when diurnal
temperatures range from 59º to 77ºF (15–25ºC), and can emerge from a depth of
4 in. (10 cm).
Rhizomes begin growth in April and May and grow rapidly in both spring and fall.
When the plant sends up new shoots in the fall, as far as 2 ft. (0.6 m) distant from the
parent plant, the rhizome also continues to grow horizontally. One plant may have as many
as 1,200 dormant buds on its rhizomes, and fragments will grow new plants. Growth of
rhizome buds may be stimulated by disturbance. As individual clumps increase in size from
rhizome extension each year, they eventually merge to become a continuous stand of
quackgrass.
Impacts. Quackgrass is an aggressive invader that can grow over 200 new shoots and
extend its spread by as much as 10 ft. (3 m) a year. As a cool-season grass, quackgrass
begins to grow in early spring, enabling it to outcompete species that begin growth later.
By utilizing soil moisture and nutrients, the rhizomes of quackgrass compete with culti-
vated crops as well as with native grasses and forbs in grassland communities. Because
quackgrass dominates disturbed land for several years after invasion, it can alter natural
succession, and hinder restoration of damaged areas. It can prevent native woody species
from regenerating. Quackgrass infestations increase dramatically after fires, also altering
succession. Alleleopathic toxins from the shoots and roots contribute to the ability of
plants to suppress native species. In direct contrast to decreasing the biodiversity in an eco-
system, quackgrass also provides cover in grasslands for small rodents, birds, and
waterfowl.
One of the most difficult invaders to control in cultivated fields, quackgrass can signifi-
cantly reduce crop yields or reduce productivity of pasture and rangeland. Seeds that con-
taminate grain crops lower the value of the harvest.
Management. Physical control in agricultural fields can be accomplished by tillage and
crop rotation, in conjunction with herbicide applications. Although a single effort can
increase the infestation by spreading rhizome fragments, two years of intensive cultivation,
repeated each time new growth appears, depletes the rhizome reserves. Fields may also be
mowed or intensely grazed before tilling, a method that also will reduce reserves. Early
spring burns in mixed grasslands, done repeatedly, or burning twice a year have been
successful.
Chemical control has had mixed results due to various timing of application and lack of
repeated attempts. One effective method is application of a postemergent selective to grasses,
such as fluazifop, sethoxydim, or clethodim. Infestations in natural areas are more difficult
to control with herbicide sprays because of potential damage to nontarget plants. Spot appli-
cations or application while native species are dormant is possible. Glyphosate and nicosul-
furon are nonselective postemergents. Herbicides should be applied in early spring or fall
when quackgrass is 6–8 in. (15–20 cm) tall and actively growing.
No biological control is known.

Selected References
“Agropyron repens.” Species Abstracts of Highly Disruptive Exotic Plants at Pipestone National
Monument. Northern Prairie Wildlife Research Center. U.S. Geological Survey, 2006. http://
www.npwrc.usgs.gov/resource/plants/exoticab/pipeagro.htm.
 WEST INDIAN MARSH GRASS n 489

Batcher, Michael S. “Element Stewardship Abstract, Elytrigia repens.” Edited by Mandy Tu and Barry
Meyers-Rice. Global Invasive Species Team, Nature Conservancy, 2001. http://www.invasive.org/
weedcd/pdfs/tncweeds/elytrep.pdf.
“Quackgrass.” Non-Native Plant Species of Alaska. Alaska Natural Heritage Program. Environment and
Natural Resources Institute. University of Alaska, Anchorage, 2006. http://akweeds.uaa.alaska.edu/
pdfs/species_bios_pdfs/Species_bios_ELRE_ed.pdf.
“Quackgrass (Elytrigia repens).” Plant Pest and Health Prevention Services (PHHPS), California
Department of Food and Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/elytrigia
-repens.htm.

n West Indian Marsh Grass

Also known as: Water straw grass, trumpet grass, trompetilla
Scientific name: Hymenachne amplexicaulis
Synonyms: Hymenachne acutigluma, Panicum amplexicaule
Family: Grass (Poaceae)
Native Range. Tropical Central and South America, from Mexico south to Uruguay, and
the West Indies.
Distribution in the United States. South of 29º N latitude in central and coastal Florida.
Also in Puerto Rico. Not yet reported elsewhere.
Description. West Indian marsh grass is a perennial grass, with robust erect culms, or
stems, 3.3–8.2 ft. (1–2.5 m) tall and less than 0.5 in. (1.2 cm) in diameter. Stems are gla-
brous and have few branches. Stems are not hollow, but filled with white pith, called aeren-
chyma. The large intercellular spaces in the aerenchyma allow oxygen to easily move from
the leaves to the roots. The glossy green leaf blades, 4–18 in. (10–45 cm) long and approx-
imately 1.5 in. (4 cm) wide, are flat. Although both the leaf sheaths and blades are generally
glabrous, hairs may line the upper margins of the sheaths and the lower margins of the
leaves. The leaf sheaths are often spongy. The leaf blade is heart-shaped at the base, with ear-
like appendages called auricles that clasp the stem. The ligule is membranous.
Prostrate stems, or cord-like stolons, extend many feet away from the parent plant. At
nodes along the stolons, roots grow to anchor the plant in the substrate.
Plants flower September to December, usually with one inflorescence per plant, but
sometimes three in succession. The inflorescence is a narrow, spike-like, cylindrical panicle,
4–20 in. (10–50 cm) long and 0.4 in. (1 cm) wide. The short-stalked spikelets, oriented
upright on the spike are lance-shaped, compressed, and less than 0.2 in. (5 mm) long,
Related or Similar Species. American cupscale, a grass native to the United States, has a
similar inflorescence but lacks the white pith in the culms and earlike appendages at the leaf
bases.
Introduction History. West Indian marsh grass is promoted as forage in seasonally
flooded land in the Tropics. Although stems are too coarse to be dried for hay, it can be
cut for green feed or for silage. It also can be used as a nutrient sink or sediment trap in pol-
luted areas. It was first described in Florida in 1908, as a rare plant. The first herbarium
record is 1957, from a pasture. It may have been intentionally introduced as forage or natu-
rally carried from the Caribbean islands by migrating birds. The second specimen, in 1977,
was also from a pasture.
Habitat. West Indian marsh grass grows in freshwater wetlands, such as floodplains,
swamp margins, and river banks. Adapted to fluctuating water levels, it prefers fertile areas
490 n GRAMINOIDS

that are seasonally inundated. It

typically grows in water less
than 6.5 ft. (2 m) deep, but is
sometimes found in water twice
that depth. Extensive colonies
are especially abundant in sites
that are enriched by nutrient
runoff, including shallow
ponds, cypress swamps, and
wet disturbed areas such as pas-
tures, ditches, and drainage
canals. The grass needs sunlight
and will tolerate only light
shade. It is not tolerant of
drought or salt, even an occa-
sional tidal impact. Based on
its native range and elevation,
it may have some adaptations
to frost.
Reproduction and Dispersal.
West Indian marsh grass repro-
duces both by seed and vegeta-
tively. It is a short-day plant,
needing about 12 hours to
flower, which may limit its
northward spread. Flowering is
also stimulated by rainfall,
which occurs in autumn in
southern Florida. Plants pro-
duce a high number of seeds,
which are viable for 1–3 years.
West Indian marsh grass is a short-day plant that grows in freshwater envi- Seeds are dispersed by stream
ronments in Florida. (Native range adapted from USDA GRIN and selected flow, floodwater, and water
references. Introduced range adapted from USDA PLANTS Database, birds. Seeds carried by migrat-
Invasive Plant Atlas of the United States, and selected references.) ing birds may account for pop-
ulations on islands.
Nodes on the submerged stolons produce roots and new plants. Pieces of the stolons or
entire rafts of plants broken loose from the banks may be carried long distances down-
stream. New infestations often follow roadsides before spreading into adjacent wetlands
because muddy tires of vehicles frequently transport seeds and stolon fragments.
Deliberate movement of plants for pasture development also may spread West Indian marsh
grass. If plants reach reservoirs, the species can easily follow canals to increase its range.
Impacts. Information on the use of West Indian marsh grass as a forage crop indicates
that it tends to suppress other species in the habitat, displacing native plants. Everglades
National Park, Big Cypress Swamp, and the St. Johns River are at risk. It creates monotypic
stands with dense mats both above and below the water, blocking light and developing a
large amount of biomass in marsh lands. The grass may alter biodiversity, resulting in fewer
 WEST INDIAN MARSH GRASS n 491

A. Dense stands of tall West Indian marsh grass can dominate freshwater wetlands. (Chris Gardiner, School of
Veterinary and Biomedical Sciences, James Cook University, Queensland, Australia.) B. The base of the leaf blade
clasps the stem. (Biosecurity Queensland, Department of Employment, Economic Development and Innovation,
Queensland Government, Australia.) C. The inflorescence is a narrow spike. (Chris Gardiner, School of
Veterinary and Biomedical Sciences, James Cook University, Queensland, Australia.) D. The lower part of the
leaves and upper part of the leaf sheaths are lined with hairs. (Biosecurity Queensland, Department of
Employment, Economic Development and Innovation, Queensland Government, Australia.) E. Roots form at
nodes along the stolons. (Biosecurity Queensland, Department of Employment, Economic Development and
Innovation, Queensland Government, Australia.)

species of native plants and animals. It threatens the survival of riparian forests, where it
dominates river banks and prevents normal plant succession.
The grass invades habitats needed by several types of native species, including birds, rep-
tiles, amphibians, and fish, reducing the resources available for feeding, shelter, and breed-
ing. Two grasses native to the eastern United States, maidencane and coast cockspur, that
provide refuge for wildlife are being displaced by West Indian marsh grass. Habitats for both
American alligators and American crocodiles are threatened, as are habitats of wading birds,
such as Wood Storks, and raptors, such as the Everglade Snail Kite. By depleting the water of
oxygen, stands may kill fish species, including killifishes, live bearers, and juvenile sun-
fishes. Several native frogs, such as leopard frogs, pig frogs, and green tree frogs are also at
risk as their habitat diminishes.
Also considered a weed elsewhere in the Tropics, West Indian marsh grass is becoming
increasingly difficult to control along drainage canals in Florida. Its dense stands alter water
quality and may worsen flooding by changing flow regimes and blocking drainage ways.
Management. Eradication of West Indies marsh grass is difficult because it both grows
and reproduces quickly. The best management is prevention. Because the plant is a prolific
producer of seeds that can be carried long distances, sites should be monitored at least twice
a year after any attempt at control.
Physical methods must remove the stolons to prevent vegetative regrowth. Mechanical
harvesters can be used on small areas as long as root fragments are removed. Because the
grass is intolerant of shade, solarization, covering the area with plastic or mulch, may be
practical for small areas. Manipulation of water level, either drying out or drowning plants,
may facilitate elimination of the grass.
492 n GRAMINOIDS

Chemical applications of herbicides can kill buried stolons. Both imazapyr and glypho-
sate are effective, killing 50–90 percent of plants. Repeated applications are necessary
because some plants can recover and produce both stolons and seeds within three months.
Burning the stubble after first spraying with glyphosate works well. Treatments should be
done prior to flowering to prevent seed development and dispersal.
An insect unofficially referred to as the Myakka bug, Ischnodemus variegatus, has potential
for biological control. Although native to South America, it was discovered feeding on West
Indian marsh grass in Florida in 2000 and now occurs throughout the central and southern
parts of the state. The insect can cause severe seasonal damage to West Indian marsh grass
by reducing its ability to photosynthesize and slowing its growth rate. It may be host-
specific. Although the grass is susceptible to two fungal diseases, caused by Curvulara lunata
and Phyllachora spp., damage is insufficient.

Selected References
Diaz, Rodrigo, and Bill Overholt. “West Indian Marsh Grass.” Weeds Gone Wild: Alien Plant Invaders
of Natural Areas, Plant Conservation Alliance’s Alien Plant Working Group, 2006. http://
www.nps.gov/plants/alien/fact/hyam1.htm.
Diaz, Rodrigo, William A. Overholt, Brent Sellers, and James P. Cuda. “Wetland Weeds: West Indian
Marsh Grass (Hymenachne amplexicaulis).” Document ENY-693 (IN491), Institute of Food and
Agricultural Sciences (IFAS), University of Florida, 2003; revised 2010.http://edis.ifas.ufl.edu/
in491.
“Factsheet—Hymenachne amplexicaulis.” Tropical Forages, n.d. http://www.tropicalforages.info/key/
Forages/Media/Html/Hymenachne_amplexicaulis.htm.
Langeland, K. A., H. M. Cherry, C. M. McCormick, and K. A. Craddock Burks, eds. “Hymenachne
amplexicaulis (rudge) Nees.” In Identification and Biology of Non-Native Plants in Florida’s
Natural Areas, 2nd ed. UF/IFAS Publication #SP 257. Institute of Food and Agricultural Sciences
(IFAS), University of Florida, 2008. http://www.fleppc.org/ID_book/Hymenachne%20
amplexicaulis.pdf.
n Shrubs
n Asiatic Colubrina
Also known as: Latherleaf, common colubrina, Asian snakeroot, Indian snakewood,
wild coffee, Asian nakedwood
Scientific name: Colubrina asiatica
Synonyms: Ceanothus asiaticus, C. capsularis, Celastrus sepiarius, Rhamnus asiatica,
R. splendens
Family: Buckthorn (Rhamnaceae)
Native Range. Coastal East Africa, Madagascar, and the Sechelles; eastern India through
Southeast Asia to New Guinea, Solomon Islands, New Caledonia, and northeastern
Australia.
Distribution in the United States. The frost-free region of the east and west coasts of cen-
tral and southern Florida, including the Florida Keys and the Everglades, where it has
become naturalized. Widespread in the Caribbean and Puerto Rico and probably in the
Virgin Islands. Also in Hawai’i.
Description. Asiatic colubrina is an evergreen shrub with either drooping or climbing,
almost vine-like, reddish-brown slender branches that can be 20 ft. (6 m) or more long.
Branchlets are slender and may slightly zigzag between the nodes. Stems on older plants
can reach 50 ft. (15 m) and be as thick as 4 in. (10 cm) in diameter. The wood is reddish-
orange and the bark is dark brown, sometimes with white striations.
The thin, papery leaves are oval or egg-shaped, 1.5–5.5 in. (4–14 cm) long and 1–2 in.
(2.5–5 cm) wide, with 3–5 distinct raised veins extending from the leaf base. Leaves are
alternate on the stem and supported by a short, slender 0.6 in. (1.5 cm) petiole. They are
glabrous, shiny dark green on the upper surface and a dull, paler green below. The apex
of the leaf is either pointed or slightly notched, and margins are toothed. Leaves produce a
thin lather when rubbed with water.
Flowering occurs from July to September, and in some places, year-round. Tiny cream to
yellow-green flowers, each with five hooded petals surrounding a nectar disk, occur in
sparse, branched clusters in the leaf axils. Green and fleshy fruit, pea-size capsules less than
0.5 in. (1.3 cm) in diameter, mature to brownish-red from September to December. Each
three-part capsule contains three tiny grayish seeds.
Related or Similar Species. Colubrina species native to the United States are all trees or
tall shrubs, with erect branches and hairy leaves. Asiatic colubrina can be distinguished by
its sprawling habit, glabrous stems and leaves, and leaves with serrated margins. Three spe-
cies, all rare and endangered, are native to Florida. Coffee colubrina, also known as green-
heart, also has shiny leaves but the margins are entire. Twigs are covered with rust-colored
hairs, which may also cover the underside of leaves. Cuban nakedwood and soldierwood
are similar to coffee colubrina.
Introduction History. Asiatic colubrina was intentionally brought to Jamaica in the 1850s
by East Asian immigrants, probably for medicinal uses or as a soap substitute. From Jamaica,
the plant spread to other Caribbean islands, the Yucatan Peninsula of Mexico, and Florida,
494 n SHRUBS

most likely by ocean currents.

The earliest records of the spe-
cies in Florida are in 1937 in
the Keys and in the 1950s in
the Everglades on the mainland.
Habitat. Asiatic colubrina
grows on the upland, higher
areas of the coast in soils with
low permeability, above mean
high tide. It can invade either
disturbed or undisturbed habi-
tats, including mangrove for-
ests, tidal marshes, beach
dunes, and coastal strands.
Hammocks of tropical hard-
woods and buttonwood are
especially vulnerable because
they are slightly above flood
level. Plants can form thickets
along elevated road right-of-
ways that have been disturbed.
Although it is aggressively
spreading along the coasts, its
northern range may be limited
by frost.
Reproduction and Dispersal.
Asiatic colubrina reproduces
both sexually and vegetatively.
Although plants can flower and
fruit in their first year of growth,
few flowers on shrubs of any age
Asiatic colubrina occupies habitats above mean tide level in Florida. will produce fruit. The upper
(Native range adapted from USDA GRIN and selected references. flowers of each cluster usually
Introduced range adapted from USDA PLANTS Database, Invasive Plant abort. Mature seed capsules are
Atlas of the United States, and selected references.) dehiscent, sometimes explo-
sively expelling seeds. Little is
known regarding germination requirements, other than loose soil and that seeds cannot ger-
minate on exposed rock surfaces. Seedlings need light and will not grow well in the shade of
the parent plants. Even so, most seedlings develop in the vicinity of mature plants, and long-
distance dispersal is not common. The fruit and seeds, buoyant and salt tolerant, remain viable
for many months after floating in salt water and are frequently dispersed by ocean currents
and storm tides. Reports of viability of seeds in the soil are conflicting. Seeds may be viable
for less than one year or for 3–5 years. Although it is speculated that the pebble-like, hard
seeds are ingested by birds for use as crop stones, resulting in another means of long-
distance dispersal, no evidence indicates that birds eat the seeds.
The ability of Asiatic colubrina to spread vegetatively is tremendous. Roots will develop
from any branch that touches the soil. Any damage to the stem stimulates sprouting from
the roots and stump. Branches or stem pieces are distributed by storms and extreme tides.
 ASIATIC COLUBRINA n 495

A. Plants have long, drooping stems. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.) B. Shiny
leaves are alternate on the stem. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.) C. Leaves have
toothed margins. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.) D. Fruit are pea-size capsules.
(Forest and Kim Starr.) E. Small flowers grow in small clusters from leaf axils. (Forest and Kim Starr.)

Plants grow rapidly. Stems can grow as much as 32 ft. (10 m) in one year, and the species
can double its range in 8–10 years. Evidence indicates that Asiatic colubrina recovers from
hurricane damage more quickly than do native plant species.
Impacts. Considered one of the most aggressive alien weeds in Florida, Asiatic colubrina
frequently grows into a thick mat of tangled stems, which can be several feet thick. Climbing
plants cover other vegetation with a dense wall of leaves and vines that is almost impen-
etrable. Stems can grow more than 23 ft. (7 m) to the top of the tree canopy, loop back to
the ground and root, and then grow upward again. Plants cover native vegetation, smother-
ing them and shading out native species. Asiatic colubrina even climbs and smothers mature
Brazilian peppertree (see Trees, Brazilian Peppertree), another alien species. Because few
plants are able to persist under the dense mat, Asiatic colubrina reduces biological diversity.
The plant changes the habitat for wildlife and disrupts species interactions. It is especially a
problem in Florida’s coastal tropical hardwood forest, which support threatened and endan-
gered plants such as West Indian mahogany, Florida thatch palm, wild cinnamon, machi-
neel, and columnar cactus, as well as bromeliads and orchids. Asiatic colubrina also
threatens rare native species, such as bay cedar and beach star in the Florida Keys.
Asiatic colubrina’s clambering habit threatens cultural resources, archaeological sites
such as Native American middens, hiding them from view and damaging them with their
roots.
Removal of invasive Australian pine (see Trees, Australian Pine) allows more light to pen-
etrate to ground level, which may increase the growth of Asiatic colubrina.
Management. Asiatic colubrina is controlled primarily by physical and chemical meth-
ods. Physical removal by hand-pulling is possible for plants which are less than 5 ft.
(1.5 m) tall, but only if the roots are small enough to be removed as well. Care must be taken
not to disturb the soil, which would encourage germination of any viable seeds in the seed
bank. Mechanical methods of removal are generally not practical because Asiatic colubrina
is growing on top of and intermixed with other species. If no native species remain,
mechanical removal is more efficient. All branches must be carefully removed from the site
because the dry fruit is easily dislodged or shattered to spread seeds.
Although labor intensive because it is often difficult to find the main trunk among the
rambling vines, chemical applications of triclopyr, either to basal bark or to cut stumps,
496 n SHRUBS

Uses of Asiatic Colubrina

S everal cultures in Southeast Asia, Oceania, and the Caribbean use Asiatic colu-
brina for medicine and other purposes. Crushed leaves are used as a detergent
or shampoo. A leaf extract may be a remedy for centipede stings and other skin irri-
tations. The plant is also used to aid digestion, as a laxative, to expel internal para-
sites, and to reduce fever. A fruit mixture is used to induce abortions and to kill fish.
Leaves are used to weave mats, and the wood is carved for utensils.

are the most effective for adult plants. The cut-stump method is recommended if the stand is
fewer than 20 individuals. With more than 20 plants, applications to basal bark are more effi-
cient. If the infestation consists of more than 100 individuals and the vines cover all existing
vegetation, triclopyr may be applied to foliage. Because of prolific sprouting from the rooted
portion of the plant, monitoring and follow-up applications are necessary. Because of the pos-
sibility of viable seeds in the soil, the site should be monitored for several years.
Biological control of Asiatic colubrina is difficult because many native species in the
buckthorn family, including three endangered species of Colubrina in Florida, may also be
susceptible. The long-horn beetle (Artimpaza argenteonota), however, may be host-specific.
No microorganisms have been reported to attack Asiatic colubrina.

Selected References
Jones, David T. “Asiatic Colubrina.” Plants Gone Wild: Alien Plant Invaders in Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/
pdf/coas1.pdf.
McCormick, Cheryl M. “Colubrina Asiatica (Lather Leaf) Management Plan.” Colubrina Task Force.
Florida Exotic Pest Plant Council (FLEPPC), 2007. http://www.fleppc.org/Manage_Plans/CA%
20Mngt%20Plan.pdf.
Schultz, Gary E. “Element Stewardship Abstract, Colubrina asiatica.” Global Invasive Species Team,
Nature Conservancy, 1992; modified 2009. http://wiki.bugwood.org/Colubrina_asiatica.
Zheng, Hao, Yun Wu, Jianqing Ding, Denise Binion, Weidong Fu, and Richard Reardon. “Colubrina
asiatica, Asiatic Colubrina, Latherleaf.” In Invasive Plants of Asian Origin Established in the US and
Their Natural Enemies. U.S. Department of Agriculture, Forest Service, FHTET-2004-05, 2004.
http://wiki.bugwood.org/uploads/Colubrina.pdf.

n Brooms
Also known as: Scotch broom (Cytisus scoparius), Spanish broom (Spartium junceum),
French broom (Genista monspessulana), Portuguese broom (Cytisus striatus)
Synonyms: Sarothamnus scoparius, Spartium scoparium, Teline monspessulanus,
Genista junceum, Cytisus monspessulanus, and others
Family: Pea (Fabaceae)
Native Range. Scotch broom is native to most of Europe, and is also from the Canary Islands
and the Madeira Islands. Spanish broom is from Mediterranean Europe and North Africa, from
Portugal and Morocco east to Turkey, Syria, and the Caucasus region. French broom is native to
the Mediterranean regions of Europe and North Africa, from Portugal and Morocco east to
 BROOMS n 497

Turkey. Less widespread than

Spanish broom, Portuguese
broom is native to Portugal,
western Spain, and Morocco.
Distribution in the United
States. Scotch broom grows in
the western states, from Mon-
tana west to Washington, south
to California; and in the mid-
western and eastern states, from
Wisconsin east to Maine, south
to Georgia, west to Alabama,
Tennessee, and Kentucky. Both
Spanish broom and French
broom grow in coastal Wash-
ington, Oregon, and California.
Spanish broom is also found in
Texas. Portuguese broom is in
Oregon and California.
Description. The taxonomy
of brooms is complex. Genera
and species are not always well
differentiated, and many hybrids
occur. Unless otherwise stated,
discussion applies to all brooms.
Scotch Broom: Scotch broom
is a perennial shrub, usually 3–
6 ft. (1–1.8 m) tall, which can
reach heights of 12 ft. (3.5 m).
Depending on growing condi-
tions, shrubs may have many
branches or a single upright Scotch broom is the most common broom in the United States, growing
stem. The woody stems are in both the West and in the East. (Native range adapted from USDA
five-angled and are star-shaped GRIN and selected references. Introduced range adapted from USDA
in cross-section. Branches are PLANTS Database, Invasive Plant Atlas of the United States, and selected
green and almost leafless, but references.)
new twigs are hairy. The softly hairy compound leaves are small, with three leaflets, each
0.12–0.75 in. (0.3–2 cm) long. Plants are deciduous, without foliage from late summer to
early spring. They have a strong, branched taproot and are nitrogen fixing.
Flowers appear before the leaves emerge. Bright yellow typical pea-shaped flowers,
0.75–1 in. (2–2.5 cm) long, grow singly or in pairs from the axils of the upper leaves.
Greenish-brown to black seed pods, flattened with hairy edges, are 1–2 in. (2.5–5 cm) long.
Each pod contains 6–22 yellow-brown, shiny, oblong seeds,—approximately 0.15 in. long
and 0.08 in. wide (4 mm by 2 mm). Individual shrubs can live 10–15 years or more.
Spanish Broom: Spanish broom, also known as weavers’ broom, is difficult to distinguish
from its close relative Scotch broom. Shrubs stand 6–10 ft. (1.8–3 m) tall and are often leaf-
less. Lance-shaped, simple (not compound) leaves 0.5 in. (1.25 cm) long, appear from
February to June. Leaves are sparse, and plants are leafless from summer to early spring.
498 n SHRUBS

Its green stems, turning brown

as they mature, are finely ribbed
and round in cross-section.
Flowering slightly later than
Scotch broom, flowers of
Spanish broom are slightly
larger, 1 in. (2.5 cm) long,
occurring in open racemes at
the ends of stems. Seed pods,
hairy all over, are 2–4 in.
(5–10 cm) long and carry 15
reddish-brown seeds each.
Because Spanish broom grows
best in full sun with limited
water, it is the most drought-
resistant species.
French Broom: Stems of
French broom, also known as
Montpelier broom, have 8–10
ridges, are covered with silky
silvery hairs, and are round in
cross-section. Leafy stems of
the evergreen plant are lined
with three-part compound
leaves. Leaflets, variable in
size but usually 0.4–0.8 in.
(1–2 cm) long, are waxy above
and slightly hairy below. Its
flowers occur in clusters of
4–10 on short shoots among
the longer stems. French broom
Spanish broom is drought resistant and also tolerates salt spray on the is the most aggressive in terms
West Coast. (Native range adapted from USDA GRIN and selected of invasion characteristics.
references. Introduced range adapted from USDA PLANTS Database, Portuguese Broom: The ap-
Invasive Plant Atlas of the United States, and selected references.) pearance of Portuguese broom,
also known as striated broom,
is most similar to Scotch broom, but the plants live longer and are larger, as much as 10 ft.
(3 m) tall and 20 ft. (6 m) across. Stems have 8–10 sides or angles. Dark-green compound
leaves, with 1–3 leaflets, are sparse, with more growing toward the ends of stems. Flowers
are pale yellow, and seed pods 0.75–1.75 in. (2–4.5 cm) long, are inflated and covered with
white hairs.
Related or Similar Species. Bridal broom is distinguished from the others by its white
flowers and purple calyxes. It has round pods, each containing only 1–2 seeds, and the pods
do not split open as they dry. Gorse (see Shrubs, Gorse), an invasive shrub with a similar
appearance and an abundance of yellow pea-shaped flowers, is distinguished by its thorny
stems.
The native California broom is a smaller plant with both smaller flowers and leaves.
 BROOMS n 499

Introduction History. Scotch

broom was introduced to
California as an ornamental in
the 1850s. Other brooms were
also introduced by horticultur-
alists as garden plants. As early
as the 1930s, the U.S. Depart-
ment of Agriculture Soil Con-
servation Service introduced
and planted brooms for erosion
control along roads and to sta-
bilize mine tailings.
Habitat. Brooms grow pro-
lifically in disturbed areas, such
as roadsides and riverbanks,
but may also invade undisturbed
land. Plants spread easily to pas-
tures, cultivated fields, dry
scrubland, native grasslands,
dry riverbeds, and clearings in
forests due to logging or fires.
They tolerate a wide range of soil
types, moisture conditions, and
sun or part shade, but cannot
grow in deep shade. They grow
best in dry, sandy soils and full
sun. Plants tolerate light frost
but die back under severely cold
winters. Scotch broom is found
primarily in mountainous
and cool coastal areas with
dry summers, at elevations as French broom is invasive in more humid regions of the West Coast.
high as 5,500 ft. (1,675 m). (Native range adapted from USDA GRIN and selected references.
The northern limit of Scotch Introduced range adapted from USDA PLANTS Database, Invasive Plant
broom is probably due to cold Atlas of the United States, and selected references.)
winter temperatures, while the
southern limit is due to summer drought. It rarely grows on limestone. Scotch broom forms
pure stands or is mixed with Spanish broom. Spanish broom occupies poor soils, including
limestone, and also tolerates salt spray and temperatures as low as 14ºF (−10ºC). It is found
below 2,000 ft. (600 m) elevation. French broom is found in more humid and subhumid
regions in its native range, and grows below 1,650 ft. (500 m) elevation.
Reproduction and Dispersal. Brooms reproduce primarily by seed. Plants produce seeds
when they are about 3 ft. (1 m) high and 2–3 years old. They flower from March through
June and sometimes again in the fall. Flowers are insect-pollinated, which accounts for
crosses between species. Each shrub is capable of producing 2,000–3,500 pods. When pods
dry and burst in late summer, during August and September, seeds are forcefully expelled.
Seeds are further distributed by water, insects, birds, and other animals, and by human
activity such as on boots, vehicles, or equipment. Approximately 50 percent of seeds remain
500 n SHRUBS

dormant, and most germinate

better after scarification or a
cold, moist period of time.
Seeds buried deeper than 4 in.
(10 cm) in the soil remain dor-
mant. With a hard coat, seeds
remain viable for over 30 years
under field conditions. Ger-
mination begins with fall rains
and continues throughout win-
ter until early to midsummer.
Seedlings have a high mortality
rate during dry summers and
are susceptible to browsing by
wildlife during the first two
years.
Plants also sprout from root
crowns after cutting or burning.
The roots themselves do not
produce sprouts.
Impacts. Brooms are ideal
invaders. Plants grow rapidly,
flower young, produce abun-
dant seed that remains viable
for years, have a long life span,
and also sprout from the root
crown. Brooms displace native
plants, causing a decrease in
biodiversity. Plants outcompete
and prevent reestablishment of
native trees in forest clearings.
Portuguese broom is invasive in California. (Native range adapted from Nutrient dynamics are altered
USDA GRIN and selected references. Introduced range adapted from in broom-dominated sites
USDA PLANTS Database, Invasive Plant Atlas of the United States, and because the plant is a nitrogen
selected references.) fixer. Brooms pose a fire hazard,
especially on inaccessible steep
slopes, because they are highly flammable and the extensive plant material increases the fuel
load. All brooms contain quinolizidine alkaloids, particularly in flowers and seeds, which
are toxic to humans, wildlife, and most domestic livestock except goats.
Management. Eradication and control is difficult because Scotch broom plants produce
abundant seed and also sprout new stems from root crowns. Prevention is critical to control-
ling brooms. Sites should be monitored after disturbances, and new infestations eliminated.
Soil moved from one site to another should be checked for seeds. Even seeds lodged in
vehicle tires can spread brooms.
Because soil disturbance brings buried seeds close to the surface where they are able to
germinate, physical removal that leaves the soil intact is preferred. Hand-pulling or
uprooting is practical only on small populations or young plants. Mechanical means can
be used on larger infestations. Stripping bark all around the base of the shrub can kill the
 BROOMS n 501

A. Small leaves of brooms may be either simple or compound. (Richard Old, XID Services, Inc., Bugwood.org.) B.
Stems of Scotch broom are distinctly angled. (Robert Videki, Doronicum Kft., Bugwood.org.) C. Bright yellow
pea-shaped flowers cover the bushes. (Utah State University Archive, Utah State University, Bugwood.org.) D.
Flattened seed pods of Scotch broom have hairy edges. (Richard Old, XID Services, Inc., Bugwood.org.) E.
Seed pods of Portuguese broom are covered with white hairs. (William M. Ciesl, Forest Health Management
International, Bugwood.org.)

plant. Regardless of method, plants must be removed below the root crown to prevent
resprouting. Because pruning during the wet season will stimulate resprouting, trimming
or mowing should be done near the end of the dry season when plants are stressed.
Cutting or mowing after flowering may also kill the plants because their nutrient level is
low at that time. Cutting or mowing may need to be repeated over several years to deplete
the plants’ resources. Removal by burning has had mixed results because fire stimulates ger-
mination. Repeated fires on newly germinated plants, however, will help to deplete the seed
bank. Flamethrowers can be used for spot treatment. Grazing may be somewhat successful.
Although brooms are unpalatable and toxic to most livestock, they are less so when young.
Angora goats and Spanish goats eat the tops of young plants, which prevents flowering
and seed formation and also eventually depletes the root reserves. Goats have the advantage
of being able to negotiate steep slopes where equipment is not practical. Desirable vegeta-
tion, however, must be protected from goats. Because their digestive system destroys seeds,
chickens can be used to reduce the seed bank. Broom does not do well in deep shade, so
planting native trees and shrubs to increase shading may be helpful.
Young, actively growing plants are most susceptible to chemical control, best accom-
plished by spraying the foliage with glyphosate. A triclopyr solution is also effective when
painted on cut stumps or applied to basal bark.
Biological control is minimal. Although some insects feed on seeds, stems, or leaves of
Scotch brooms, they do only minor damage. The twig-mining moth (Leucoptera spartifoli-
lella) from Europe was released in California in 1960 and is now well established. This small
white moth lays eggs on new growth, and the larvae tunnel through the stems and emerge
the following spring. Affected twigs and branches die, but the effect on the whole plant is
unknown. The Scotch broom seed weevil (Apian fuscirostre), also from Europe, is now estab-
lished in California, Oregon, and Washington. Adults feed on stems, while larvae feed on
seeds inside the pods. Although seed production may be reduced by 60 percent, the larvae
have little effect on reducing infestations because of the great number of seeds each plant
produces. The shoot tip leaf moth (Agonopterix nervosa) from Europe was accidentally intro-
duced in the 1920s. Larvae feed on flower buds and young leaves, decreasing seed produc-
tion, but control is limited. All three insects are specific to Scotch broom and do not feed on
502 n SHRUBS

Beneficial Uses of Brooms

B room derives its name from the fact that its stiff branches were used in Europe
to make brooms. In its native range in Europe, Spanish broom is used for to
make baskets, mats, rope, and paper. A yellow dye is made from its flowers, and
an essential oil is used in perfumes. Scotch and Spanish broom flowers are used in
Europe for medicinal purposes, including cardiovascular problems, as a diuretic,
and to induce labor. No scientific evidence attests to their value, and they are not
approved for use in the United States. The plant is useful for erosion control
because its deep root structure binds soil.

other brooms. No other insects are currently under consideration in the United States. New
Zealand, however, is studying other possible biocontrol agents, including insects and
pathogens.

Selected References
“Brooms.” Plant Pest and Health Prevention Services (PHPPS), California Department of Food and
Agriculture, n.d. http://www.cdfa.ca.gov/phpps/ipc/weedinfo/brooms.htm#anchor384748.
“Brooms Cytisus spp., Genista spp., Spartium spp.” Invasive Weeds in Forest Land. Oregon State
University, 2008. http://extension.oregonstate.edu/catalog/pdf/ec/ec1598-e.pdf.
Nilsen, Erik. “Spartium junceum.” In Invasive Plants of California’s Wildland, edited by Carla C. Bossard,
John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000. http://www.
cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=79&surveynumber=182.php
“Scotch, French, and Spanish Broom, IVM Technical Bulletin.” Noxious Weed Integrated Pest
Management Guide, 2000. http://www.ipmaccess.com/Noxbroom.html.

n Exotic Bush Honeysuckles

Also known as: Amur honeysuckle (Lonicera maackii), Morrow’s honeysuckle (L. morrowii),
Tatarian honeysuckle (L. tatarica), Bell’s honeysuckle (L. x bella)
Synonyms: Lonicera insularis, L. sibirica
Family: Honeysuckle (Caprifoliaceae)
Native Range. Amur honeysuckle is native to east-central Asia, from southern China to
Korea and Japan. Morrow’s honeysuckle is native to South Korea and Japan. Tatarian honey-
suckle is from central Asia, including southwestern Russia, Kazakhstan, Kyrgyzstan, and
western China.
Distribution in the United States. Collectively, the four most aggressive species of exotic
bush honeysuckle grow in the majority of the country. Distribution of individual species is
scattered. Amur honeysuckle is naturalized in the eastern and midwestern states, from
New England west to North Dakota, south to Texas, and east to Georgia and the Atlantic sea-
board. It is not known in Minnesota, South Dakota, Louisiana, or Florida. Morrow’s honey-
suckle is primarily found in the northeastern and Atlantic states, from Maine south to
Georgia, and in the midwestern states, from Minnesota to Arkansas. It also grows in the
Rocky Mountain region, from Montana south to New Mexico. Tatarian honeysuckle is wide-
spread from the East Coast to the West Coast, with the exception of the extreme south and
 EXOTIC BUSH HONEYSUCKLES n 503

the desert states of Nevada and

Arizona. It also grows in Alaska.
Bell’s honeysuckle grows from
Maine south to North Carolina,
west to Missouri, and north into
the Dakotas.
Description. Although five
species and one hybrid of exotic
bush honeysuckles are known,
four species are considered to
be the most invasive. Species
may be difficult to distinguish.
All species are deciduous,
upright shrubs reaching 6–20 ft.
(1.8–6 m) tall. The glabrous twigs
do not have thorns. Bark is light
tan, and frequently flaking.
Older stems are often hollow,
with a brown pith. The opposite
dark-green leaves are oval or egg-
shaped, 1–2.5 in. (2.5–6 cm) on
short petioles. Different species
may sometimes be distinguished
by the amount of hairiness.
The paired tubular flowers,
less than 1 in. (2.5 cm) long,
are showy and grow in leaf axils.
Depending on the variety, flow-
ers bloom from May to June
and range from creamy white or
yellow to pink or crimson.
Species also differ in corolla and Found in the eastern half of the United States, Amur honeysuckle is
pedicel length. The dark-red to especially adapted to limestone soils. (Native range adapted from USDA
orange or occasionally yellow GRIN and selected references. Introduced range adapted from USDA
fleshy berries, paired in leaf axils PLANTS Database, Invasive Plant Atlas of the United States, and selected
references.)
like the flowers, mature from
September to October. Berries
may remain on the plant during winter, and each berry contains 2–6 small seeds.
Growing as tall as 30 ft. (9 m), Amur honeysuckle usually has several stems, with arching
branches and stringy tan bark. Its leaves, 1.5–3.5 in. (3.5–9 cm) long, taper to a long
pointed tip, and are slightly hairy and pale on the lower surface. The fragrant flowers,
0.75–1 in. (2–2.5 cm) long, are white or pink, fading to yellow. Berries are less than
0.25 in. (6 mm) in size and remain on the plant until mid-winter.
Short and broad, usually 5–8 ft. (1.5–2.5 m) tall and 6–10 ft. (2–3 m) wide, Morrow’s
honeysuckle becomes a tangled mound of foliage and branches covering the ground. Its
gray-green leaves are 1–2.5 in. (3–6 cm) long, wrinkled above and softly pubescent under-
neath. Flowers, 0.6–0.75 in. (1.5–2 cm) long, are also softly pubescent, supported by
504 n SHRUBS

densely hairy peduncles, or

short stalk. The white flowers
change to yellow as they age.
The red berries, which mature
June through August, are about
0.25 in. (6 mm) in diameter.
Tatarian honeysuckle grows
3.3–12 ft. (1–3.5 m) tall and
10 ft. (3 m) wide. Usually
multi-stemmed, the upper
branches of the shrub arch into
a dense mass of fine twigs.
Long flat scales on the bark pro-
duce little shredding. The gla-
brous leaves are 1.25–2.5 in.
(3–6 cm) long. The sessile flow-
ers (with no peduncle) are also
glabrous, usually pink but vary-
ing from red to white. The
red, or rarely yellow, berries,
0.25 in. (6 mm), occur singly
or in pairs with the bases fused
together. Although they ripen
from June through August, they
remain on the plant in winter.
Bell’s honeysuckle, a hybrid
between Tatarian honeysuckle
and Morrow’s honeysuckle, has
intermediate characteristics
between the two species and is
probably more common than
Morrow’s honeysuckle grows in a wide range of habitats, from the Rocky either parent. Difficult to distin-
Mountain states to the East Coast. (Native range adapted from USDA guish, Bell’s honeysuckle has a
GRIN and selected references. Introduced range adapted from USDA round growth habit, 4–10 ft.
PLANTS Database, Invasive Plant Atlas of the United States, and selected (1.2–3 m) tall, usually with an
references.)
equal spread. The leaves,
slightly hairy on the lower sur-
face, are 1–3 in. (2.5–7.5 cm) long. Because it is a hybrid, Bell’s honeysuckle flower color
is variable, usually pink fading to yellow. The red, or rarely yellow, fruit is 0.25–0.5 in.
(6–13 mm). Its roots are shallower than those of other exotic bush honeysuckles, less than
6 in. (15 cm) deep, and spreading beyond the crown of the shrub. Other hybrids may exist
but may not yet have become established in natural landscapes.
Related or Similar Species. Winter honeysuckle, also known as fragrant honeysuckle or
sweet breath of spring, and European fly honeysuckle, also known as dwarf honeysuckle,
are less invasive species of exotic bush honeysuckles. Winter honeysuckle was introduced
to the eastern United States from China in 1945. It is a deciduous or semievergreen shrub,
usually 3.2–10 ft. (1–3 m), but sometimes reaching 15 ft. (4.5 m) tall. The plant has an erect
and wide-spreading crown of tangled slender branches. Leaves are 0.5–3.5 in. (1.5–9 cm)
 EXOTIC BUSH HONEYSUCKLES n 505

long, and paired flowers are

0.5 in. (1.2 cm) long. Berries
are less than 0.5 in. (1.3 cm).
European fly honeysuckle is a
rounded shrub, 3.3–10 ft. (1–
3 m) tall and 10–12 ft. (3–
3.7 m) wide, with spreading
arching branches.
Several native plants, includ-
ing honeysuckles, resemble
exotic bush honeysuckles.
Canadian honeysuckle grows at
high elevations and reaches a
maximum height of 6.5 ft.
(2 m). Red honeysuckle, yellow
honeysuckle, and grape honey-
suckle are twining vines, not
shrubs. Native honeysuckles
are also distinguished by their
solid stems. Flowers of exotic
bush honeysuckles have hairy
styles, while flowers of the na-
tive species, except for swamp
fly-honeysuckle, do not. The
fruit of sweetberry honeysuckle
and bearberry honeysuckle,
also called twinberry honey-
suckle, is blue or black. The
slender twigs of coralberry are
purple to brown, and the leaves
of bush honeysuckle are finely
toothed and lance-shaped. Tatarian honeysuckle is widespread in the United States, and like all
Introduction History. Amur species of exotic bush honeysuckles, grows in disturbed open areas.
honeysuckle was brought to (Native range adapted from USDA GRIN and selected references.
New York Botanical Gardens in Introduced range adapted from USDA PLANTS Database, Invasive Plant
Atlas of the United States, and selected references.)
1898, and by 1931, it was avail-
able from at least eight nurseries.
Morrow’s honeysuckle was introduced to North America in 1875, and Tatarian honeysuckle
arrived as early as 1752. Exotic bush honeysuckles were promoted for wildlife cover and soil
erosion control, as well as for ornamental use, and are still sold in both private and state-run
plant nurseries. Tatarian honeysuckle has been used for reclamation of mining sites.
Habitat. Somewhat intolerant of shade, exotic bush honeysuckles are usually found in
open areas, such as abandoned fields, pastures, roadsides, and forest edges. Disturbed sites
are common locations. They also invade canopy gaps in woodlands, caused by grazing,
windthrow, or insect defoliation. Dense infestations are common beneath bird perches, such
as trees, shrubs and fence rows. Amur honeysuckle grows in a wide range of soil types and is
especially adapted to calcareous soils. It tolerates wet conditions, such as stream banks that
overflow, for short periods of time. It is adapted to cold. Morrow’s honeysuckle, Tatarian
506 n SHRUBS

honeysuckle, and Bell’s honeysuckle are found in the widest range of habitats, and also grow
in swampy areas, sandy plains, and lakeshores. All three species occupy soils that are poorly
drained to well drained, with low nutrient content.
Reproduction and Dispersal. Exotic bush honeysuckles reproduce primarily by seed, but
will resprout from the roots after the aerial parts are damaged. Bumblebees are the major
pollinators, and Bell’s honeysuckle may also be pollinated by hummingbirds. Although
information is sparse, exotic bush honeysuckles appear to be prolific in production of ber-
ries and seeds. Plants may produce fruit at 3–5 years of age, after reaching 3.3 ft. (1 m) tall.
One typical Bell’s honeysuckle in Wisconsin, approximately 6.5 ft. (2 m) tall, produced over
3,500 berries and over 20,000 seeds in one year. In southern Ohio, Amur honeysuckle and
European fly honeysuckle may produce 1.2 million berries per plant. Fruit-eating birds,
especially American Robins; White-tailed deer; and small mammals, such as deer mice, dis-
perse the seeds. Although most seeds have short-term viability and the soil seed bank appears
to be low, some seeds can germinate after 12 years. Little information is available regarding
germination requirements, which vary according to species, but some seeds may require a
cold dormant period or scarification from passage through an animal’s digestive tract.
Because plants are tolerant of only moderate shade, more germination takes place in sunny
sites. Establishment is best where litter cover is sparse and few herbaceous plants offer com-
petition. Cultivated land can be a barrier to the advance of exotic bush honeysuckles, while
forested areas, which provide habitat for birds which disperse seeds, can promote its spread.
Impacts. By aggressively creating a dense shrub layer, exotic bush honeysuckles shade
and crowd out native plants. They have a competitive advantage because they leaf out earlier
in spring and retain leaves later in the fall. They alter the habitat by decreasing light and
depleting soil moisture and nutrients. Plants may also be alleleopathic. Exotic bush honey-
suckle infestations restrict the growth of native plant seedlings and annual herbs, thereby
affecting natural biodiversity and interfering with succession in forest habitats. They may
also compete with native honeysuckles for pollinators, resulting in reduced seed production
from natives. Although the fruit is rich in carbohydrates, it supplies too little of the fat and
nutrients required by migrating birds. Berries of Amur honeysuckle are mildly poisonous.
Birds, particularly American Robins, that use exotic bush honeysuckles for nesting sites
may lose more eggs and young to predation due to the lack of protection afforded by the
thornless branches.
Management. Control or eradication of exotic bush honeysuckles requires a commitment
of 3–5 years. While it may be impossible to restore highly infested areas to a natural state, it
can be accomplished in newly infested areas. Sites with bare soil are susceptible to invasion
and must be monitored for seedlings.
Physical removal of seedlings and small shrubs is possible by pulling or digging, but soil
disturbance will promote seedling growth. Any root not removed may resprout. Repeated clip-
ping or trimming of branches may prevent dense stands from forming. Clipping in winter,
however, will encourage vigorous resprouting. All plant parts, especially fruit, must be bagged
and properly disposed. Prescribed annual burns will kill the top of shrubs and limit regrowth.
Burning should be done at least every other year, before seeds are dispersed, for several years.
Chemical application of glyphosate or triclopyr is frequently necessary. Foliar sprays are
appropriate where no risk to nontarget plants exists, such as in large monospecific thickets.
If grasses are present, the broadleaf selective triclopyr should be used. Individual shrubs can
be cut and the stumps treated with either glyphosate or triclopyr, but triclopyr or imazapyr
is best for basal bark applications. A flush of seedlings may follow herbicide treatments, but
can themselves be treated.
 EXOTIC BUSH HONEYSUCKLES n 507

What Method Works Best?

I n a scientific study of the effects of Amur honeysuckle (Lonicera maackii), Sahar

Haghighat, a biology student at Monmouth College, concluded that the best
method for increasing the biodiversity of understory plants where Amur honey-
suckle has invaded is to physically uproot and remove entire plants, including the
root system.
Working with the hypothesis that management or elimination of Amur honey-
suckle would increase the understory biodiversity, she compared three manage-
ment methods to ascertain their effects. She outlined three experimental plots
and one control plot, each 10 x 10 m2, with a buffer zone in between to ensure that
the control treatments were completely separate. Plots were in the forested areas
of LeSueur Nature Preserve in Monmouth, Illinois, which is a restored prairie with
prairie grasses and typical midwestern trees. Plot 1 was the control, where nothing
was done. In Plot 2, all Amur honeysuckle stems were clipped or trimmed, to
reduce their shading effect, to 30 cm above the ground. In Plot 3, the same amount
of clipping was combined with application of herbicide, a 50 percent concentra-
tion of glyphosate isopropylamine salt solution, painted on rather than sprayed
to prevent damage to surrounding plants. In Plot 4, plants were totally removed,
including all roots, by digging out or uprooting.
Beginning in early summer 2008 and ending in summer 2009, each plot was
sampled five times, recording species abundance and growth stage. Analysis
included a calculation of importance levels of each species. While Amur honey-
suckle was of course the most important species in the plots, its values dropped
over the course of the project. Other plants, primarily opportunistic nonnative
invaders such as garlic mustard (Alliaria petiolata) increased in abundance in the
treated plots. Although all four plots were similar at the start of the project, they
became increasingly different in terms of species composition as the project pro-
gressed, indicating that Amur honeysuckle was indeed responsible for the low bio-
diversity in sites where it dominated. While all treatments were followed by an
increase in biodiversity, total removal of the Amur honeysuckle plants resulted in
the biggest change. The most significant increase in biodiversity and biggest differ-
ence from the control plot was in the experimental plot where Amur honeysuckle
plants were uprooted.
Although results may be different with a greater number of samples or other
refinements of methodology, the conclusion of this simple but well-done study
was that physical removal of Amur honeysuckle by uprooting, while initially the
most difficult and time-consuming, is the best method to restore biodiversity of
the understory in wooded areas.
Source: Summarized from “Response of Lonicera maackii to Different Removal Methods
and the Ecological Effect upon Native Species,” by Sahar Haghighat, Department of Biology,
Monmouth College, Monmouth, IL, 61462, USA, May 12, 2010.

Because exotic bush honeysuckles appear to be free of diseases, insects, and predators, no
biological control is available. An aphid (Hyadaphis tatariacae) feeds on the tips and shoots of
Tatarian honeysuckle, and possibly on Morrow’s honeysuckle and Bell’s honeysuckle. The
resulting “witch’s broom” tuft of twigs may interfere with flowering. Any potential
508 n SHRUBS

organisms, however, must be host-specific to ensure that native honeysuckle species are not
affected.

Selected References
Batcher, Michael S., and Shelly A. Stiles. “Element Stewardship Abstract for the Bush Honeysuckles.”
Global Invasives Team, Nature Conservancy, 2000. http://www.invasive.org/weedcd/pdfs/
tncweeds/loni_sp.pdf.
“Bush Honeysuckles.” Southeast Exotic Pest Plant Council (SEEPPC) Invasive Plant Manual, 2003.
http://wiki.bugwood.org/Archive:SEEPPC/Bush_Honeysuckles_-_Lonicera_spp.
Munger, Gregory T. “Lonicera spp.” In: Fire Effects Information System. U.S. Department of Agriculture,
Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, 2005. http://www.fs
.fed.us/database/feis/plants/shrub/lonspp/all.html.
Williams, Charles E. “Exotic Bush Honeysuckles.” Weeds Gone Wild: Alien Plant Invaders of Natural
Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2005. http://www.nps.gov/
plants/alien/fact/pdf/loni1.pdf.

n Gorse
Also known as: Common gorse, furze, Irish furze, whin
Scientific name: Ulex europaeus
Synonyms: None
Family: Pea (Fabaceae)
Native Range. Central and western Europe and the British Isles, where it is not aggressive
and is cultivated as hedgerows to enclose livestock.
Distribution in the United States. Pacific Coast, from California, north to Washington.
Scattered in southern California as well as on the Atlantic coast, from Massachusetts south
to North Carolina, where it is not an aggressive invader. Also in Hawai’i.
Description. Gorse is a very prickly evergreen shrub, usually growing 1.5–10 ft.
(0.5–3 m) tall, but occasionally reaching 16 ft. (5 m). The woody shrub branches densely
from the base of the plant. The stout branches are longitudinally ridged. Young green twigs
are covered with gray to reddish-brown hairs, while older branches turn brown and become
stiff, intertwined, and thornlike. Young plants are mat-like in form, but as plants age, they
tend to grow outward, forming a ring of growing vegetation around a dry, dead center.
One plant may be as large as 30 ft. (9 m) in diameter. Under sustained windy conditions,
plants may be sculpted into cushion or mat forms. Individual plants live for about 30 years.
Gorse has different types of leaves according to the age of the plant. Seedlings grow in a
compact rosette-leaf pattern. Juvenile plants, usually up to 2–4 in. (5–10 cm) tall, have com-
pound leaves typical of the pea family, with three leaflets. Leaves on larger plants are alter-
nate and reduced to awl-like stiff spines. Branches may appear leafless because of
the reduced leaf structure. Branches and twigs terminate in a bottle brush of spiny leaves
0.5–2 in. (1.3–5 cm) long.
Gorse has two types of roots, a fibrous system of lateral roots in the top few inches of soil
and a deeper taproot.
Bloom season varies with latitude, but generally occurs from March through May. In
Hawai’i, flowering begins in December. Inflorescences, containing either single flowers or
small clusters, grow in leaf axils and near the ends of twigs. One plant may have thousands
of flowers, which completely cover the bush. Shiny, bright yellow flowers are distinctly pea
 GORSE n 509

like, with a two-lipped corolla.

Flowers are 0.8–1 in. (2–
2.5 cm) long, and the calyx is
densely covered with soft hairs.
Fragrant flowers attract pollina-
tors, and seed pods mature
about two months later. The
fruit is a very hairy pod, 0.5–1
in. (1.3–2.5 cm) long, usually
containing 2–6 hard olive-
brown kidney shaped seeds. A
fleshy, yellow protrusion on
one end of the seeds is rich in
oil and protein.
Related or Similar Species.
Gorse may be confused with
one of the brooms, such as
Scotch broom (see Shrubs,
Brooms), which are also inva-
sive species, but those plants
have no spines. Gorse also
forms denser, more impen-
etrable thickets. Dwarf gorse,
reaching only 10 in. (30 cm) tall,
is native to eastern England and
western France, Spain, and
Portugal. It has smaller flowers
and softer spines, and is not
naturalized in the United States.
Introduction History. Gorse
was probably brought to the
East Coast by immigrants, who Although present on both coasts, gorse is invasive only in the West.
used the plant extensively in (Native range adapted from USDA GRIN and selected references.
Europe, including for animal Introduced range adapted from USDA PLANTS Database, Invasive Plant
fodder and as a yellow dye. Atlas of the United States, and selected references.)
Gorse was introduced to the
West Coast, possibly to Mendocino County, California, or to Oregon, prior to 1894. By
the 1950s, it had spread to western Washington, western Oregon, and northern
California, with localized infestations on 15,000 ac. (6,000 ha) in California and on
25,000 ac. (10,000 ha) in Oregon. Gorse was introduced to Hawai’i as a hedge plant and
ornamental sometime before 1910.
Habitat. Gorse most frequently invades disturbed sites and can be found growing on
sand dunes, gravel bars, overgrazed pastures, logged areas, and burned sites. It can also
occupy forests, riparian sites, coastal scrub, and coastal cliffs. Because gorse is a nitrogen
fixer, it is capable of growing on infertile soils and most every soil type, including silty,
rocky, and clay. It can grow on serpentine soils, occasionally on calcareous sites, and can
colonize barren mine dumps. Tolerant of acidity, it prefers a pH of 4.5–5.0, but also requires
trace elements, such as boron, to grow well. It grows best in shady exposures but is
510 n SHRUBS

A. Gorse is a densely branched woody shrub. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.)
B. Leaves on large stems are reduced to stiff spines. (George Markham, USDA Forest Service, Bugwood.org.)
C. The hairy seed pods contain hard seeds. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.) D.
Pea-shaped flowers grow in showy spikes. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.)

intolerant of heavy shade. Gorse requires moist, but well-drained, soils because the
nitrogen-fixing bacteria are rendered ineffective under flooded conditions.
Gorse requires a temperate climate and cannot tolerate drought. Its sensitivity to extremes
of heat and cold limits its spread away from the coast. Its latitudinal range may be limited by
photoperiod. Short summer days limit both flowering and thorn formation.
Reproduction and Dispersal. Although gorse reproduces both vegetatively and sexually,
it spreads predominantly by seed, and is a prolific seed producer. Plants can produce 14 mil-
lion seeds per ac. (35 million per ha) each year. The upper inch (2.5 cm) of soil may contain
2,000 seeds per sq. ft. (20,000 per m2). Pods are explosively dehiscent, opening forcefully
when dry and ejecting the seeds 3.3–10 ft. (1–3 m) from the parent plant. Although small,
seeds are too heavy to be carried by wind, but are dispersed by water, ants, ground birds
such as quail, other animals, and human activity. Improperly dumped garden waste, hikers,
machinery, vehicles, and fill dirt may also be responsible for seed distribution.
Because the seed coat is very hard and impenetrable by water, seeds are viable for
30–50 years. Seeds usually germinate in the wetter winter and spring months, but germina-
tion may occur any time of year if conditions are suitable. Germination increases substan-
tially, either due to more light or higher soil temperatures, when mature plant cover is
removed. Heat, particularly from fires, stimulates germination, but temperatures above
212ºF (100ºC) are lethal to seeds. Plants need a minimum of 18 months to grow from a
seedling to reproductive age.
Plants also vigorously resprout from stumps. Stems touching the ground will root, and
some evidence suggests that gorse also spreads by creeping rhizomes.
Impacts. Although initially slow to spread, gorse impacts the environment in a variety of
ways. It forms dense, spiny thickets that remain impenetrable even when plants are dead. It
outcompetes both native and cultivated plants, in part due to its nitrogen-fixing bacteria,
which allow it to colonize soils that are poor in nitrogen. Nitrogen can accumulate at
20–30 pounds per ac. (22–34 kg per ha) every year. Gorse leaf litter makes the soil more
acid and less suitable for native species. By replacing grassland, gorse thickets result in a loss
of valuable pasture. By displacing native plants, it also decreases native wildlife habitat. In
Hawai’i, where gorse occupies approximately 35,000 acres (14,000 ha) on both the island
of Hawai’i and Mau’i, it threatens mamane forests and native birds, such as the palila.
 GORSE n 511

Because plants grow rapidly, a lot of dry plant litter accumulates. A new stand can accu-
mulate 5,500 lb. per acre (6,000 kg per ha) of dry material in its first year. Dry matter accu-
mulation in older stands can be 90–180 lb. per acre (100–200 kg per ha) every year. The oils
in the foliage and litter are highly flammable, not only making gorse an extreme fire hazard
but also causing fires to burn hotter.
Gorse produces abundant pollen, and the major pollinator is the honey bee, making the
plant attractive to commercial beekeepers.
Management. The most effective control for gorse is rapid response, eliminating the shrub
when it first invades. In order to be effective, management efforts must be long term because of
the longevity of viable seeds. Preventing seed production is an option, and Hawai’i has tried
restricting commercial bee keepers from setting hives near gorse. Reseeding with native plants
should follow methods of control or eradication. Because gorse is intolerant of shade, it may be
suppressed by taller plants. Research in both chemical and biological control is being con-
ducted in Australia and New Zealand where gorse is also seriously invasive.
The spiny nature of gorse makes it difficult to remove by physical means. Young plants,
less than two years old and under 5 ft. (1.5 m) tall, may be removed by hand-pulling or dig-
ging, but all root pieces must be removed. Repeated cutting of aerial parts may deplete
nutrient stores in roots, especially if done when reserves are lowest, just as plants begin to
flower. In spite of the fact that burning will not remove either the roots or the seed bank
in the soil, fires are most efficient in removing large plants. Although fire will kill seeds in
the top 0.4–0.8 in. (1–2 cm) of soil, the deeper ones are stimulated to germinate. Repeated
burning, however, may reduce the seed bank. Physical removal of the upper part of the
shrub, down to the stumps, may be necessary to gain access to the base of the plant for her-
bicide application. Cutting or burning will also reduce seed production. Any physical
removal of the top portion of plants should be followed by herbicide application to prevent
resprouting. Grazing, particularly by angora goats, can control seedling and resprouts, but
mature plants are too woody and spiny to be palatable to herbivores. A minimum of two
years of grazing of new growth is needed to show any reduction in gorse. The digestive sys-
tem in chickens destroys many weed seeds, including gorse, making chickens effective at
reducing the seed bank.
A number of herbicides, such as dicamba, triclopyr, 2,4-D + triclopyr, picloram, tordon
and glyphosate, may provide chemical control of gorse. Some types, however, are not effec-
tive on mature plants because of the structure of the spiny leaves and their waxy cuticle,
which prevents absorption of the herbicide. Seedlings are effectively eliminated by picloram,
glyphosate, or triclopyr. Cut stumps of mature plants are best treated with glyphosate.
Because different herbicides have different effects on nontarget species, the choice of which
to apply depends on what remaining native vegetation must be protected. Regardless of the
herbicide used, treatments must be repeated.
No biological means of control are totally effective at controlling or eradicating gorse. The
gorse weevil (Apion ulicis), from France, is established in California and Oregon. The larvae
feeds on seeds inside the pod. When the pods ripen and open, the adults emerge to eat
spines and flowers. Effects are limited in cool coastal climates because the adult weevils
die inside the pod before the pods ripen and burst. The gorse spider mite (Tetranychus lintea-
ris), introduced to and established in Hawai’i, damages plants but does not reduce popula-
tions. Colonies of the spider mites weave a tent-like web on the gorse plants and eat leaves
of the mature plant. Effectiveness of other insects now established in Hawai’i, such as a
foliage-eating moth (Pempelia genistella) and gorse thrips (Sericothrips staphylinus), is not yet
known.
512 n SHRUBS

Selected References
Hoshovksy, Marc. “Element Stewardship Abstract, Ulex europaeus.” Global Invasive Species Team,
Nature Conservancy, 1989; updated 2001. http://www.imapinvasives.org/GIST/ESA/esapages/doc
umnts/ulexeur.pdf.
Markin, George. “Ulex europaeus L., Common Gorse.” In The Woody Plant Seed Manual, Agricultural
Handbook No. 727, edited by Franklin T. Bonner and Robert P. Karrfalt. U.S. Department of
Agriculture, Forest Service, 2008. http://www.fs.fed.us/rm/pubs_other/wo_AgricHandbook727/
wo_AgricHandbook727_1140_1142.pdf.
Motooka, P., et al. “Ulex europaeus.” In Weeds of Hawai’i’s Pastures and Natural Areas: An Identification
and Management Guide. College of Tropical Agriculture and Human Resources, University of
Hawai’i at Manoa, 2003. http://www.ctahr.hawaii.edu/invweed/WeedsHI/W_Ulex_europaeus.pdf.
“Ulex europaeus.” Pacific Island Ecosystems at Risk (PIER). Hawaiian Ecosystems at Risk (HEAR), n.d.
http://www.hear.org/pier/species/ulex_europaeus.htm.

n Japanese Barberry
Also known as: No other names
Scientific name: Berberis thunbergii
Synonyms: Berberis thunbergia var. atropurpurea, Berberis sinensis, Berberis japonica
Family: Barberry (Berberidaceae)
Native Range. Japan
Distribution in the United States. Eastern half of the United States, from Georgia north to
New England, west to the Great Plains states, and from North Dakota south to Nebraska.
Also in Wyoming and Washington.
Description. Japanese barberry is a compact and spiny deciduous or semievergreen shrub
with many branches. It usually grows 2–3 ft. (0.6–0.9 m) tall, but occasionally reaches 8.2 ft.
(2.5 m). The deeply grooved brown branches form slight zigzags, rather than growing
straight. Plants are hairless, but have one very sharp, simple spine in each leaf axil. The inner
bark and wood is yellow. Leaves grow in tight, alternate bunches close to the branches. They
are spoon-shaped or narrowly oval and wedge-shaped at the base. Leaves have smooth mar-
gins and vary in size, 0.5–1.5 in. (1.3–4 cm) long. Plants are distinguishable by the leaf
color, ranging from bright green to burgundy. In warmer southern climates, plants may
retain leaves during the winter. The root system is shallow, but tough.
An abundant number of small pale-yellow flowers, 0.3–0.4 in. (8–10 mm) long, appear
from March to May, depending on geography. Clusters of 2–4 flowers hang in small umbels
along the entire stems of the shrub. The small, oblong fruits are bright red berries, approx-
imately 0.3 in. (8 mm) in size, maturing from July through October. Although slightly juicy,
they are solid. The clustered berries remain on the stems until spring.
Related or Similar Species. Common barberry, a nonnative plant from Europe and
western Asia, was formerly used extensively by early settlers for hedgerows, dye, and jam.
However, because it was host to the black stem grain rust transmitted by the fungus
Puccinia graminis, grain crops were devastated. Beginning in 1916, steps were taken to eradi-
cate common barberry. Berberis x ottawensis is a hybrid of common barberry and Japanese
barberry. American barberry, the only barberry native to North America, is found in dry
woodland. It can be distinguished by its sharply toothed leaves and three-prong spines.
Introduction History. Japanese barberry was introduced to the United States as an ornamen-
tal plant in 1875, when seeds from Russia were sent to Arnold Arboretum in Boston. In 1896, it
was planted at the New York Botanical Garden and recommended as a substitute for the
 JAPANESE BARBERRY n 513

common barberry because it is

not a host for the black stem
grain rust. In 1910, Japanese bar-
berry became naturalized in the
Northeast after it escaped cultiva-
tion from plantings at vacation
home sites. It is still a popular
landscape plant, with several cul-
tivars sold in nurseries.
Habitat. Although it first col-
onizes open fields and pastures,
Japanese barberry grows in a
variety of habitats, including
roadsides, railroad and utility
right-of-ways, and fencelines. It
grows best with at least part
sun, but because it is shade toler-
ant, it can form dense stands in
several types of forest, such as
floodplain, planted, and succes-
sional. It is less common, how-
ever, in forests dominated by
oaks or on north-facing slopes.
Well-drained sites are preferred,
but it also occupies wetlands.
Japanese barberry does not com-
pete well with grasses. Severe
drought or extremely cold win-
ters do not negatively impact
either plants or seed production.
Deer generally avoid Japanese
barberry but browse other veg- Imported as a substitute for common barberry, Japanese Barberry is
etation, giving the invaders more prominent in the northern Great Plains and eastern states. (Native range
room to spread. adapted from USDA GRIN and selected references. Introduced range
Reproduction and Dispersal. adapted from USDA PLANTS Database, Invasive Plant Atlas of the
United States, and selected references.)
Japanese barberry reproduces
both sexually and vegetatively.
Plants are prolific seed producers, and the germination rate is as high as 90 percent. Birds,
most often ground birds such as Wild Turkey, Bobwhite Quail, Ring-necked Pheasant,
and Ruffed Grouse, eat the fruit and deposit seeds elsewhere. Other birds drop seeds when
they rest on power lines or on trees at forest edges or roadsides. Small mammals, such as
rabbits, also eat the fruit and distribute seeds.
Plants sprout from rhizomes, and branches will root when in contact with moist soil.
Impacts. Japanese barberry plants can form dense, continuous stands that threaten natu-
ral areas by displacing native plants and reducing habitat and forage for native birds and
mammals. Because it is one of the first plants to produce leaves in spring, it shades out native
understory plants. Growth can be rapid, especially for seedlings, as much as 2–4 ft. (0.6–
1.2 m) in one year. Shrubs also change soil chemistry, altering nitrogen content and
514 n SHRUBS

A. Shrubs are compact and spiny. (Richard Old, XID Services, Inc., Bugwood.org.) B. Twigs slightly zigzag, with
a sharp spine in each leaf axil. (James H. Miller, USDA Forest Service, Bugwood.org.) C. Spoon-shaped leaves
may remain on the plant in warmer climates. (James H. Miller, USDA Forest Service, Bugwood.org.) D. The
oblong fruit are bright red berries. (Richard Old, XID Services, Inc., Bugwood.org.) E. Flowers hang in short
clusters. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology, University of Connecticut, Bugwood.org.)

biological activity in the soil. The species raises the soil pH and, by retaining its leaves, may
reduce the depth of litter where it is semievergreen.
Management. Because Japanese barberry seeds are abundant and dispersed easily, reduc-
ing seed production should be a priority in control. Whatever type of physical control is
used to eliminate Japanese barberry, care should be taken to minimize soil disturbance.
Plants are easy to pull out of forest soils in early spring, an option that is feasible only if num-
bers are low. Hand-pulling or digging out whole plants is effective when plants are less than
3 ft. (0.9 m) tall. All connected roots must be removed and remains properly disposed of to
prevent accidental dispersal of seed. Uprooting plants, which can be done any time of year,
reduces both the population and seed development. Plants are easier to see in winter, how-
ever, when native species are leafless. Mowing or cutting, as short as possible, effectively
controls the spread of Japanese barberry by reducing flower and seed development, but will
not kill plants. Mowing should be done at least once each growing season. Fire kills Japanese
barberry and is a good option, but only in fire-adapted communities.
Because of detrimental effects on nontarget native plant species, chemical applications
should be used only on plants that are difficult to remove by hand. Foliar sprays, such as
glyphosate and triclopyr, are effective when temperatures are above 65ºF (18ºC). Both her-
bicides can also be used on cut stumps, a more selective method than spraying, any time the
soil is not frozen.
No biological controls are available.

Selected References
“Japanese Barberry.” Invasive Plant Atlas of New England (IPANE). University of Connecticut, 2009.
http://nbii-nin.ciesin.columbia.edu/ipane/icat/browse.do?specieId==26.
“Japanese Barberry.” Southeast Exotic Pest Plant Council (SEEPPC) Invasive Plant Manual, 2003.
http://www.invasive.org/eastern/eppc/barberry.html.
“Japanese Barberry.” Weed of the Week. U.S. Department of Agriculture, Forest Service, Invasive
Plants, 2005. http://www.na.fs.fed.us/fhp/invasive_plants.
 KOSTER’S CURSE n 515

Miller, Arthur E. “Black Stem Rust Quarantine.” Invasive Plants of the Eastern U.S., 2003. http://www
.invasive.org/eastern/other/barberry.html.
Swearingen, Jil M. “Japanese Barberry.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/ALIEN/fact/
beth1.htm.

n Koster’s Curse
Also known as: Soapbush, clidemia, camasey, nigua
Scientific name: Clidemia hirta
Synonyms: Melastoma hirta, Clidemia crenata, M. elegans, C. Elegans, M. hirtum
Family: Melastome (Melastomataceae)
Native Range. Tropical Central America and South America, from Mexico to Paraguay
and northern Argentina. Also in the Greater and Lesser Antilles in the Caribbean, including
Puerto Rico and the Virgin Islands.
Distribution in the United States. Hawai’i.
Description. Koster’s curse is a branched perennial shrub, either scrambling or growing
upright. Branches and twigs are round and slender, and the plant is covered with straight
red bristles or hairs that become lighter with age. When many stems arise from one root-
stock, the shrub can become very dense. Depending on habitat conditions, mature shrubs
may be 3.3–16.5 ft. (1–5 m) tall, shorter in exposed sites and taller in moist, shady locations.
The oval leaves, 2–7 in. (5–18 cm) long and 1–3 in. (3–8 cm) wide, are opposite on the stem
and grow on short petioles, 0.2–1.2 in. (0.5–3 cm) long. Each leaf has 5–7 prominent veins
running from the base to the pointed tip. Many lateral veins intersect the longer veins, mak-
ing the leaf appear checkered or pleated like seersucker fabric. Leaves are slightly hairy, and
leaf margins are smooth or slightly scalloped or serrated. During dry seasons, the plant may
lose its leaves, but will generate new leaves when adequate rain falls. Plants have a shallow
lateral root system with many fine rootlets.
Small white flowers, each with five petals, grow in short panicles, approximately 1 in.
(2.5 cm) long, both at the ends of branches and in leaf axils. The calyx is hairy. Purple to
black, fleshy berries, 0.3 in. (7 mm) in diameter and covered with bristles, taste a bit like
blueberries. Each four-celled berry contains 100 or more very small coffee-colored seeds.
Related or Similar Species. Other members of the Melastome family present in Hawai’i
include velvet tree (see Trees, Velvet Tree) and two Tobouchina species, cane ti and glory-
bush, all native to tropical South America. Velvet tree is a branching plant, as tall as 45 ft.
(13.5 m), with large, velvety leaves. Its leaves are as long as 3 ft. (1 m), green on the upper
surface and purple below. Its upright inflorescences of tan flowers are not showy. Both
Tobouchina species are smaller shrubs. Cane ti reaches 9 ft. (2.5 m) tall, with smaller leaves,
3 in. (7.5 cm) long. Its flowers are pink, with four petals and bright yellow anthers. Because
it does not spread by seeds, glorybush is a less invasive plant. It grows to 12 ft. (3.5 m) tall,
with 5 in. (12.5 cm) long leaves, and five-petaled purple flowers. Bristletips, native to the
Himalayas and southwestern China, is also a smaller shrub 6.5–13 ft. (2–4 m) tall, with pen-
dulous branches. Its leaves, 3–6 in. (8–16 cm) long, have only five parallel veins, and its
pinkish-purple flowers grow in drooping clusters.
Introduction History. In the 1880s, a man named Koster imported coffee plants from
Central or South America to develop a plantation in Fiji and accidentally introduced Clidemia
seeds. The plant quickly spread from Fiji to other Pacific Islands and became known as
Koster’s curse because of its invasive properties. Although the date it reached Hawai’i is
516 n SHRUBS

unknown, it was first recorded

on O’ahu in 1941. It may have
been an intentional introduction
through the horticulture trade or
as a specimen in a botanical gar-
den or zoo. The tiny seeds could
also have been a contaminant
and brought to the island acci-
dentally. By 1957, it was recog-
nized as a noxious plant, and by
1988, it covered approximately
247,100 ac. (100,000 ha) on
O’ahu alone. Plants are found
on almost all the major islands,
and it is still expanding its range.
Habitat. Even in its native
range, Koster’s curse spreads
most rapidly in disturbed soil,
including floodplains, pastures,
agricultural fields, and road-
sides. It rarely occurs in forests
in its native range and appears
to be more tolerant of shade in
locations to which it has been
introduced. The plant grows in
a wide range of soil types, but
needs moisture. Rainfall in its
native range, where it grows
from sea level to 5,000 ft.
(1,500 m) elevation, is 47–157 in.
(1,200–4,000 mm). It, there-
Koster’s curse is primarily an invader of disturbed sites in Hawai’i. (Native fore, has the potential to invade
range adapted from USDA GRIN and selected references. Introduced all wet and mesic habitats
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the below 5,000 ft. (1,500 m)
United States, and selected references.) elevation, and may extend even
higher. Although it grows best
in full sun, it is tolerant of shade and can grow in both open and closed habitats, even with
100 percent canopy cover, under both native and introduced trees. Koster’s curse is pri-
marily a weed on disturbed sites, where gaps, caused by storms, feral pigs (see Volume 1,
Vertebrates, Mammals, Feral pig), landslides, and fire, occur in the forest subcanopy.
Infestations are especially linked to disturbances by feral pigs. The shrub can also invade
undisturbed areas, but its populations remain low.
Reproduction and Dispersal. Koster’s curse reproduces both by seed and vegetatively.
Where conditions are continually moist, Koster’s curse will flower and fruit year-round. It
is a prolific seed producer. A large plant can produce more than 500 fruit each year, each
containing 100 seeds, for a total of 50,000 seeds annually. Seeds are dispersed in several
ways. Many seeds are distributed locally by nonnative species, such as birds, feral pigs,
and mongooses, as well as by humans. Feral pigs carry seeds in their hooves. Hunters,
 KOSTER’S CURSE n 517

A. Several stems can sprout from one rootstock. B. Prominent veins give the leaves a distinct pattern. C. Small flowers
grow in the leaf axils or at the ends of branches. D. Fleshy berries are covered with bristles. (Forest and Kim Starr.)

hikers, and vehicles are responsible for long-distance dispersal. Seeds adhere to hikers’
boots, which may account for the spread of Koster’s curse from trail to trail or to other
islands. Infestations in abandoned forest clearings formerly used to grow marijuana indicate
another probable vector for long-distance dispersal. Seeds remain viable in the soil for four
years, and plants can reach maturity and fruiting size in 6–10 months.
Stems sprout from the root stalk and from cut stumps and also root where they contact
moist soil. Even loose leaves have been known to grow roots.
Impacts. In its native range, Koster’s curse lives only a few years before dying, due to dis-
ease or insect damage. However, in the absence of disease and insects where it has been
introduced, the shrub can create monotypic thickets. Its high seed production, high seedling
establishment, large seed bank, and early maturation give it the means to be a significant
invader in a broad range of environments. Due to its rapid growth, Koster’s curse may domi-
nate the understory in forests after only one year. By shading out understory native plant
species, it inhibits forest regeneration and natural plant succession. It reduces biodiversity
and is a serious threat to understory plants on tropical islands. It is considered one of the
worst alien plants in Hawaiian natural ecosystems and is also a weed in plantation agricul-
ture. Sheep will not eat it, and it is toxic to goats.
Management. Because disturbance is the key to major invasions, control of feral pig pop-
ulations will limit the amount of disturbed ground, as well as decrease the seed dispersal.
Hikers and hunters should thoroughly wash their boots after leaving infected areas, so as
to minimize seed dispersal. Because of the abundant seed production and seedbank, any
control must be done on new stands before fruit sets for the first time.
Most physical efforts at control, such as pulling up or cutting plants, fail unless they are
followed by herbicide treatments. Plants are quickly replaced from the soil seed bank, and
uprooted plants often resprout or reroot. Because of resprouting, mowing or cutting plants
only once is not effective. Any physical control must be repeated for as long as 10 years to
deplete root reserves and ensure eradication.
Chemical applications of a broadleaf herbicide offer the best short-term control but are
not practical in many areas of Hawai’i. Variation in results may be due to differences in
518 n SHRUBS

Is Koster’s Curse Always a Curse?

I n its native range in Puerto Rico, Koster’s curse is beneficial because it colonizes
disturbed areas and is food for wildlife. It is used in Brazil as treatment for a skin
infection, Leishmania braziliensis.

application techniques, and more research is needed, but more than one herbicide applica-
tion is usually necessary. Herbicides should be used on seedlings and on uprooted plants to
prevent sprouting. Glyphosate is effective as a foliar spray. A triclopyr solution is sometimes
effective when applied to foliage, cut stump, or to basal bark.
Biological control is the best possibility for a long-term solution. Although biological
agents fail to eliminate the plant, they decrease its competitive advantage. Two insects that
have been released in Hawai’i have met with only partial success. A thrips (Liothrips urichi)
from Trinidad, released in 1953, is effective in open areas, such as pastures and fields, but
not in shaded locations. A beetle (Lius peisodon), however, attacks plants in the shade. A fun-
gus (Colletotrichum gloesporioides), introduced in 1986, provides some control in drier sites,
both open and shaded, but is ineffective in wetter habitats. Three moths (Antiblemma acclina-
lis, Carposina bullata, and Mompha trithalama) released in the 1970s have been unsuccessful.

Selected References
Francis, John K. “Clidemia hirta (L.) D. Don.” U.S. Department of Agriculture, Forest Service,
International Institute of Tropical Forestry, San Juan, Puerto Rico, n.d. http://www.fs.fed.us/
global/iitf/pdf/shrubs/Clidemia%20hirta.pdf.
Gerlach, Justin. “Clidemia hirta.” ISSG Global Invasive Species Database, 2006. http://www.issg.org/
database/species/ecology.asp?fr=1&si=53.
Nelson, R. “Koster’s Curse” Untamed Science, 2010. http://www.untamedscience.com/biodiversity/
plants/flowering-plants/dicotyledons/myrtales/melastomataceae/clidemia/kosters-curse.
Tunison, Tim. “Element Stewardship Abstract, Clidemia hirta.” Global Invasive Species Team. Nature
Conservancy, 1991; modified 2009. http://wiki.bugwood.org/Clidemia_hirta.

n Lantana
Also known as: red sage, yellow sage, prickly sage, big sage, blacksage, flowered sage,
lantana wildtype, largeleaf lantana, prickly lantana
Scientific name: Lantana camara
Synonyms: Lantana aculeata, L. camara var. nivea, L. camara var. aculeata
Family: Verbena (Verbenaceae)
Native Range. Tropical North and South America, specifically the Caribbean Islands,
including Bermuda, the Bahamas, Greater Antilles, Lesser Antilles, and Puerto Rico, and
from Mexico south to Colombia and Venezuela. Because lantana has been cultivated for
more than 300 years as an ornamental, it is difficult to determine exactly where it originated.
Distribution in the United States. Southern United States, from North Carolina west to
California. Also on all the main Hawaiian Islands, Puerto Rico, and the Virgin Islands.
Description. Lantana has many cultivars, making it a highly variable species in its growth
habit, flowers, leaves, growth rates, shade tolerance, spines, and toxicity to livestock.
 LANTANA n 519

Lantana is a perennial shrub

with multiple stems growing
from its base, at or close to
ground level. It may be as tall
as 10 ft. (3 m). It is usually erect
in open habitats, but prostrate,
or occasionally clambering
(climbing onto and over shrubs
and trees) in scrubland or for-
ests. It may grow in individual
clumps or in thickets. Ever-
green in the Tropics and Sub-
tropics, lantana is deciduous in
cooler climates. Stems are four-
angled. Stems on plants that
grow on the U.S. mainland usu-
ally have no spines, but the
variety in Hawai’i usually has
prickly stems. Yellow-green to
green leaves are oval and broad-
ly lance-shaped, 0.8–6 in.
(2–15) cm long and 0.8–2.5 in.
(2–6 cm) wide. They are oppo-
site on the stem and stiff, with
serrated edges. Covered with
small rough hairs, the leaf surfa-
ces feel rough, like fine sand-
paper. When crushed, leaves
are aromatic, reminiscent of
black currents.
The inflorescences, growing
in the leaf axils near the ends Lantana has many cultivars and is widely available as an ornamental.
of the stems, are compact, flat- (Native range adapted from USDA GRIN and selected references.
topped flowerheads, about Introduced range adapted from USDA PLANTS Database, Invasive Plant
1 in. (2.5 cm) in diameter. Flo- Atlas of the United States, and selected references.)
wer color is variable, typically
becoming darker as the flowers age, changing from yellow to orange to red, or from white
to pink to lavender. The central flowers and the outer ring of flowers are different colors.
Lantana produces abundant fruit. Round, fleshy berries, 0.2 in. (6 mm) in diameter, hang
in clusters like a blackberry. They are green when immature and purplish black when ripe.
Each individual fruit contains 1–2 seeds.
Related or Similar Species. Now endangered, pineland lantana, also called depressed
shrubverbena, is native to Florida. It is distinguished from the invasive lantana by a dis-
tinctly tapered leaf base and yellow flowers. The two species, however, hybridize, making
identification difficult.
Introduction History. Lantana was introduced to Europe from Brazil in the seventeenth
century. Although it may have been introduced to Europe multiple times, no records exist.
In the 1700s, lantana was a favorite ornamental in Europe, leading to many cultivars. In
520 n SHRUBS

A. Many stems grow from the base of the plant. (Rebekah D. Wallace, Bugwood.org.) B. Opposite leaves are
toothed. (Forest and Kim Starr.) C. Round berries are fleshy. (Richard Old, XID Services, Inc., Bugwood.org.)
D. and E. Flat-topped clusters of showy flowers almost cover the shrub. (Forest and Kim Starr.)

the 1800s and 1900s, lantana became widely distributed worldwide, with hundreds of cul-
tivars and hybrids. No record indicates exactly when lantana was introduced to the main-
land United States, but it was taken to Hawai’i as an ornamental in 1858. It became
naturalized on the Hawaiian Islands by 1871 and was widespread. Plants are sold in nurser-
ies throughout the United States.
Habitat. Because so many cultivars and varieties exist, lantana has a broad tolerance to
environmental factors and occupies a variety of habitats, including forests, riparian sites,
pastures, and citrus groves. It thrives in both shady and sunny sites, but generally grows best
in open, sunny locations, such as wasteland, forest edges, beaches, transportation corridors,
and burned or logged forests. Somewhat shade tolerant, it is found beneath the canopy of
open forests, but cannot grow in heavy shade. Lantana usually colonizes disturbed sites,
and natural fires stimulate its growth. It benefits from destructive activities of foraging ani-
mals, such as pigs, cattle, sheep, or goats. It grows well on all types of well-drained soils,
but cannot tolerate boggy conditions. Although resistant to both drought and salt spray, it
does not grow on saline soils. Able to thrive in poor soils and quick to invade disturbed sites,
plants may be used in reclamation of mine spoils.
In Hawai’i, lantana occurs in low elevations that range from dry to moist, predominantly
along roadsides, in vacant lots, pastures, forest, shrubland, and natural grassland.
Worldwide, it is found in regions with 10–160 in. (250–4,000 mm) of annual rainfall. It
grows from sea level to 6,500 ft. (2,000 m) elevation, but most varieties are limited by frost.
Stems die back when temperatures drop to 28ºF (-2 C), but they resprout in spring.
Reproduction and Dispersal. Reproduction is both sexual and vegetative. Under favor-
able temperature and moisture conditions, lantana has an almost continuous bloom period.
In seasonal climates, peak flowering usually coincides with the rainy season. Flowers are
pollinated by insects, especially butterflies and thrips. Reports of self-fertilization are con-
flicting. One plant produces about 12,000 fruit. Birds, and occasionally other animals, eat
the fruit, which accounts for long-distance dispersal of the seeds. Six nonnative birds,
including the Chinese Turtledove and the Common Myna (see Volume 1, Vertebrates,
Birds, Common Myna), disperse lantana seeds in Hawai’i, where no native birds have been
observed feeding on the fruit. Fruit that is not eaten becomes dry and remains on the shrub
for some time. The low germination rate of seeds is enhanced by passage through a digestive
 LANTANA n 521

tract. Germination requires high light conditions, and seedlings do not survive beneath the
parent bushes. Given suitable conditions, germination can take place all year. Plants grow
rapidly and may reach 10 ft. (3 m) tall in 3–4 years.
Plants can regenerate from the base after stems are damaged. Large plants can survive
fires and mowing because new stems grow from the base, but not from broken or damaged
roots. Stems will root where they contact the soil, and new plants can be propagated by
cuttings.
Impacts. Lantana is a pest in both natural areas and in agriculture. By first growing on
creek banks and roadsides, the shrub more easily invades adjacent disturbed natural ecosys-
tems. It often forms dense thickets in disturbed forest and pasture, crowding out native spe-
cies and reducing biodiversity. It dominates the understory layer of forests, where it
suppresses growth of native plants. It is thought to have alleleopathic compounds, which
limit growth of plants nearby. By interfering with forest reproduction, lantana can change for-
est communities into shrubland. In Hawai’i, lantana displaces native plants in dry habitats.
Lantana can become the dominant understory shrub in orchards, plantations, or open
forests. It is a weed in agricultural crops and reduces the carrying capacity of pasture. It is
a serious economic pest in citrus groves in Florida, where its alleleopathic substances reduce
vigor and productivity. Thick stands impede access to commercial forests and create fire
hazards by increasing the biomass. Spiny varieties hinder human access and interfere with
recreational activities, such as hiking.
Leaves and unripe fruit are poisonous. If ingested by livestock, such as cattle, sheep,
goats, or horses, or by wildlife, the plant can cause liver failure and death. Most animals
avoid the plant. In home garden settings, children have been poisoned by eating the unripe
fruit, occasionally causing death.
Management. An integrated management approach, including physical removal, burn-
ing, chemicals, shading, prevention, and revegetation, is most effective. The best prevention,
however, is to not purchase and plant lantana in the home garden and to maintain healthy
ecosystems with native plant biodiversity. Rapid revegetation of disturbed areas, such as
burns or cleared sites, will prevent lantana seedling development. Lantana cannot reestab-
lish under the dense shade of closed tree canopies.
Physical methods range from cutting off flowers to removal of entire plants. Spread can
be prevented by removing flower heads or fruit clusters from the plant before they are ripe.
Small plants can be hand-pulled. Fire or mowing reduces the biomass, thereby reducing the
amount of chemicals required for herbicide applications. Fire is ineffective, however, unless
followed by chemical treatments for several years. Digging out plants as follow-up is labor
intensive and not practical for large stands.
Chemical applications work best when clumps are first cut and herbicide is applied to the
stumps or to the new sprouts. Regrowth should be sprayed when it is 1.5–5 ft. (0.5–1.5 m)
tall. Foliar applications of glyphosate are not very effective, failing to prevent regrowth from
the base. Fluroxypyr plus aminopyralid sprayed twice in six months works well, as does
fluroxypyr or imazapyr applied to the base of the plant.
Between 1902 and 1969, more than 20 insects were introduced to Hawai’i for biological
control, with varying results. The most effective agents are a caterpillar (Hypena strigata),
which defoliates plants; a fly (Ophiomyia lantanae), which destroys seeds; and the lace bug
(Teleonemia scrupulosa), which damages the plant by feeding on leaves. Those introduced
insects are now a problem on ornamental lantana shrubs, indicating that any biological con-
trol will be controversial because of the landscape plant industry and private gardens. A ben-
efit of biological control is that it reduces the volume of individual plants, making other
522 n SHRUBS

Uses of Lantana

P lanted worldwide as an ornamental, lantana may also have value as an herbal

medicine. It is used as a folk medicine for many ailments, including chicken
pox, asthma, eczema, tumors, high blood pressure, tetanus, and rheumatism.
Boiled leaves are applied to the skin to alleviate pain and reduce swelling.
Lantana oil is used for itchy skin, as an antiseptic, and for such skin conditions as
leprosy and scabies. Alkaloids may relieve constipation and lower blood pressure.
Leaves contain an insecticidal or antimicrobial substance. Potatoes stored with lan-
tana leaves, for example, sustain no damage from potato tuber moths
(Phthorimaea operculella). Because of its toxicity, lantana offers possibilities for
biocides or herbicides. Research has shown that a leachate can kill waterhyacinth.

control methods easier. No biological agent has totally controlled lantana, perhaps due to its
genetic diversity. Different research is required for specific areas of the world, because what
is effective in Australia may not control different varieties of lantana in Florida or in South
Africa.

Selected References
Francis, John K. “Lantana camara.” U.S. Department of Agriculture, Forest Service, International
Institute of Tropical Forestry, San Juan, Puerto Rico, n.d. http://www.fs.fed.us/global/iitf/pdf/
shrubs/Lantana%20camara.pdf.
MacDonald, Greg, Brent Sellers, Ken Langeland, Tina Duperron-Bond, and Eileen Ketterer-Guest.
“Lantana species.” Excerpted from Identification and Biology of Non-Native Plants in Florida’s
Natural Areas, by K. A. Langeland and K. Craddock Burks. IFAS Publication SP 257. Center for
Aquatic and Invasive Plants, University of Florida, IFAS, 2008. http://plants.ifas.ufl.edu/node/223.
Motooka, P., et al. “Lantana camara.” In Weeds of Hawai’i’s Pastures and Natural Areas: An Identification
and Management Guide. College of Tropical Agriculture and Human Resources, University of
Hawai’i, Manoa, 2003. http://www.ctahr.hawaii.edu/invweed/WeedsHI/W_Lantana_camara.pdf.
Walton, Craig. “Lantana camara (Shrub).” ISSG Global Invasive Species Database. 2006. http://
www.issg.org/database/species/ecology.asp?fr=1&si=56.

n Multiflora Rose
Also known as: Rambler rose
Scientific name: Rosa multiflora
Synonyms: Rosa cathayensis
Family: Rose (Rosaceae)
Native Range. Eastern Asia, including eastern China, Korea, Taiwan, and Japan.
Distribution in the United States. Most of the contiguous United States except for the
northern Rocky Mountain states and Great Plains states, from the Dakotas west to Idaho,
and the desert states of Nevada and Arizona. Absent from Alaska, Hawai’i and Puerto Rico.
Description. Multiflora rose is a thorny, perennial shrub that can reach 10–15 ft.
(3–4.5 m) tall and be 9–13 ft. (2.7–4 m) wide. Many long, arching stems, called canes, which
arise from the root crown, are covered with hard, recurved thorns. Leaves, on 0.5 in.
 MULTIFLORA ROSE n 523

(1.3 cm) long petioles, are alter-

nate and pinnately compound,
about 3–4 in. (8–10 cm) long.
Each leaf has 4–11 oval or
oblong leaflets, which are
sharply toothed. The base of
each leaf has a pair of fringed
or finely dissected bracts called
stipules.
Showy panicles of fragrant
flowers appear in May and
June. Usually white, sometimes
pink, flowers are 0.5–1 in.
(1.3–2.5 cm) in diameter, with
five petals. Clusters of fruit,
called rose hips, hang from each
panicle. The rose hips may be
either glabrous or pubescent
and become bright red as they
mature in mid-to-late summer.
Later in the fall, they become
leathery. They do not burst to
release seeds and remain on
the plant. Each fruit contains
an average of seven seeds,
which are yellowish to tan and
irregularly shaped, 0.08–0.16 in.
(2–4 mm) long.
Related or Similar Species.
More than 80 species or sub-
species of roses, some native,
some escaped from cultivation, Because it was introduced multiple times and used for several purposes,
and some confined to gardens, multiflora rose is widespread in the United States. (Native range adapted
grow in the eastern United from USDA GRIN and selected references. Introduced range adapted
States. Registered cultivars nu- from USDA PLANTS Database, Invasive Plant Atlas of the United States,
and selected references.)
mber an additional 8,000.
Maccartney rose was brought
from Asia to southeast Texas for use as a hedge row. Because it sprouts vegetatively and
forms dense thickets, it has become invasive along the Gulf coast. Stems have both recurved
and straight thorns. Each leaflet on the pinnately compound leaf is 1–3 in. (2.5–7.5 cm)
long. White flowers with many yellow anthers grow in small clusters. Although named for
the Cherokee Indians who cultivated it in the 1800s, Cherokee rose is native to China.
Introduced by a plantation owner in 1759, this rose naturalizes so readily that it was
believed to be native as late as 1916. It has many large, reddish-orange prickles, a three-
leaflet compound leaf, and white flowers for a short time in early spring. It is a serious weed
in the rich soils of the “black belt” in Alabama, where it occupies hundreds of acres of land.
The dog rose, a scrambling species from Europe and western Asia, is not invasive and is
prized for its large rose hips as a natural food source of vitamin C. The vines have recurved
524 n SHRUBS

A. Canes are long and arching. (Ohio State Weed Lab Archive, The Ohio State University, Bugwood.org.) B.
Leaflets have serrated margins. (James H. Miller, USDA Forest Service, Bugwood.org.) C. Leaf bases have
fringed bracts. (James H. Miller, USDA Forest Service, Bugwood.org.) D. Showy flowers are usually white.
(James H. Miller, USDA Forest Service, Bugwood.org.) E. The fruit, rose hips, remain on the plant. (James H.
Miller, USDA Forest Service, Bugwood.org.) F. Plants have prominent recurved thorns. (Chris Evans, River to
River CWMA, Bugwood.org.)

or hooked spines and pinnately compound leaves with 5–7 leaflets. Fragrant flowers can be
deep pink or white but are usually pale pink. Sweetbriar rose from Europe, and rugosa rose
from China, are both common and widespread in the eastern United States, but are generally
not invasive. Young stems of sweetbriar rose are smooth, but become prickly and hairy with
age. Flowers are pink to pale pink. Leaves of rugosa rose are a deep glossy green, heavily
veined and wrinkled. Stems are very spiny, densely covered with needlelike thorns.
Flowers range from white or yellow to pink or purple, and the rose hips are large, 1 in.
(2. 5 cm) in diameter.
Native rose species that may be confused with multiflora rose include pasture rose,
climbing prairie rose, swamp rose, prickly rose, smooth rose, and Virginia rose, all with pri-
marily pink flowers 2–3 in. (5–7.5 cm) in size. Two shrubby berry species are also similar.
Allegheny blackberry has 1 in. (2.5 cm) white flowers, while purpleflowering raspberry
has 2 in. (5 cm) pinkish-purple flowers. None of the native roses are invasive.
Introduction History. Multiflora rose has been introduced into the United States many
times since the late 1700s, both as garden plants and as root stock for ornamental roses.
An introduction from Japan in 1886, of plants to be used as grafting stock for ornamental
roses, is well documented. Multiflora rose is no longer used and is not available in nurseries.
Beginning in the 1930s, the U.S. Soil Conservation Service recommended multiflora rose
to check soil erosion and for use as living fences to control livestock. State conservationists
recommended it as wildlife cover for ground birds and cottontail rabbits and as food for
songbirds. Over 14 million free cuttings from state conservation departments were planted
in West Virginia in the 1940s–1960s, and over 20 million were planted in North Carolina.
However, some states such as Kentucky refused to promote multiflora rose, with the result
that Kentucky as a whole is relatively free of invasion. Multiflora rose, however, is prominent
in forested land in Kentucky. Plants were also planted on medians of highways, both as
crash barriers and to reduce the glare of oncoming headlights. Rose hips are also used in
tea for their vitamin C content.
 MULTIFLORA ROSE n 525

Habitat. A prolific grower, multiflora rose invades old fields and agricultural land in the
southeastern United States. In the northeastern and midwestern states, it is predominantly a
weed in pastures and other unplowed lands. On the Great Plains, it was frequently used as
wind breaks. It can also be found in prairies, along roadsides, on hillsides, fence rows,
right-of-ways, margins of swamps and marshes, and in open woods. Multiflora rose grows
best on deep, well-drained soils, but tolerates a wide range of soil and environmental condi-
tions. It grows in both shade and sun, but does best in full or partial sun. It grows in either
damp or dry soils, but cannot tolerate standing water. It is less vigorous in the northern
states because of cold winters, and is not found where winter temperatures drop below -
28ºF (−33ºC). In contrast, south of central Georgia, it is only found as deliberate plantings
because it needs cold temperatures to stimulate germination.
Reproduction and Dispersal. Multiflora rose reproduces both sexually and vegetatively.
A large plant may have 40–50 panicles, with an average of 50–60 rose hips each, meaning that
an average mature plant can produce about 500,000 seeds per year. Approximately 90 percent
of the seeds are viable. Long-distance dispersal is accomplished by winter-feeding birds, such
as Cedar Waxwings, Mockingbirds, Starlings, Red-winged Blackbirds, and American Robins,
which eat the fruit and expel the seeds. Passage through a bird’s digestive tract scarifies
the seeds, which facilitates germination. Seedlings are often concentrated beneath places
where birds perch. Deer also eat the fruit. Seeds remain viable for as long as 20 years and
germinate easily. Seedlings grow close to the ground for the first 1–2 years. Local spread is
due to the habit of the long, drooping canes, which root where the tips touch the soil.
Impacts. Multiflora rose has both economic and ecological impacts. Over 45 million areas
in the United States are infested with multiflora rose. Infestations lower land values of agri-
cultural, forestry, and recreational areas. When used as a hedge between fields, it may lower
crop yield by siphoning nutrients away from crops. Pastures on hilly terrain or steep slopes
are often infested. Forage is reduced, and cattle avoid grazing pastures that support the
thorny plants, resulting in a decrease in beef production.
Because of its prolific growth and tendency to form dense, impenetrable thickets, it
crowds out native plants, not only displacing the plant communities, but also the wildlife
and insects that depend on them.
Management. Type and effectiveness of physical control depends on the size of plants or
the infested area. Although small seedlings can be pulled, removing plants by hand is time-
consuming. Repeated mowing will prevent establishment of seedlings, but it should be done
3–6 times per growing season for 2–4 years. Easily done in flat grassy pastures, mowing is
difficult in wooded or brushy areas. Mowing cannot be done on mature clumps that are over
10 ft. (3 m) tall and 20 ft. (6 m) in diameter. Bulldozing of large plants and thickets is an
option, but subsequent control of seedlings, which sprout on disturbed soil, is necessary.
Burning has not been tried with multiflora rose, but it has been partly successful in Texas
on Macartney rose, where 90 percent of top growth was killed. Regrowth from root crowns,
however, occurred within two weeks. Periodic burning combined with spraying with herbi-
cides is necessary to completely kill plants. Dead plant debris remaining from herbicide
treatment can be efficiently burned.
Any chemical control requires monitoring and follow-up because of the long-lived soil
seed bank. Application of a foliar spray, such as glyphosate, 2,4-D, picloram, dicamba, tri-
clopyr, or fosamine, in spring offers control if repeated for two years. Picloram, however,
is not effective if applied directly to the soil. An advantage to fosamine is that it affects only
woody species, leaving herbaceous species undamaged. Application of glyphosate or other
herbicides to freshly cut stumps is very effective.
526 n SHRUBS

Multiflora rose is tolerant to many North American insects and diseases. Any introduced
biological control must not damage ornamental roses. Three possibilities are well established
in the United States. Larvae of the European rose chalicid (Megastigmus aculeatus var. nigro-
flavus), a small torymid wasp, hatch within the seeds and feed on them. Although native to
the same world region as multiflora rose, this insect was discovered in a New Jersey nursery
in 1917, where rose seed was imported from Japan for creation of root stocks. The insect
spread slowly from that location. After overwintering in the empty seed, the insects emerge
as adults the next summer to lay eggs and renew the cycle. Adult chalicids are limited flyers
and remain with the plant unless the seeds are transported. Because of the limited flying
ability of the adults, the species is not widespread in the United States, and would have to
be introduced locally where multiflora rose is invasive. Because most plants were distributed
by cuttings, not seed, distribution of chalicid wasp lagged behind that of multiflora rose.
Because the rose chalicid has been found to infect 95 percent of seeds imported from
Japan, it is expected that it will infect the same percentage in the United States. Although
it may take 3–5 decades, it has the potential to drastically reduce the invasiveness of multi-
flora rose.
The virus-like rose rosette disease, which is native to North America and spread by a tiny
native eriophyid mite (Phyllocoptes fructiphilus), causes flowers to develop abnormally and
leaves and shoots to turn deep red or purplish. Stems are enlarged and become thornier.
Plants also develop a broom-like growth from profuse sprouting of lateral buds. When tem-
peratures drop below 14ºF (-10ºC), the affected canes die. The disease was first reported in
California and Wyoming in 1941, on both ornamental roses and on the wild native Woods’
rose, which is common to the Rocky Mountains, western Plains, California and Arizona. The
disease spread eastward, and was found on a multiflora rose in a Nebraska nursery in 1964
and in Pennsylvania and West Virginia by 1994, sparking concern over its potential damage
to ornamental roses. This pathogen has been fatal to multiflora rose from the Great Plains to
Pennsylvania and Maryland. It also affects ornamental roses, causing a yellow pattern on the
leaves, increased thorniness, clumped and wrinkled foliage, and broom-like lateral bud
growth. The pathogen is disseminated by air currents and can also be transmitted through
grafting affected plants.
The rose stemgirdler (Agrilus aurichalceus) is a small brownish-golden metallic colored
beetle, 0.2–0.4 in. (5–9 mm) long. Originally from Europe, it is now established in eastern
North America. As the larvae burrow beneath the bark, they girdle the stem, which kills
the stem above the girdle. A drawback is that the stemgirdler may also affect other rose spe-
cies and some blackberries and raspberries (Rubus spp.).

Selected References
Amrine, J. W., Jr. “Multiflora rose.” In Biological Control of Invasive Plants in the Eastern United States, by
R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-04, 2002. http://
www.invasive.org/eastern/biocontrol/22MultifloraRose.html.
Bergmann, Carole, and Jil M. Swearingen. “Multiflora Rose (Rosa multiflora).” Weeds Gone Wild: Alien
Plant Invaders of Natural Areas. Plant Conservational Alliance’s Alien Plant Working Group, 2009.
http://www.nps.gov/plants/alien/fact/pdf/romu1.pdf.
Eckardt, Nancy, and TunyaLee Martin. “Element Stewardship Abstract, Rosa multiflora.” Global
Invasive Species Team, Nature Conservancy, 1987; revised 2001; modified 2009. http://
wiki.bugwood.org/Rosa_multiflora.
 RATTLEBOX n 527

n Rattlebox
Also known as: Chinese wisteria, Spanish gold, scarlet wisteria, red sesbania,
purple sesbane, false poinciana, black acacia
Scientific name: Sesbania punicea
Synonyms: Sesbania tripetii, Daubentonia punicea
Family: Legume (Fabaceae)
Native Range. South America, specifically Uruguay, southern Brazil, southern Paraguay,
and northeastern Argentina.
Distribution in the United States. Southern United States, from Texas and Arkansas east
to Florida and as far north as
Virginia. It has recently been
recorded west of the Rocky
Mountains in the Central
Valley of California.
Description. Rattlebox is a
deciduous woody shrub or small
tree, growing as tall as 15 ft.
(4.5 m). The gray or reddish-
brown bark is covered with len-
ticels. Mature trees have an open,
spreading crown. The alternate
leaves are pinnately compound,
4–8 in. (10–20 cm) long, with
10–40 dark green leaflets.
Oblong leaflets, each about 1 in.
(2.5 cm) long, have pointed tips
and are arranged opposite on
the leaf stem, with no terminal
leaflet. Leaflets are droopy.
Orange-red or coral-colored
flowers, with the characteristic
shape of legumes or peas, appear
in spring and early summer.
Flowers, each 0.8–1.2 in.
(2–3 cm) long, hang clustered
in dense sprays up to 10 in.
(25 cm) long. The dark-brown
woody seed pods, which hang
in clusters on short stalks, are
3–4 in. (7.5–10 cm) long with a
pronounced pointed tip. Pods
are distinguished by four wing-
Rattlebox grows best in climates with long, hot summers, in California
like projections running the and the southern states. (Native range adapted from USDA GRIN and
length of the seed pod. The 3– selected references. Introduced range adapted from USDA PLANTS
10 seeds are loose inside parti- Database, Invasive Plant Atlas of the United States, and selected
tions within the pod, making a references.)
528 n SHRUBS

A. Rattlebox has a spreading crown. (Charles T. Bryson, USDA Agricultural Research Service, Bugwood.org.)
B. Compound leaves have no terminal leaflet. (Karan A. Rawlins, University of Georgia, Bugwood.org.) C. Seed
pods are distinctively shaped. (Karan A. Rawlins, University of Georgia, Bugwood.org.) D. Showy flowers grow
in long clusters. (Karan A. Rawlins, University of Georgia, Bugwood.org.) E. Loose seeds inside the pods make
a rattlling sound. (Karan A. Rawlins, University of Georgia, Bugwood.org.)

rattling sound similar to a rattlesnake when shaken. Seeds are released when the pod shatters,
but seed pods often remain on the plant through the winter.
Related or Similar Species. The genus Sesbania has many species, both annual and
woody. Some are native to the United States, and some have commercial value. The three
described below are all deciduous and have the distinctive rattling seedpods.
Bagpod, also called bladderpod, is an annual, native to the southeastern United States,
that reaches 13 ft. (4 m) tall. Found on disturbed moist or wet sites, it has a smooth stem
and few or no branches. Stem tips have dense white hairs. Leaves are alternate and pinnately
compound, but longer than rattlebox leaves, as long as 12 in. (30 cm). Leaves have 20–40
leaflets, each 1.2 in. (3 cm) long, with smooth margins. Leaflets are very hairy when they
first emerge, but become smooth as they mature. Flowers usually have yellow petals, but
they are variable and often pink or red. The fruit is a dry, smooth inflated pod 0.8–2.4 in.
(2–6 cm) long, containing two seeds. Green plants are not favored by livestock, but dried,
mature seed pods may be eaten by goats and cattle in fall and winter. This plant, however,
is toxic, especially the seeds. Freshly dried seeds are more toxic than older ones remaining
on the plants. Bagpod often grows intermixed with hemp sesbania.
Hemp sesbania, native to the United States, is an annual growing to 13 ft. (4 m) tall, with
few to no branches. Its bark is smooth, and stem tips are not hairy. Leaves can be as long as
12 in. (30 cm), with 20–70 leaflets. Linear or oblong leaflets, 1.5 in. (3.5 cm) long, have
smooth margins and pointed tips. Leaflets are smooth above, but waxy beneath. The yellow
flowers may be streaked or spotted with purple. Seedpods are smooth, 4–8 in. (10–20 cm)
long, and contain 30–40 seeds each. It grows in disturbed sites and along water courses as
far north as New York. Hemp sesbania is a noxious weed in Arkansas because it invades
crops, such as peanuts and cotton.
Drummond rattlebox, also native to the United States, is a woody perennial shrub or
small tree growing 2–15 ft. (0.6–4.5 m) tall. The green or light-brown bark is smooth.
Leaves are alternate, 4–8 in. (10–20 cm) long, with 20–50 leaflets, each 0.5–1.5 in. (1.2–
4 cm) long. The dull-green leaflets fold up in response to hot sun. Although it has many
branches in its canopy, their wide separation gives the tree an open appearance. The 0.5
in. (1.2 cm) flowers are yellow, sometimes streaked with red, hanging in 2 in. (5 cm) long
clusters. The shrubby tree occupies low-lying wet sites. In the northern part of its range,
 RATTLEBOX n 529

the higher aerial parts freeze back in cold weather, but the lower woody parts remain
unharmed.
Introduction History. Rattlebox was introduced to California in 1930, and probably
much earlier to the East Coast. Because of its showy coral-colored flowers and persistent, rat-
tling seed pods, this species is still sold and planted as an ornamental. It has escaped culti-
vation and invaded natural habitats.
Habitat. Rattlebox prefers moist areas with long, hot summers and is found along rivers
and streams, where it receives full sun. It is also somewhat shade tolerant, and young plants
can be found growing in the shade of mature rattlebox trees. It can survive freezing condi-
tions for a few days. Leaves are damaged at 23ºF (−5ºC), and the entire plant will die at
14ºF (−10ºC).
Reproduction and Dispersal. Rattlebox reproduces solely by seeds. Each plant produces
hundreds of seed pods and thousands of seeds each year. Pods continue to fall from the
branches almost all year. Buoyant seed pods, which can float for up to 10 days, are dispersed
by water and have a high germination rate, especially after the seed coat has been abraded by
sandy running water. Seeds that fail to germinate remain dormant in the soil for years. Seed
counts have averaged over 100 per sq. ft. (1,000 per m2). Seedlings grow quickly, as much
as 3.3 ft. (1 m) in the first year. Plants become reproductively mature in their second year.
Impacts. Rattlebox displaces native vegetation, which in turn decreases habitat for wild-
life and reduces biodiversity. Its worst impact is around water bodies, such as rivers, lakes,
and stream banks. Dense thickets that cover 50–100 percent of the ground can interrupt
water flow and alter water quality. Thickets also block access for recreational uses. By colo-
nizing gravel bars and islands, the plants increase channel roughness, which increases flood
potential.
All parts of rattlebox, especially the seeds, are poisonous to birds, reptiles, humans, and
other mammals. Even a few seeds can be fatal to birds. The toxic component is saponic gly-
cosides. Symptoms of ingestion include nausea and vomiting, associated with abdominal
pain and diarrhea. Severe reactions result in respiratory failure or death. Rattlebox is
included on the list of toxic plants for horses.
Management. The best control is to limit planting of rattlebox. However, if stands are
small, young plants can be removed by physical means before seeds are produced. Because
the root system is not large, pulling plants from wet soil is not difficult. Mulches provide
some control but are not feasible in large areas. Attempts to drown the plant by flooding
are not effective, and the plant is difficult to control in wetlands. During flooding, the lower
part of the stem splits and grows new roots, which help prop up the plant in unstable,
mucky soils. Mowing is effective but usually cannot be accomplished in wet soils where
rattlebox grows. Large plants can be cut, and the stumps treated with herbicides. Physical
removal must be repeated for several years because seeds remaining in the soil seed bank
will continue to sprout. Whatever method of removal is used, it is important to avoid dis-
turbing the site.
Little research has been conducted on chemical control of rattlebox. Use of glyphosate
and triclopyr was not successful in site trials in Florida. Trees sprayed with glyphosate in
California turned yellow, but results were not conclusive.
No biological controls are currently used in the United States. Three insects from
Argentina are being investigated in South Africa. The sesbania flower beetle (Trichapion latri-
vetre) and the sesbania seed weevil (Rhyssomatus marginatus) attack seed pods and seeds,
reducing production as much as 98 and 84 percent, respectively. The sesbania stem borer
(Neodiplogrammus quadrivittatus), also a weevil, damages and kills trees.
530 n SHRUBS

Selected References
Hall, David W., Vernon V. Vandiver, and Jason A. Ferrell. “Hemp Sesbania, Sesbania exaltata (Raf.)
Cory.” Institute of Food and Agricultural Sciences (IFAS), University of Florida. 2009. http://
edis.ifas.ufl.edu/fw039.
Hall, David W., Vernon V. Vandiver, and Brent A. Sellers. “Bagpod (Bladderpod), Sesbania vesicaria
(Jacq.) Ell.” Institute of Food and Agricultural Sciences (IFAS), University of Florida.2009. http://
edis.ifas.ufl.edu/fw038.
Hunter, John C., A. Gerrit, and J. Platenkamp. “The Hunt for Red Sesbania.” Cal EPPC News (Quarterly
newsletter of the California Exotic Pest Plant Council) 11(2): 4–6, 2003. http://www.cal-ipc.org/
resources/news/pdf/cal-ipc_news5109.pdf.
Russell, Alice B, James W. Hardin, Larry Grand, and Angela Fraser. Poisonous Plants of North Carolina.
North Carolina State University, 1997. http://www.ces.ncsu.edu/depts/hort/consumer/poison/
Sesbapu.htm
“Sesbania drummondii (Poisonbean).” Native Plant Database, Lady Bird Johnson Wildflower Center,
2010. http://www.wildflower.org/plants/result.php?id_plant=SEDR.
“Spanish gold, rattlebox, Sesbania punicea.” Center for Aquatic and Invasive Plants, University of
Florida, IFAS, 2009. http://plants.ifas.ufl.edu/node/418.

n Tropical Soda Apple

Also known as: Sodom apple
Scientific name: Solanum viarum
Synonyms: S. chloranthum, S. viridiflorum, S. khasianum var. chatterjeeanum
Family: Potato (Solanaceae)
Native Range. South America, from Brazil south through Paraguay to northeastern
Argentina and Uruguay.
Distribution in the United States. Southeastern states, from Texas east to Florida, and
north to Tennessee and North Carolina. Also in New York State, Pennsylvania, and California.
Description. Tropical soda apple is a perennial shrub that grows 3–6 ft. (1–2 m) tall, and
is usually as broad as it is tall. While the sturdy stems persist in mild winter climates, they
may die back during unfavorable seasons. Deeply lobed leaves, somewhat resembling oak
leaves, grow on distinct petioles. Leaves are alternate, 4–8 in. (10–20 cm) long, and
2–6 in. (5–16 cm) wide. Lobes are broad and pointed. Leaves and stems are covered with
soft, fine hairs and appear velvety except for the prickles. Most parts of the plant, including
leaves, stems, pedicels, petioles, and calyxes, have thick, white to yellowish, straight prickles
or thorns up to 0.4 in. (1 cm) long. Rigid prickles line the midvein and secondary veins on
both surfaces of leaves and are even more concentrated on the petioles.
The root system is extensive. Feeder roots, just a few inches below the soil surface, are
0.25–1 in. (0.6–2.5 cm) in diameter and can extend 3–6 ft. (1–2 m) away from the root
crown. Roots have adventitious buds which sprout into new plants.
Although plants may flower any time of year, most flowering occurs from September
through May. The small flowers with five white petals, curved backwards as is typical in
the potato family, grow in small terminal clusters on the stems below the leaves. Fruit, each
about the size of a golf ball, 1–1.5 in. (2.5–4 cm) in diameter, is light green with dark-green
streaks when immature, resembling a tiny watermelon. As they mature, the smooth skin
turns a dull yellow, and the fruit looks like small apples and smell like apples or plums.
Although smooth, the skin is leathery, surrounding a thin layer of pale green mucilaginous
pulp. The reddish-brown bitter seeds, 0.1 in. (2.5 mm), are also mucilaginous and slightly
flattened.
 TROPICAL SODA APPLE n 531

Related or Similar Species.

Several prickly Solanum species
are found in the southeastern
United States. Two nonnative
species and one species native
to the western United States
are considered invasive.
Although turkey berry, also
known as bushy white sola-
num, gully-bean, Thai eggplant,
devil’s fig, or susumber, was
introduced to Florida more
than 100 years ago, it has been
recognized as invasive only
recently. It is a pantropical
weed with an unknown origin,
probably West Africa, Central
or South America, or Asia.
Turkey berry is found in
Alabama, Florida, Maryland,
Hawai’i, Puerto Rico, and the
Virgin Islands. A prickly tree-
like shrub as tall as 10 ft.
(3 m), it grows in a variety of
habitats, from wetlands to
rocky hillsides. Its leaf shape is
highly variable and may be
evenly lobed, oddly lobed, or
unlobed. Stems and leaves,
which may have either straight
or curved prickles, are covered
with stellate (star-shaped) hairs. Tropical soda apple has rapidly increased its range since it was identified
Flower stalks are covered with in Florida in 1988. (Native range adapted from USDA GRIN and selected
glandular hairs. Petals in the references. Introduced range adapted from USDA PLANTS Database,
terminal flowers are bright Invasive Plant Atlas of the United States, and selected references.)
white and are not recurved.
Turkey berry is grown for its edible yellow fruit the size of grapes. Flower and fruit appear
year-round in tropical and subtropical climates, and seeds are dispersed by birds. Plants
grow into thickets because of sprouts from lateral rhizomes. Turkey berry invades disturbed
areas, such as pastures, roadsides, forest clearings, and damp waste areas. It is also cultivated
in home gardens for its fruit, even though recent studies indicate that it may be carcinogenic
to humans and poisonous to animals.
Wetlands nightshade, also called wetlands soda apple, aquatic soda apple, Tampico soda
apple, or scrambling nightshade, is native to Mexico, Belize, and the West Indies. First
recorded on the Dry Tortugas in 1974, it had spread to mainland Florida by 1983.
Wetlands nightshade is limited to Florida. It grows along rivers and in cypress stands in
either full shade or full sun. The plant has a clambering vine-like habit, growing as tall as
15 ft. (4.5 m) and forming large, tangled stands. The long, narrow leaves are distinctive,
532 n SHRUBS

A. Infestations of shrubs displace forage plants in pastures. (Charles T. Bryson, USDA Agricultural Research
Service, Bugwood.org.) B. Stems and leaf veins have sharp thorns. (Florida Division of Plant Industry Archive,
Florida Department of Agriculture and Consumer Services, Bugwood.org.) C. Five white petals on the flowers
curve backward. (Rebekah D. Wallace, Bugwood.org.) D. The small striped fruit ripens to yellow. (Florida
Division of Plant Industry Archive, Florida Department of Agriculture and Consumer Services, Bugwood.org.)

up to 6.3 in. (16 cm) long and 0.6–2.2 in. (1.5–5.5 cm) wide, with wavy, lobed, margins. All
prickles, whether on stems or on leaves, are curved, not straight. Veins on the upper surface
of leaves have straight hairs, and leaf surfaces and stems are covered with stellate hairs. It is
tolerant of frost and temporary high water. The plant flowers from May to January. Small,
pea-size, red berries, which have no dark markings when green, hang in clusters opposite
the leaves. Dispersal is by seeds and by stem fragments transported in water. New stems
emerge annually from the plant’s base, and plants also root at leaf axils. Wetlands nightshade
prefers areas that are regularly flooded. It has infested approximately 500–750 ac. (200–300
ha) of marshlands, cypress swamps, and other riparian habitat in southwestern Florida.
Wetland nightshade displaces desirable plants, such as pickerelweed and forms dense thick-
ets that block access and waterways for both wildlife and humans.
Silverleaf nightshade, which is also known as tomato weed or white horsenettle, is a per-
ennial shrub native to northeastern Mexico and southwestern United States. Listed as inva-
sive in Florida and many other states, it was probably transported in ballast or as bedding
in livestock railroad cars. Although it may grow to 3 ft. (1 m) tall, the stems die back to
the root system in winter. The dark-green to pale, grayish-green leaves are variable, usually
lance shaped, 1–4 in. (2.5–10 cm) long and 0.4–1 in. (1–2.5 cm) wide. Margins may be
slightly lobed or wavy. All leaves and stems are pubescent, making the plant appear silvery
green. Leaves are covered with stellate hairs, and both stems and leaves have yellow or
brown colored prickles. The inflorescence, a cyme, has 1–7 flowers with bright blue to pur-
ple, or sometimes white, corollas. Fruit is a dehiscent berry 0.4–0.6 in. (1–1.5 cm) in diam-
eter, and one plant may have 40–60 fruits, each with 60–120 tiny seeds. Silverleaf
nightshade is a weed in cultivated land, orchards and pastures, primarily in regions with
low rainfall.
Several other introduced nightshades can be invasive locally. Heartleaf horsenettle is a
forb, while purple African nightshade and shrubby nightshade are shrubs. Climbing night-
shade and orangeberry nightshade can be both shrubby and vine-like.
Several native species may be confused with tropical soda apple. Cockroach berry, also
called Devil’s apple or love-apple, is a shrub found in the southeastern United States,
Texas, and the West Indies. It is distinguishable from tropical soda apple by its smaller red
 TROPICAL SODA APPLE n 533

fruit and stems that are more densely covered with prickles. Carolina horsenettle, a forb
native to the southeastern states, has become a weed of disturbed areas further north and
further west. Both plants and prickles are usually shorter, and flowers are lavender to white.
Western horsenettle, also known as Torrey’s nightshade, is native to Kansas, Arkansas,
and Texas, but now occurs in Florida and southern Georgia. Flowers are purple with a green
center, and it has pale-yellow fruit. Plants are covered with stellate hairs. Buffalobur is native
to the Great Plains but is an occasional weed in the southeastern states. Leaves are pinnatifid
(deeply lobed) and covered with stellate hairs and straight prickles. The very prickly calyx
completely covers the fruit.
Introduction History. The method of arrival of tropical soda apple into the United States
is not known, but seeds were a likely contaminant in seed shipments or in hay. Seeds may
have been in bags of manure used in composting, or attached to someone’s shoes. Plants
were probably present in Florida as early as 1981, before it was identified and collected in
1988. (Reports in the early 1960s are now known to have been misidentifications of cock-
roach berry.) By 1990, plants had infested 24,700 ac. (10,000 ha) in Florida; by 1993,
400,300 ac. (162,000 ha); and by 1995, 1,235,500 ac. (500,000 ha).
Habitat. Tropical soda apple invades agriculture land, riparian areas, recreation areas,
and disturbed sites that are seasonally wet. Soil disturbance, such as disking, trampling by
cattle, rooting by hogs, or even cleaning of ditches enhances establishment of plants. It is
commonly found in damp waste areas, such as pastures, roadsides, and ditch banks, and
in sugarcane fields, rangeland, and citrus groves. It grows in both open and semi-shaded
sites, and prefers sandy soils on poorly drained level land. Although plants can survive very
wet soils, more than three weeks of standing water will kill them. Seeds will sprout, how-
ever, when the water recedes.
Reproduction and Dispersal. Although tropical soda apple’s main reproduction method
is through abundant seed production, it also reproduces vegetatively. Plants flower and pro-
duce fruit all year and may have both mature and immature fruit at the same time. Plants
average 125 fruit, each containing 180–420 seeds, meaning that each plant can produce
approximately 50,000 seeds, of which 70 percent are viable. In delta soils in Mississippi,
8–10 plants can produce approximately one million seeds each year. Seeds are dispersed
locally by livestock, especially cattle, and by wildlife, which are attracted to the sweet-
smelling fruit. Raccoons, deer, feral hogs, and birds eat the fruit and expel the seeds. Seeds
cannot germinate within the fruit, and scarification in a digestive tract promotes germina-
tion. One cow patty can hold 150 seeds, and plants are often concentrated where cattle rest.
Spread to other counties and states is frequently due to the movement of livestock that have
recently ingested tropical soda apple. Seeds are also inadvertently dispersed as contaminants
in hay, grass seed, grass sod, or on machinery. Approximately 20 percent of the seed crop
remains dormant for up to several months. Germination usually occurs during a dry season,
October to May in Florida, but depends on temperature, light, and age of the seed. Older
seeds lose viability, and seeds deeper than 6 in. (15 cm) in soil probably will not germinate.
White seeds are not viable. Plants can grow from seed to maturity in 105 days.
Root segments, branches, and old root crowns may sprout new plants.
Impacts. Tropical soda apple is both a pest in agricultural land and a threat to native eco-
systems. It is a competitive weed in agricultural crops, such as vegetables. The plant is an
alternate host for a number of viruses and pathogens that cause disease in crops, such as
tomatoes, cucumbers, potatoes, and tobacco. It also carries a fungal pathogen (Alternaria sol-
ani) that also causes disease in food crops. Several crop pests, including tobacco and tomato
hornworms, Colorado potato beetle, tobacco budworm, tomato pinworm, green peach
534 n SHRUBS

aphid, silverleaf whitefly, and soybean loper, use tropical soda apple as an alternate host.
Prickles harm both foraging animals and humans who harvest crops.
Plants infest thousands of acres of pasture and lawn in the southeastern United States.
The rapid growth of tropical soda apple lowers carrying the capacity of pastures by displac-
ing forage plants. From 1992 to 1993, infestations in pastureland in Florida doubled, to
388,300 ac. (157,145 ha). Although cattle eat the fruit, they avoid the prickly leaves and
stems. Cattle undergo heat stress when they are forced to remain in sunny sites because
impenetrable, prickly thickets impede their movement to shade.
Costs due to control of tropical soda apple or to crop damage are difficult to assess, but
the plant has the potential to be a major problem in the southern states. In 1994, losses to
the cattle industry in Florida were estimated at more than $11 million. Costs could extend
into the billions of dollars annually.
Infestations of tropical soda apple disrupt ecosystems, displacing native plants and
reducing biodiversity in natural areas. The plant infests oak and palm hammocks and
cypress (Taxodium spp.) island stands in Florida. By creating a physical barrier, the prickly
plants restrict movement of wildlife. Its rapid growth interferes with restoration of phos-
phate mine reclamation. Plants can create a monoculture covering 50 or more acres.
Although eaten by animals, fruit of tropical soda apple contains a glyco-alkaloid called
solasodine, a substance poisonous to humans. A lethal dose is 200 fruit.
Management. Because tropical soda apple spreads quickly to form large patches, early
detection and removal is important. It is also important to prevent fruit and seed develop-
ment and dispersal of seeds. Cattle that may have eaten tropical soda apple fruit should be
held for several days before being shipped elsewhere. All equipment, including vehicles,
mowers, tractors, clothing, and shoes, must be thoroughly cleaned to prevent transport of
seeds. Physical control may be accomplished by hand-pulling small populations. Entire
plants, including fruit, stems, and roots, should be removed and destroyed. Large stands
may be mowed to prevent formation of flowers and fruit.
Chemical applications are effective on new sprouts after plants have been mowed.
Although glyphosate, imazapyr, picloram, dicamba, and triclopyr are all effective, triclopyr
works best.
Because many economically important crops, including peppers, tomatoes, tobacco,
eggplant, and potato, are closely related to tropical soda apple, any use of biological con-
trol must be host-specific to the invasive species of Solanum. Of the numerous agents
investigated, two pathogens and four insects offer potential. Ralstonia solanacearum, a
soil-born bacterium, causes tropical soda apple plants to wilt and die, but it can also attack
crops if they come into contact with contaminated soil, equipment, or irrigation water.
A strain of the tobacco mild green mosaic tobamovirus (TMGMV U2) is lethal to tropical
soda apple. A defoliating leaf beetle (Leptinotarsa texana) and a leaf-feeding tortoise beetle
(Metriona elatior) do not appear to harm Solanum-related crops, but more testing is
required. A weevil (Anthonomus tenebrosus) destroys the contents of flower buds, which
inhibits fruit production. A leaf-feeding beetle from South America (Gratiana boliviana),
which is host-specific to tropical soda apple, was released in 2003. Although a high pop-
ulation of insects is necessary for total control, the beetles affect the plant’s competitive
ability by reducing its vigor and the number of fruit. Small infestations require a release
of 100–300 beetles, while dense stands need 300–500 individuals, all of which will repro-
duce. Because the beetles do not feed in winter, tropical soda apple plants will rebound
where winters are mild.
 YELLOW HIMALAYAN RASPBERRY n 535

Benefits of Tropical Soda Apple

T ropical soda apple has been cultivated in Mexico and India as a source of ste-
roids, made from the solasodine that causes the fruit to be poisonous to
humans. These steroids are useful in cancer treatment, Addison’s disease, rheu-
matic arthritis, and in the production of contraceptives.

Selected References
Coile, Nancy C. “Tropical Soda Apple, Solanum viarum Dunal: The Plant from Hell.” Botany Circular
No. 27, Florida Department of Agriculture and Consumer Services, Division of Plant Industry,
1993; revised 1996. http://www.doacs.state.fl.us/pi/enpp/botany/botcirc/TSA-circ27-1996.pdf.
Cuda, J. P., D. Gandolfo, J. C. Medal, R. Charudattan, and J. J. Mullahey. “Tropical Soda Apple,
Wetland Nightshade, and Turkey Berry.” In Biological Control of Invasive Plants in the Eastern
United States, by R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-04, 2002.
http://www.invasive.org/eastern/biocontrol/23SodaApple.html.
National Biological Information Infrastrucure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Solanum viarum (Shrub).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=265&fr=1&sts=&lang=EN.
Sellers, Brent, Jay Ferrell, J. Jeffrey Mullahey, and Pat Hogue. “Tropical Soda Apple: Biology, Ecology
and Management of a Noxious Weed in Florida.” SS-AGR-77. Agronomy Department, Florida
Cooperative Extensive Service, Institute of Food and Agricultural Sciences, University of Florida,
1993; revised 2009. http://edis.ifas.ufl.edu/pdffiles/UW/UW09700.pdf.

n Yellow Himalayan Raspberry

Also known as: robust blackberry, wild blackberry, wild raspberry, yellow raspberry,
cheeseberry
Scientific name: Rubus ellipticus
Synonyms: Rubus flavus, R. gowreephul
Family: Rose (Rosaceae)
Native Range. Foothills of the Himalaya Mountains, from Kashmir east through Nepal
and southern China, and in tropical and subtropical India and Southeast Asia.
Distribution in the United States. Hawai’i.
Description. Yellow Himalayan raspberry is a robust evergreen shrub with very long
stems or canes that may reach 15 ft. (4.5 m). Berry plants like this are often called brambles
because of the arching and intertwining prickly stems. The hardy, perennial roots grow new
stems each year, increasing the density and size of the stand. Stems, leaves, and inflores-
cences are densely covered with conspicuous prickles and reddish hairs. Some hairs are
stout and curved, while others are fine and straight. Leaves are alternate and palmately com-
pound, on prickly petioles about 1 in. (2.5 cm) long. The three leaflets are thick and hairy,
with sawlike toothed margins. Leaflets are broadly oval, 2–4 in. (5–10 cm), with the termi-
nal leaflet being slightly larger. It is the only raspberry species in Hawai’i with light-green
oval leaves that are blunt and do not terminate in a point. The underside of leaves is lighter
in color than the top, with downy hairs.
536 n SHRUBS

About 20 small white flow-

ers with five petals each are
clustered on compact panicles,
2–5 in. (5–10 cm) long, at the
ends of the stems. The prickly
flowers are pollinated by in-
sects. Fruits are oval or round
yellow berries, about 0.5–1 in.
(1.5–2.5 cm) long. It is the only
yellow fruited raspberry in
Hawai’i.
Related or Similar Species.
Hawai’i has both native and
introduced species of raspberry
and blackberry. Yellow Hima-
layan raspberry is considered
the worst, but other nonnatives
are also considered extremely
invasive. The native Hawaiian
blackberry, or akala, is the only
deciduous raspberry species in
Hawai’i. Growing in mesic to
wet forests and subalpine
woodlands at 2,200–10,000 ft.
(660–3,070 m), it has dark
pink flowers and less prickly,
pale-colored stalks. Another
native blackberry, akalakala, is
rare, with a more localized
distribution.
Several nonnative Rubus spe-
Yellow Himalayan raspberry is one of several Rubus species that are cies have been naturalized in
invasive in Hawai’i. (Native range adapted from USDA GRIN and selected Hawai’i. Himalayan blackberry
references. Introduced range adapted from USDA PLANTS Database, is a sprawling evergreen shrub
Invasive Plant Atlas of the United States, and selected references.) with canes as long as 10 ft.
(3 m). In spite of its name, it is
native to Western Europe. Canes, which can be erect, decumbent, or trailing, are angled
and furrowed, with straight or curved prickles, 0.25–0.4 in. (0.5–1 cm) long. Young canes
are pubescent, but become almost glabrous as they age. The five leaflets in the compound
leaves, glabrous on the upper surface but fuzzy below, are large and broad, with coarsely
toothed margins. Leaves on flowering canes are smaller, with 3–5 leaflets, and all petioles
have hooked prickles. The inflorescence is a large terminal cluster emerging from lower leaf
axils. Peduncles and pedicels are prickly. The white or rose-colored flowers, 0.8–1 in.
(2–2.5 cm) in diameter, have broad petals. Fruit is a typical berry, a cluster of round, black,
succulent drupelets, 0.8 in. (2 cm) long. Because Himalayan blackberry canes will root
where the tips contact soil and suckers may emerge from roots or cuttings, the plant creates
impenetrable thickets on disturbed land and in riparian areas. It is commonly found in pas-
tures, right-of-ways, creek gullies, and stream edges, where it tolerates periodic flooding of
 YELLOW HIMALAYAN RASPBERRY n 537

A. Arching canes can be 15 ft. (4.5 m) long. B. Stems are covered with prickles and stiff hairs. C. The terminal
leaflet is larger than the other two leaflets. D. Small flowers are white. (Forest and Kim Starr.)

both fresh and brackish water. It grows on many types of soil, but requires moisture. Plants
are reported to grow at elevations over 6,000 ft. (1,830 m).
Himalayan blackberry was introduced to North America in 1885, probably for its berries,
and is now widespread in the western states, several midwestern and eastern states, and in
Hawai’i. It spreads locally by vegetative means and long-distance by seed when berries are
eaten by birds or mammals. Thickets may produce 700–130 seeds per sq. ft. (7,000–
13,000 per m2). By dominating range and pasture, Himalayan blackberry brambles shade
out and displace native species. Livestock avoid the prickly plant, and thickets may prevent
even large animals from reaching water. It hinders access for maintenance and recreational
activities, and its thick jumble of branches creates a fire hazard.
Less prickly than yellow Himalayan raspberry, sawtooth blackberry, which is native to
the eastern United States, is the most widespread nonnative raspberry in Mau’i. It is distin-
guishable by its white flowers, black fruits, and pointed leaf tip. It is considered a serious
pest in wet forests and pastures on several islands. Native to tropical Asia, West Indian rasp-
berry is a small prickly shrub 6 ft. (1.8 m) tall, with white flowers and red fruit. It has 5–7
long, pointed leaflets. This plant is found on most islands of Hawai’i, where it displaces
native plants in pastures and impedes passage because of its dense growth. Snowpeaks rasp-
berry, native to India and Southeast Asia, has 5–9 leaflets, which are also pointed. Lavender
flowers and dark-red to black fruit with white hairs distinguish it from yellow Himalayan
raspberry. It occurs on several islands with similar impacts.
Introduction History. Deliberately introduced to the town of Volcano on the island of
Hawai’i for its tasty yellow fruits in the early 1960s, yellow Himalayan raspberry has since
escaped from cultivation. People also like the plant as an ornamental, and in some cases to

Brambles of Himalayan Blackberry

B rambles of Himalayan blackberry may be so dense and intertwined that young

sheep and goats that become entrapped may die from hunger or thirst if not
rescued.
538 n SHRUBS

check soil erosion. Because it is a noxious weed in Hawai’i, it is illegal to transport the plant
between islands. Seeds, however, are unknowingly carried in tree fern trunks, which are
widely shipped from the island of Hawai’i as an ornamental. Plants have been found in sev-
eral places on Mau’i, indicating that more than one arrival to that island was responsible. It is
likely that yellow Himalayan raspberry can be spread to wherever the tree ferns are sold in
the state. Seeds can sprout and plants can be found growing directly on the tree ferns.
Habitat. Yellow Himalayan raspberry tolerates several types of habitat, from semimoist to
moist sites, including wet places in forests and along forest edges, both in full sun and deep
shade. In Hawai’i, it is found in open canopy forests, shady rainforests, pasture or rangeland,
grassland, agricultural areas, and disturbed sites. The raspberry is found primarily on the
wet windward side of Hawai’i and might not be able to tolerate the drier rainshadow sides
of the islands. It is especially invasive to areas damaged by feral pigs (see Volume 1,
Vertebrates, Mammals, Feral pig). The plant survives in deep shade, taking advantage of for-
est openings as they occur. It is especially invasive to pasture and forest near Hawai’i
Volcanoes National Park at 4,000 ft. (1,200 m). It is found growing at elevations ranging
from 2,000 ft. (600 m) to 8,530 ft. (2,600 m), with annual rainfall of 49–275 in. (1,250–
7,000 mm). The plant tolerates some frost.
Reproduction and Dispersal. Clumps of yellow Himalayan raspberry brambles increase
in size by sprouting stems from root suckers and underground shoots, especially after dis-
turbance by cutting or by fire. Each stem lives two years, producing fruit in the second year
and dying two months after flowering. New stands originate from seeds, often when the fruit
is eaten by birds and mammals, including humans, and then passed through the digestive
tract and deposited in a distant location. Outlying, sporadic populations at mid-elevation
forests on the eastern side of the island of Hawai’i are probably due to birds spreading seeds.
Seeds can also be accidentally transported in clothing and camping gear, a significant disper-
sal means in Hawai’i Volcanoes National Park. New plants can also grow from pieces of roots
or cuttings of fresh wood not properly disposed of.
Impacts. Yellow Himalayan raspberry bushes form impenetrable thickets that displace
native vegetation and impede wildlife movement. Arching stems can climb over and smother
other plants. Thickets are often several feet or meters wide, and up to 15 ft. (4.5 m) tall.
Because it grows in many habitats and smothers smaller plants, the raspberry threatens species
diversity in Hawai’i. The native Hawaiian raspberry, akala, has no prickles for protection and is
more likely than yellow Himalayan raspberry to be used as forage by feral sheep, goats, and pigs.
The plant is considered a major threat to the Ola’a Forest Tract of Hawai’i Volcanoes
National Park, especially in areas disturbed by feral pigs. The worst infestation is around
farm lots near the town of Volcano and around the summit of Kilauea, where the region
receives 50–275 in. (1,270–7,000 mm) of rain annually.
Management. Control is not easy. Because of the size of dense thickets and the difficulty
of handling the very prickly stems, it is difficult to impossible to eradicate these raspberry
plants by physical removal. Every piece of the root must be removed and preferably burned
so that it does not have the opportunity to resprout. Physical efforts to eradicate the plant in
the Ola’a Forest Tract failed, largely because of the dense thickets where basal stem diame-
ters may exceed 2 in. (5 cm). Even on smaller plants, it is very difficult to extract all the
roots. Because yellow Himalayan raspberry is especially invasive to areas disturbed by feral
pigs, it is important to first eliminate the pigs.
Chemical applications can be applied in several ways, including foliar, stem injection, cut
stump, or basal stem. Glyphosate or triclopyr, both systemics, are preferred because the her-
bicide is transported to the roots. Foliar sprays are not recommended in dense rainforest
 YELLOW HIMALAYAN RASPBERRY n 539

Beneficial Uses of Yellow Himalayan Raspberry

C ontrol of this invasive shrub is made more difficult because the delicious fruits
are desirable for food. Fruits are also used to create a natural, purplish blue
dye. The inner bark is used as a natural remedy in traditional Tibetan medicine,
and the plant is useful as a renal tonic and an antidiuretic.

conditions because they will kill nontarget plants as well as yellow Himalayan raspberry.
Experiments conducted at the University of Hawai’i at Manoa indicate that four herbicides
(imazapyr, metsulfuron, triclopyr amine, and triclopyr ester) used in cut-stump treatments
killed 95–100 percent of yellow Himalayan raspberry plants within two years. Although
10–15 percent resprouted, about one-half later died. Sprouts on plants treated with metsul-
furon were slightly more likely to survive. Applications of picloram and 2,4-D + triclopyr
killed only 65–75 percent of targeted plants. None of the herbicide applications significantly
affected native plants
The possibility always exists that any introduction of biological controls might adversely
affect native plant species in Hawai’i. Four moths and a sawfly were introduced to control
sawtooth blackberry. Not only were they considered to be ineffective, but one species was
also observed feeding on the native Hawaiian raspberry species. Results of research attempts
specifically to control yellow Himalayan raspberry are not yet available. A possibility exists
in plant rust fungi from the southeastern United States.

Selected References
Benton, Nancy. “Yellow Himalayan Raspberry.” Weeds Gone Wild: Alien Plant Invaders of Natural
Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/
plants/alien/fact/pdf/ruel1.pdf.
Hoshovsky, Marc C. “Rubus discolor.” In Invasive Plants of California’s Wildland, edited by Carla C.
Bossard, John M. Randall, and Marc C. Hoshovsky. Berkeley: University of California Press, 2000.
http://www.cal-ipc.org/ip/management/ipcw/pages/detailreport.cfm@usernumber=71&survey
number=182.php.
McCarney, Perry. “Invasive Alien Species: Yellow Himalayan Raspberry.” Helium Sciences: Biology,
2002–2010. http://www.helium.com/items/984556-invasive-alien-species-yellow-himalayan
-raspberry.
Motooka, P., et al. “Rubus ellipticus.” In Weeds of Hawai’i’s Pastures and Natural Areas: An Identification
and Management Guide. College of Tropical Agriculture and Human Resources, University of
Hawai’i at Manoa, 2003. http://www.ctahr.hawaii.edu/invweed/index.html.
Santos, Gregory L., Linda W. Cuddihy, and Charles P. Stone. Control of Yellow Himalayan Raspberry
(Rubus ellipticus Sm.) with Cut Stump Herbicide Treatments in Hawaii Volcanoes National Park.
Technical Report 80. Cooperative National Park Resources Studies Unit, University of Hawai’i at
Manoa, 1991. https://scholarspace.manoa.hawaii.edu/handle/10125/7167.
Starr, Forest, Kim Starr, and Lloyd Loope. “Rubus ellipticus, Yellow Himalayan Raspberry.”
U.S.Geological Survey, Biological Resources Division. Haleakala Field Station, Mau’i, HI, 2003.
http://www.hear.org/starr/hiplants/reports/pdf/rubus_ellipticus.pdf.
n Trees
n Australian Pine
Also known as: Beach she-oak, casuarina, coast she-oak, ironwood, beefwood,
horsetail tree, scaly bark oak, whistling pine
Scientific name: Casuarina equisetifolia
Synonyms: C. litorea, C. littorea
Family: Casuarina (Casuarinaceae)
Native Range. Australia and neighboring regions, including Southeast Asia, Indonesia,
Borneo, the Philippines, New Guinea, and New Caledonia.
Distribution in the United States. California, Arizona, Texas, Alabama, Florida, Hawai’i,
Puerto Rico, and the Virgin Islands.
Description. Australian pine is an evergreen angiosperm that resembles a conifer because
of its needle-like twigs and cone-like fruits. Although it generally grows 50–80 ft. (15–25 m)
tall, with an open, irregular crown of erect, spreading, or drooping branches, it can reach as
tall as 130 ft. (40 m). Trunk diameter is as much as 1–1.5 ft. (0.3–0.5 m), but those in cop-
piced thickets are much smaller. Smooth on small trunks, the reddish-brown to gray bark
becomes thick, rough, furrowed, and peeling on older trees. What appear to be grayish-
green pine-like needles are narrow jointed twigs, 4–8 in. (10–20 cm) long and less than
0.04 in. (1 mm) wide. The twigs, drooping in clusters, have tiny but distinct ridges running
between the nodes (joints). The twigs separate at the nodes, like a horsetail. The leaves are
tiny pointed scales, 6–8 gathered in whorls that encircle the joints on the twigs. The
needle-like twigs are deciduous, shedding continually all year.
Plants have a shallow fibrous root system that spreads near the surface, but roots can also
penetrate deep into the soil if moisture is available. Under wet conditions, roots will form a
dense mat.
Trees are monoecious, meaning that male and female flowers are on the same tree. Many
tiny male flowers grow in narrow terminal spikes, 0.25–1 in. (0.6–2.5 cm) long, at the end
of the twigs. Female flowers are borne on short stalks in small, round, light-brown clusters
further down along the non-shedding part of the branch. Flowers are wind-pollinated.
Abundant fruit form in brown woody clusters, or fruiting heads, 0.75 in. (2 cm) long and
0.5 in. (1.3 cm) wide, that resemble pine cones. Green fruiting heads are covered with a
mat of white hairs before they ripen and become rusty colored. When the cluster dries, it
releases the tiny, 0.25 (6 mm) one-seeded winged nutlets, called samaras.
Related or Similar Species. Two other Casuarina species, Australian river oak and
Brazilian oak, are also invasive. Both are known by several common names. Australian river
oak, the smallest Casuarina in Florida, 33–50 ft. (10–15 m) tall, has a longer male flower
spike, 0.4–2 in. (1–5 cm), smaller cones, and 8–10 scaly leaves in a whorl. Because it has
no tolerance for salt spray or salt water, it does not grow on beaches. Brazilian oak trees have
no female flowers in Florida. This species may be distinguished by its longer deep green
twigs, smooth gray branches, and 12–16 scaly leaves in a whorl. In Hawai’i, it has shorter
male flower spikes and smaller fruiting heads. Although Australian pine is the only species
 AUSTRALIAN PINE n 541

naturalized in the United States,

it readily hybridizes with these
two species, resulting in trees
with intermediate characteris-
tics that are difficult to identify.
Introduction History. The
U.S. Department of Agriculture
introduced seeds of Australian
pine to Florida in 1898 for
stabilization of ditch and canal
banks and for use as shade,
shelterbelts, hedges, ornamen-
tals, and timber. The tree became
naturalized both in Florida
and in the West Indies before
1920. Populations of Australian
pine rapidly expanded in 1960
after Hurricane Donna created
newly disturbed sites for its
establishment. In Hawai’i, plants
were recorded on Kaua’i in
1882 and on O’ahu in 1895.
Habitat. Australian pine is
predominantly found in tropi-
cal and subtropical climates
in coastal habitats, such as
beaches and estuaries, and in
riparian sites. It is frequent on
disturbed sites, such as filled
wetlands, clearings, and road-
sides, and can occupy poor soils
because it is a nitrogen fixer. Australian pine was used for bank stabilization along canals and ditches
The tree has a wide range of in Florida as early as the late 1800s. (Native range approximated from
environmental tolerances and USDA GRIN and selected references. Introduced range adapted from
can grow on several types of USDA PLANTS Database, Invasive Plant Atlas of the United States, and
substrate, including sand or selected references.)
shell beaches, rocky coasts,
sand bars or sand dunes, and on limestone. It can withstand partial water logging and can
live in slightly saline mangrove habitats, but does not do well in heavy, clay soils. In
Florida, Australian pine is usually found in coastal swamps or in hammocks in the
Everglades, where it is salt tolerant and wind resistant. The mean maximum monthly tem-
perature in its native range is 50–91ºF (10–33ºC), and it is not frost hardy. Average annual
rainfall in its native range is 28–79 in. (700–2,000 mm), often with a six- to eight-month dry
season, but it can grow with as little as 25 in. (640 mm) or as much as 169 in. (4,300 mm) of
annual rainfall. It can also grow in semiarid regions and from sea level to approximately
5,000 ft. (1,500 m) elevation.
In their native Australia, Australian pine grows along the coast, Brazilian oak grows in
swamps, and Australian river oak grows along rivers. In Florida, Australian pine is more
542 n TREES

A. The forest floor under an Australian pine stand is barren of other plants. (Forest and Kim Starr, U.S.
Geological Survey, Bugwood.org.) B. What appear to be needles are twigs with scaly leaves. (Amy Ferriter,
State of Idaho, Bugwood.org.) C. Cone-like fruit form in brown woody clusters. (Forest and Kim Starr, U.S.
Geological Survey, Bugwood.org.)

frequent along beaches and other dry sites, while Brazilian oak, the most frost hardy, is
found along roads and fence lines. Australian river oak is primarily limited to deliberately
planted trees. In Hawai’i, Australian pine is found along the sandy coast and other lowlands.
Reproduction and Dispersal. Except for Brazilian oak, which has no female flowers in
Florida, Casuarina species reproduce both sexually and vegetatively. Although trees in
Hawai’i and Puerto Rico can produce flowers and fruit year-round, most trees in the
United States flower twice a year, February to April and September to October. Fruit
matures in June and December. After reaching 4–5 years of age, each plant produces thou-
sands of winged seeds. Seeds are dispersed by birds, animals, water, and wind. Birds, par-
ticularly exotic parrots and parakeets, eat the seeds, and seeds may also adhere to animals’
feet and fur. The fruiting heads float in rivers and currents, and the wings on the seeds en-
able them to be wind-blown. Although seeds need only 4–8 days to germinate under ideal
conditions, they remain viable for as long as a year. Seedlings, sensitive to fire, drought,
and flooding, grow most rapidly in the first seven years, as much as 6.5 ft. (2 m) in the first
year alone. Growth rates in Florida vary with conditions, ranging from 1.6 ft. (0.5 m) to over
10 ft. (3 m) per year. The plant, however, is intolerant of shade, and seedlings can easily be
outcompeted by grasses and sedges.
When the trunk of Australian pine or Brazilian oak is cut, it will produce many sprouts
from the stump, resulting in a thicket of small trees. Brazilian oak also produces many
new trees from root suckers, even without injury, but Australian pine and Australian river
oak do not.
Impacts. Australian pine alters the habitat, displaces native species, and contributes to
soil erosion. Dense, monospecific thickets alter the light, temperature, soil chemistry, and
hydrology of the habitat, which displaces native dune and beach plant species. Growing
quickly to produce heavy shade, Australian pine also depletes soil moisture and lowers the
water table. The thick litter layer has alleleopathic properties, which inhibit the growth of
other plants.
It offers little to no wildlife habitat, displacing native songbirds except for migrating gold-
finches, which eat the seeds. The twigs and foliage have little food value, and the thick litter
layer covering the ground does not favor insect populations. In the Everglades, fewer
rodents, such as cotton mouse, hispid cotton rat, and marsh rice rat, inhabit groves of
 AUSTRALIAN PINE n 543

Australian pine. Rodents are an important ecological link between plants and predators,
such as raptors and snakes. Australian pine destroys breeding sites of threatened or endan-
gered species in the Everglades, particularly the only remaining nesting sites in the United
States of the American crocodile. Because baby turtles get trapped in the maze of tree roots,
the most productive nesting sites for the green sea turtle and loggerhead turtle are at risk.
The gopher tortoise is also vulnerable.
The trees have a shallow root system, providing little support in storms. Downed
Australian pine trees blocked escape routes in Florida after a 1945 hurricane. The root sys-
tem can also break sewer and water lines and crack pavement. Because Australian pine dis-
places soil-binding vegetation, erosion is more likely to occur. The tall trees do not trap sand
as well as shorter plants, resulting in flatter topography instead of the diversity in habitat
provided by a swell and swale system. The pollen causes significant respiratory allergic
reactions.
Management. Most physical means of removal are ineffective because cutting or damage
to the tree induces sprouting. Small stands may be removed by digging out seedlings and
small trees. Periodic burning of large monospecific stands, combined with herbicide use
on resprouts, may be used for trees greater than 3 in. (8 cm) in diameter. However, fire
can also make the soil too alkaline, rendering it unsuitable for native plant species.
Applied in any method—basal bark, cut stumps, hack-and-squirt, spray, or injection—
chemical treatments are effective. Triclopyr works well on foliage of shoots and saplings.
Glyphosate, dicamba, and picloram are best for cut surfaces.
Although Australian pine is a tree with few insect problems, searches are being conducted
for biological agents. No natural enemies exist in North America. A twig girdler (Oncideres
cingulata) and an Australian pine borer (Chrysobothris tranquebarica) are only effective on
small trees. Both a leaf notcher weevil (Artipus floridanus) and a spittlebug (Clastoptera
undula) do little damage. Tests are being conducted on a defoliating moth (Zauclophora pel-
odes) and a host-specific seed-feeding wasp (Bootanelleus orientalis). Pests that attack trees
in plantation settings or other places in Australian pine’s introduced range throughout the
world also deserve notice. In India, a stem borer kills shoots, and a Rhizoctonia species causes
seedling damping off. A root fungus (Trichosporium visiculosum) causes a blister disease in
India, especially in wet, crowded conditions. A lymantriid moth (Lymantria xylina) is a sig-
nificant pest of both Australian pine and Brazilian oak in China. A root rot is caused by a
fungus (Clitocybe tabescens) in Florida, but it also attacks 210 other species. A stem canker
and dieback is caused by a fungus (Diplodia natalensis) in Puerto Rico. Puerto Rico and
India see few seedlings because of predation of seeds by ants.

Uses of Australian Pine

A ustralian pine is widely used in regions where it has been introduced, espe-
cially in developing countries. Because of its rapid growth, it is often used for
coastal reclamation and erosion control. Although brittle, the wood has a variety
of uses, including oars, paving, wheels, roofing, and boats. The tannin has been
used to cure alligator skins in Florida. In Hawai’i, the hard wood was used for war
clubs and for beaters to pound tapa cloth, but now is used primarily for fuel.
544 n TREES

Selected References
Elfers, Susan C. “ Element Stewardship Abstract, Casuarina equisetifolia.” Global Invasive Species Team,
Nature Conservancy, 1988. http://wiki.bugwood.org/Casuarina_equisetifolia
“Horsetail Casuarina.” In Common Forest Trees of Hawaii (Native and Introduced). Agricultural
Handbook No. 679 by Elbert L. Little Jr. and Roger G. Skolmen. U.S. Department of Agriculture,
Forest Service, 1989. Reprint by College of Tropical Agriculture and Human Resources,
University of Hawai’i at Manoa, 2003.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Casuarina equisetifolia (Tree).” ISSG Global Invasive Species Database, 2010. http://
www.issg.org/database/species/ecology.asp?fr=1&si=365.
“River-oak Casuarina.” In: Common Forest Trees of Hawaii (Native and Introduced). Agricultural
Handbook No. 679 by Elbert L. Little Jr. and Roger G. Skolmen. USDA Forest Service. 1989.
Reprint by College of Tropical Agriculture and Human Resources, University of Hawai’i at Manoa,
2003.
Rockwood, D. L., R. F. Fisher, L. F. Conde, and J. B. Huffman. “Casuarina.” n.d. http://www.na.fs.fed
.us/pubs/silvics_manual/volume_2/casuarina/casaurina.htm.

n Brazilian Peppertree
Also known as: Christmasberry, Florida holly, Pimienta de Brazil, copal, schinus, wilelaiki
Scientific name: Schinus terebinthifolius
Synonyms: Sarcotheca bahiensis, Schinus antiarthriticus, Schinus mellisii, Schinus mucronulatus,
Rhus terebinthifolia
Family: Sumac (Anacardaceae)
Native Range. Southern Paraguay, northeastern Argentina, Uruguay, and southeastern
Brazil.
Distribution in the United States. Widespread use as an ornamental in tropical,
Mediterranean, and desert climates of California, Texas, Florida, Alabama, and Hawai’i, as
well as Puerto Rico and the Virgin Islands. It has not reached the limits of its potential range,
and is invasive in Hawai’i and Florida.
Description. Brazilian peppertree grows either as a small broadleaf evergreen tree, usually
6.5–23 ft. (2–7 m) tall, or as a shrub. The trunk can be as large as 3 ft. (1 m) in diameter, but
hidden by densely intertwining and arching branches. Shrubby trees with multiple trunks
occur when many stems sprout from the root. The slightly rough bark is gray or dark, but
young stems are yellow-green. The resinous sap and crushed leaves are aromatic, with a tur-
pentine or peppery smell. Alternate leaves are pinnately compound, shiny and dark green
above and pale green below. The number and size of leaflets is variable, ranging from as
few as 4 to as many as 13. Leaflets are elliptical and pointed at each end. They can be as small
as 0.6 in. (1.5 cm) or as large as 3 in. (7.5 cm) long and 1.25 in. (3 cm) wide, with the ter-
minal leaflet being the largest. Leaves have a reddish color, often with a reddish midrib, and
may be either entire or finely toothed. Distinctive parallel veins pattern the leaflets from the
midrib.
Trees flower in the fall, September through November, followed by the fruit from
December to February. About 10 percent of populations flower again in spring, March to
May. Flower clusters, 2–3 in. (5–7.5 cm) long, consisting of several small greenish-white
flowers with yellow petals and yellow centers, grow in the leaf axils near the ends of
branches. Although male and female flowers are not distinctively different, they occur on
separate male and female trees. Male flowers last only one day, while female flowers last
 BRAZILIAN PEPPERTREE n 545

up to six days. Flowers are

pollinated by many insects,
especially flies, wasps, and
butterflies. Female trees pro-
duce an abundance of fruit,
which remains ripe on the tree
for eight months. Occurring in
clusters like the flowers, the
fruits are small, 0.16–0.25 in.
(4–6 mm), glossy green berries
which turn red when ripe.
They have a brown pulp. Seeds
are dark brown and tiny, one
in each berry. A pound contains
over 36,500 seeds (80,600
per kg).
Related or Similar Species.
Peruvian peppertree, also called
California peppertree, can be
distinguished by its weeping
foliage and hanging flower
panicles, almost a foot long
(30 cm). The pale yellow flow-
ers are followed by long, hang-
ing clusters of green berries,
first turning red, and then black
as they ripen. The long com-
pound leaves have 20–40 leaf-
lets. The bark is dark brown
and deeply fissured, peeling off
in large flake-like plates. This
species is also becoming inva- Commonly planted as an ornamental or for shade in California and
sive in some areas. southern states, Brazilian peppertree is still expanding its range. (Native
Introduction History. Listed range adapted from USDA GRIN and selected references. Introduced
in seed catalogs as an ornamen- range adapted from USDA PLANTS Database, Invasive Plant Atlas of the
tal in 1832, Brazilian pepper- United States, and selected references.)
tree was imported to Florida
by the 1840s. In the mid-1920s, a gardener gave friends hundreds of seeds, which dispersed
the plant around the state. By the 1950s, the tree was considered an invasive pest. By 1969,
it had reached the Everglades. It is still planted as an ornamental tree in several states.
Habitat. Brazilian peppertree invades disturbed sites, such as fields, ditches, roadsides,
and drained wetlands. It also invades pinelands and grows densely in dry to mesic pastures
and forests, as well as in urban environments. In Florida, trees can dominate some ham-
mocks, pinelands, and mangrove forests. In Hawai’i, it is widespread in mesic to dry envi-
ronments, and can be dense on wastelands. It is very drought resistant and can survive
both fire and high winds. Brazilian peppertree is a pioneer species and does not invade
well-established native ground cover, but once it gains a foothold, the trees grow fast and
outcompete native plants. It is tolerant of partial shade, growing slowly beneath a canopy
546 n TREES

A. The trunk of large trees may be hidden unless low-hanging branches are pruned. (Forest and Kim Starr, U.S.
Geological Survey, Bugwood.org.) B. Flower clusters grow from leaf axils. (James H. Miller, USDA Forest Service,
Bugwood.org.) C. Compound leaves have elliptical leaflets. Fruits are glossy red berries. (Joseph M. DiTomaso,
University of California-Davis, Bugwood.org.)

until some disturbance opens the site to vigorous growth. It is somewhat tolerant of salinity,
invading mangrove regions and low-saline ditches. Although seedlings die in deep water,
mature trees can withstand up to six months of flooding. It is absent, however, from the
parts of the Everglades that are flooded for more than six months. Its northern spread is lim-
ited because it cannot survive cold temperatures.
Reproduction and Dispersal. The invasive nature of Brazilian peppertree is enhanced by
its tremendous seed production, effective dispersal, and high germination rate. Trees mature
in 3 years and can live for more than 30 years. Birds and mammals spread the seeds, and
birds are critical in removing the fruit pulp. Because germination is improved by scarifica-
tion, seeds benefit from the digestive tracts of birds and mammals. Raccoons and opossums
especially feed on Brazilian peppertree seeds, and sometimes their droppings contain hun-
dreds of germinating seeds. Seeds are also dispersed by water. Between 30 and 60 percent
of the seeds are viable, but for only 2–9 months. Most germination takes place in January
and February, soon after the fruit ripens. Between 66 and 100 percent of seedlings survive.
The plant also propagates by resprouting from roots and from cuttings.
Impacts. Brazilian peppertree’s major threat is to biodiversity of both plants and animals.
Capable of invading both terrestrial and seasonally aquatic environments, it grows in dense
thickets that shade out native plants and dominate pine forests. It replaces functioning eco-
systems with dense, monotypic stands, decreasing habitat for native wildlife. Brazilian pep-
pertree has displaced local populations in Florida of federally endangered species, such as
beach clustervine and the endangered sedge, beachstar. Compared with native pinelands,

U.S. Champion

T he largest Brazilian peppertree in the United States, 52.75 in. (134 cm) in diam-
eter at breast height and 35 ft. (10.7 m) tall, is in Florida.
Source: John K. Francis, U.S. Department of Agriculture, Forest Service, International
Institute of Tropical Forestry, San Juan, PR. N.d.
 BRAZILIAN PEPPERTREE n 547

Brazilian peppertree stands have smaller populations and lower diversity of birds. Based on
chemical irritants of its sap and leaves, the tree is suspected of allelopathy, but no field evi-
dence exists to prove so.
Because it is in the same family as poison ivy, poison oak, and poison sumac, contact with
the sap or leaves can cause a rash in persons with sensitivity. Pollen can cause respiratory
problems, as can smoke from burning plants. If eaten in excess, berries and seeds can kill
raccoons, deer, and horses.
Considered one of the most aggressive trees in Florida, Brazilian peppertree infested an
estimated 113,500 ac. (280,000 ha) in central and southern Florida in the mid-1990s. A
negative effect on tourism is expected as Brazilian peppertree replaces more of the natural
ecosystems visitors come to see. Visitors may also choose to avoid areas where the tree is
common, therefore eliminating risk of resin-caused dermatitis.
Management. The best management strategy is to not grow Brazilian peppertree as an
ornamental or allow it to be sold or transferred. Cut trees must be disposed of properly to
prevent inadvertent seed dispersal. Because seeds are viable for a short time and no substan-
tial seed bank remains in the soil, it is possible to totally eradicate small stands. Female trees
can be selectively removed, therefore eliminating seed production. Physical control is pos-
sible if the entire plant, especially the roots, is removed. Even small root pieces, 0.25 in.
(0.6 cm) diameter, can sprout. Fire will kill the aerial parts, but not the roots, which will
produce new shoots. Seeds, however, will not germinate after a fire. Browsing by goats will
control Brazilian peppertree but will not kill it.
Foliar applications of herbicides are an effective chemical control, but coverage is difficult
with large, leafy stands. The technique is better used for seedlings. Triclopyr or glyphosate
leave no soil residuals, so nontarget plants are not harmed unless their leaves are acciden-
tally sprayed. Imazapyr is also good, but because it has more soil residual, it harms nontarget
species. Glyphosate or triclopyr can also be applied to trunks that are freshly cut. Triclopyr,
with an oil additive that enables it to penetrate the bark and take the herbicide down to the
cambium, can be wiped on basal bark with no need to first girdle the tree. This method
requires weeks to take effect, but defoliation and termite activity are clues that it has worked.
Trials with 2,4-D were not effective. Several herbicides, such as bromacil and hexazinone,
which remain in the soil, are absorbed by tree roots and provide long-term effectiveness.
They are slow to act, however, and a problem for nontarget species of hardwoods such as
oaks and maples, which also absorb the herbicides from the soil.
Although more than 200 insects feed on Brazilian peppertree in its native range, no bio-
logical controls for North America are currently known. Three Brazilian species were

Beneficial Uses of Brazilian Peppertree

B ecause it is fire-resistant, Brazilian peppertree has been planted for fire breaks.
Although the wood is too brittle for lumber, it is useful for stakes and posts.
The abundant nectar supplies honey bees, and the foliage and twigs can be
browsed by goats. The bright red berry clusters are used as Christmas decorations.
Dried fruits are used in Brazil as a substitute for black pepper. The bark, when
steeped in a hot bath, is used as a folk remedy in South America to cure rheuma-
tism and back pain. Other folk remedies include use for skin ulcers, bronchitis,
gout, arthritis, diarrhea, and infertility.
548 n TREES

released in Hawai’i—a seed-feeding beetle (Lithraeus atronotatus = Bruchus atronotatus), a

leaf-rolling moth (Episimus utilis), and a stem-galling moth (Crasimorpha infuscata). Two
did only minor damage, and one failed to establish. A species of wasp (Megastigmus transvaa-
lensis) originally from South Africa, was accidentally introduced into both Hawai’i and
Florida, where it was found feeding on Brazilian peppertree fruit. It appears to kill up to
80 percent of the seeds it attacks, giving this wasp good potential for control. A sawfly
(Heteroperreyia hubrichi) which causes defoliation; a thrips (Pseudophilothrips ichini), which
attacks the roots; and other insects are being studied as possibilities.

Selected References
Cuda, J. P., J. C. Medal, and D. H. Habeck. “Classical Biological Control of Brazilian Peppertree (Schinus
terebinthifolius) in Florida.” ENY-820 (IN114). University of Florida Institute of Food and
Agricultural Science (IFAS) Extension, 1999; revised 2009. http://edis.ifas.ufl.edu/in114.
Hight, S. D., J. P. Cuda, and J. C. Medal. “Brazilian Peppertree.” In Biological Control of Invasive Plants in
the Eastern United States, by R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-
04, 2002. http://www.invasive.org/eastern/biocontrol/24BrazilianPeppertree.html.
MacDonald, Greg, Brent Sellers, Ken Langeland, Tina Duperron-Bond, and Eileen Ketterer-Guest.
“Brazilian Pepper-tree, Schinus terebinthifolius.” Excerpted from the University of Florida Institute
of Food and Agricultural Science (IFAS) Extension, Circular 1529, Invasive Species Management
Plans for Florida. Center for Aquatic and Invasive Plants, 2008. http://plants.ifas.ufl.edu/node/405.
Masterson, J. “Schinus terebinthifolius (Brazilian Pepper).” Smithsonian Marine Station at Fort Pierce, FL,
2007. http://www.sms.si.edu/irlspec/Schinus_terebinthifolius.htm.
Motooka, P., et al. “Schinus terebinthifolius.” In Weeds of Hawai’i’s Pastures and Natural Areas; An
Identification and Management Guide. College of Tropical Agriculture and Human Resources,
University of Hawai’i at Manoa, 2003.

n Carrotwood
Also known as: Tuckeroo tree, carrot weed.
Scientific name: Cupaniopsis anacardioides
Synonyms: Cupania anacardioides, Cupania anacardioides var. parvifolia
Family: Soapberry (Sapindaceae)
Native Range. North and east coasts of Australia and New Guinea.
Distribution in the United States. Carrotwood is a popular landscape tree that is very
adaptable and widely planted. It has become naturalized on both east and west coasts of
Florida. In 1996, its distribution paralleled that of mangroves, and inland populations were
just beginning to develop. Also in California, Texas, and Hawai’i.
Description. Carrotwood is a broadleaf evergreen tree, usually with one slender trunk,
which may rapidly grow as tall as 35 ft. (10.5 m). The outer bark is dark gray, but the inner
bark, which gives the tree its name, is orange-colored. The large compound leaves, alternate
on the stem, are evenly pinnate, meaning that the leaf terminates in a pair of leaflets. The
petioles have a swollen base. The 4–12 stalked leaflets are a shiny oblong, yellowish green,
and leathery. They are 8 in. (20 cm) long and 3 in. (7.5 cm) wide, with untoothed but wavy
margins, and tips that are rounded or slightly indented.
From January to March, depending on locality, many inconspicuous flowers grow in
branched clusters from leaf axils. Each cluster is 3–14 in. (7.5–35.5 cm) long. Although
flowers are unisexual, both types occur in each cluster. The small flowers, with five white
to greenish petals less than 0.1 in. (3 mm) long, grow on short pedicels. Male flowers have
 CARROTWOOD n 549

eight stamems. Fruit, on a short-

stalk, is a woody capsule, about
1 in. (2.5 cm) in diameter, with
three distinctly ridged segments
or lobes. Both flowers and fruit
are minutely pubescent. The
capsule ripens to yellow in
April to June, before drying to a
brown color and splitting to
reveal the seeds. Three shiny
black seeds in each fruit are
covered by a red-orange aril, or
fleshy tissue.
Related or Similar Species.
The unusual fruit and color
of the wood make this tree
distinctive.
Introduction History. Carrot-
wood was brought to St. Lucie
County in eastern Florida in
1955 as a cultivated plant. In
1968, a separate commercial
introduction placed the plant
into large-scale propagation as
an ornamental in Sarasota. It was
a popular landscape plant in the
late 1970s and the early 1980s,
but by 1990, seedlings were
found in various natural habitats.
Carrotwood was recommended
as a street tree in Honolulu.
Habitat. Carrotwood is a very Especially popular as an ornamental tree in Florida, carrotwood is
durable urban tree that is used in tolerant of saline conditions and poor soils. (Native range adapted from
private gardens and commercial USDA GRIN and selected references. Introduced range adapted from
landscaping. It occupies dis- USDA PLANTS Database, Invasive Plant Atlas of the United States, and
turbed and undisturbed sites in selected references.)
both herbaceous and scrub com-
munities. Growing in both sun and shade, it can tolerate conditions of salt, nutrient-poor soils,
and poor drainage. Although most common in wet locations, such as freshwater marshes and
riverbanks, it has a wide range of tolerance to soil moisture and can also be found in dry sites,
such as hammocks and pinelands. Its tolerance of salinity enables it to invade coastal habitats,
including mangrove and cypress swamps, beach dunes, and coastal strands.
Cold temperatures, approximately 22ºF (-6ºC), may be the limiting factor in the potential
northward distribution of carrotwood. Although used as an ornamental plant in California,
no reports have indicated that carrotwood is becoming naturalized, probably because of
California’s drier climate.
Reproduction and Dispersal. Carrotwood reproduces both sexually and vegetatively, but
it spreads primarily by seed. Flowers are most likely pollinated by bees. Plants seed freely,
550 n TREES

A. Carrotwood trees usually have a single trunk. B. Large compound leaves are leathery, with shiny oblong
leaflets. C. Fruits are woody capsules. D. Each capsule has three compartments or segments. E. The hard seeds
have a distinctive red aril. (Forest and Kim Starr.)

and the seeds are attractive to birds. Populations beneath telephone poles and lines, as well
as island populations, can be attributed to birds. Fish Crows in Florida are especially impor-
tant in carrying seeds from inland to coastal island habitats. Clumps of seedlings, however,
are more likely the result of small mammals, which gather the fruit. Seeds are also distrib-
uted by water.
When the top part of the tree is damaged or cut, multiple sprouts emerge from the stump.
If left unpruned, the result is a multi-trunked tree.
Impacts. Carrotwood is especially a problem in low, moist areas and a significant pest in
mangrove habitats, which are important biological resource areas. Mangrove swamps are
critical sites for wading and diving birds and provide protected nursery habitat for commer-
cial and sport fish, crabs and other crustaceans, and invertebrates. These important habitats,
as well as coastal hammocks, are in jeopardy because they are also removed from the natural
landscape by development and by tropical storms and hurricanes. Carrotwood can develop
monotypic stands in mangrove and coastal hammock sites, with densities of more than two
plants per square foot (20 per m2).
In Hawai’i, carrotwood grows in dense monocultures, crowding and outcompeting native
plants for light and nutrients. Carrotwood also grows with other invasive plants in Hawai’i.
Management. Because carrotwood is a common landscape plant, many management sug-
gestions are directed to private homeowners. A major goal is to prevent distribution or dis-
persal of seeds. Physical prevention of seed development includes removing mature
landscape plants. After cutting close to the ground, the stump should be treated with herbi-
cides to prevent resprouting. Dead trees may be left in place to decay.
Chemical treatments, which include triclopyr or glyphosate, are the most commonly used
and most effective controls of carrotwood. Triclopyr is the best application for basal bark or
to cut stumps. Glyphosate has marginal effectiveness, and treatments usually need to be
repeated. Any applications of herbicides in mangrove swamps or other wetlands must be
made with care because of the sensitivity of those water environments.
No biological controls are known for carrotwood.
 CHINABERRY n 551

Selected References
Langeland, K. A. “Natural Area Weeds: Carrotwood (Cupaniopsis anacardiodes).” SS-AGR-165.
University of Florida Institute of Food and Agricultural Sciences (IFAS) Extension, 2001; revised
2009. http://edis.ifas.ufl.edu/ag111.
Lockhart, Chris. “Carrotwood (Cupaniopsis anacardioides).” Weeds Gone Wild: Alien Plant Invaders of
Natural Areas. Plant Conservation Alliance, Alien Plant Working Group, 2009. http://
www.nps.gov/plants/alien/fact/cuan1.htm.
National Biological Information Infrastructure (NBII) and IUCN/SSC Invasive Species Specialist Group
(ISSG). “Cupaniopsis anacardioides (Tree).” ISSG Global Invasive Species Database, 2005. http://
www.issg.org/database/species/ecology.asp?si=641.

n Chinaberry
Also known as: Pride of India, umbrellatree, Persian lilac, Indian lilac, white cedar,
China tree, bead tree, ‘inia (Hawai’i), alelaila or pasilla (Puerto Rico)
Scientific name: Melia azedarach
Synonyms: Melia japonica, Azedarach amena, A. sempervirens, A. vulgaris, Melia angustifolia,
M. australasica, M. floribunda, and others.
Family: Mahogany (Meliaceae)
Native Range. Temperate and tropical southeastern China, possibly India and Southeast
Asia. Naturalized in Japan, India, Sri Lanka, Indonesia, Papua New Guinea, northern
Australia, and the Solomon Islands.
Distribution in the United States. Southern half of the United States, from California to
Florida, north to Utah, Missouri, Virginia, and New York. Also in Hawai’i, Puerto Rico,
and the Virgin Islands.
Description. Chinaberry is a fast-growing deciduous tree usually less than 30 ft. (10 m)
tall, although some plants reach 50 ft. (15 m) in height. Its rounded crown may cover an
area 20 ft. (6 m) in diameter, making it a desirable shade tree. Although a single trunk
may be as thick as 2 ft. (0.6 m) in diameter, the tree often has many smaller trunks because
it readily sprouts from its roots. Stout stems and twigs, ranging in color from glossy olive-
green to brown or slightly purplish red, are covered with light brown spots. Leaf scars from
dropped leaves are three-lobed and prominent. The bark, either dark chocolate or reddish
brown, is also covered with light-brown spots, lenticels, and becomes fissured with age.
The wood is soft and white. The bipinnately, or sometimes tripinnately, compound leaves,
1–2 ft. (0.3–0.6 m) long and 9–16 in. (23–41 cm) wide, give the tree a lacy appearance.
Leaves are alternate, on long petioles, and either glabrous or covered with very fine hairs.
They emit a musky odor when crushed. Toothed leaflets are lance-shaped, 1–3 in.
(2.5–8 cm) long and 0.5–1.2 in. (1.3–3 cm) wide, tapering towards the tip. The upper sur-
face of leaflets is dark green, with a sparsely hairy light-green midvein. The lower surface is a
lighter green with no hairs. Leaves turn golden yellow in the fall. The root system is shallow,
within the top 28 in. (70 cm) of the soil.
In early spring, from March through May, loose drooping clusters of small fragrant flow-
ers emerge in leaf axils on the new growth toward the ends of the branches. Each star-
shaped flower, with five narrow pink or lavender petals around a central dark tube, is about
0.75 in. (2 cm) in diameter. Round yellowish-tan berries, about the same size as the flowers
and also hanging in clusters, are initially mucilaginous and sticky, but become leathery,
552 n TREES

brown seed capsules when

mature. The hard berries, each
containing 1–6 seeds, remain
on the tree after the leaves fall.
Related or Similar Species.
Texas umbrella chinaberry, also
called Texas umbrella tree, is
characterized by upward arch-
ing branches and drooping foli-
age that resembles an umbrella.
This variety is often pollarded
(limbs cut back to trunk), caus-
ing a dense cluster of branches
to sprout from the top of the
pruned tree. Common elder-
berry, a native shrub, can be
distinguished by its white flow-
ers and dark purple berries.
Introduction History. The
precise time period when
Chinaberry was introduced to
the United States is not clear. It
may have been brought to the
United States in the late 1700s
by a French botanist, or intro-
duced to Charleston, South
Carolina, or Georgia in the
mid-1800s. It has been a popu-
lar ornamental shade tree in
southern states for over
200 years, where it is believed
Chinaberry has been a popular shade tree in the southern states for over to bring good luck. It was intro-
200 years. (Native range adapted from USDA GRIN and selected duced as an ornamental to
references. Introduced range adapted from USDA PLANTS Database, Hawai’i in 1840. The tree is still
Invasive Plant Atlas of the United States, and selected references.) sold in nurseries.
Habitat. Chinaberry can
grow in a variety of habitats, but is most invasive in riparian zones or disturbed sites.
Usually found below 1,000 ft. (300 m) elevation, it is reported to grow as high as 9,000 ft.
(2,700 m) elevation in Hawai’i. The tree most often invades disturbed areas, such as road
right-of-ways, fencerows, forest margins, and open areas, or after a site has been burned,
cleared, or mowed. However, it is also found in relatively undisturbed sites, such as flood-
plain hammocks, marshes, and upland woods and grasslands. It is frequently found in open
pine forests in South Carolina when fires are suppressed and after eradication of other inva-
sive species, such as kudzu. It is often present around rural home sites, both occupied and
abandoned, in the southeastern states. In Hawai’i, Chinaberry invades dry pastures and both
dry and moist forests. It tolerates a wide range of soil moisture, as well as heat, drought, and
poor soils. It grows most vigorously in full sun and is only somewhat shade tolerant.
Although predominantly a tropical or subtropical species, the tree tolerates some frost.
 CHINABERRY n 553

A. Trees have a rounded crown. (Chuck Bargeron, University of Georgia, Bugwood.org.) B. Compound leaves
give the tree a lacy look. (Karan A. Rawlins, University of Georgia, Bugwood.org.) C. Star-shaped flowers are
small. (Chris Evans, River to River CWMA, Bugwood.org.) D. Old bark becomes fissured. (Karan A. Rawlins,
University of Georgia, Bugwood.org.) E. Berries hang in clusters. (Cheryl McCormick, University of Florida,
Bugwood.org.) F. Bark on twigs and young trunks is speckled with lenticels. (James H. Miller, USDA Forest
Service, Bugwood.org.)

Reproduction and Dispersal. Chinaberry reproduces and spreads both by seed and by
vegetative means. Plants produce abundant seed, which can be carried long distances by
birds or downstream by water in riparian areas. Because seeds require no dormancy period
and can germinate as soon as they mature, seedlings are numerous. Seeds can survive severe
desiccation and remain viable for at least two years. Colonies of trees sprouting from root-
stocks can create dense, monotypic thickets.
Impacts. Plants grow rapidly from numerous root sprouts to create dense thickets that
crowd out and shade out native species. Plants can reach a height of 19.5–26 ft. (6–8 m)
in only 4–5 years. By creating monocultures, Chinaberry decreases the biodiversity in native
ecosystems. Because Chinaberry is resistant to insects and pathogens, it is able to outcom-
pete native plants. The plant also has alleleopathic properties. The leaf litter changes the
chemistry of the soil, reducing the aluminum levels and making the soil more alkaline.
Leaf litter can also increase the available nitrogen, comparable to nitrogen-fixing legumes.
The wood is brittle, and large trees are prone to dropping limbs, posing a hazard to people
and structures.
All parts of the plant, especially the fruit, are poisonous to humans, some livestock, and
small mammals, including cats and dogs. Symptoms, which appear a few hours after inges-
tion, include stomach irritation, vomiting, diarrhea, breathing difficulty, and weakness or
paralysis. Cattle and some birds usually eat the fruit without harm. However, ingestion of
too many seeds may paralyze birds. Bees and butterflies do not use the flowers.
Management. Maintenance of a healthy ecosystem, combined with early detection and
control, will prevent Chinaberry invasions in natural areas. Where plants have become
firmly established, total eradication is unlikely. The best management for existing
Chinaberry trees is to limit or prevent seed production. Sites should be monitored for new
growth and potential seed production for 3–5 years. Chinaberry can be identified
on color-infrared aerial photographs, a factor that is useful for documenting its spread
or for monitoring control efforts. Physical methods of controlling chinaberry, such as
554 n TREES

Uses of Chinaberry

T he hard berries of Chinaberry trees are used for beads and in rosaries. Fruit and
leaves can be used like mothballs to prevent insect damage to clothing or books.
When stored with food, leaves act as an insect repellant. Extracts of the bark and
fruit have medicinal properties and have been used for many purposes, such as an
antiseptic, an emetic, a diuretic, and an astringent. Parts of the plant are reputed
to reduce fevers, treat rheumatism, and kill parasitic roundworms. A peptide from
leaf tissue has been shown to be effective against the herpes simplex virus.

hand-pulling, are only effective on very small seedlings. Because plants vigorously resprout
from root fragments, all parts of the root must be removed. Cutting or mowing down trees
will prevent development of seeds, but herbicides should be applied to the sprouts to pre-
vent formation of thickets of trees.
Chemical applications are the most effective means of controlling Chinaberry. Seedlings
and small trees can be killed by spraying foliage with triclopyr, glyphosate, or imazapyr, but
root systems of larger trees will not be affected. Foliar sprays are best done in summer and fall.
Spraying trees taller than 20 ft. (6 m) or large stands is not practical because of the amount of
foliage to be covered. Herbicides, such as imazapyr, picloram, or triclopyr, applied to cut
stumps or girdled trunks are the best approach for large trees. This technique does best in late
summer and fall, when the trees are translocating nutrients to the roots. Basal bark or cut
stump application of triclopyr has proven to be most effective method of eradication.
Because Chinaberry is practically insect and disease free, no biological controls are
known.

Selected References
Batcher, Michael S. “Element Stewardship Abstract, Melia azedarach, Chinaberry, Umbrella Tree.”
Global Invasive Species Team, Nature Conservancy, 2000. http://www.imapinvasives.org/GIST/
ESA/esapages/documnts/meliaze.pdf.
Motooka, P., L. Castro, D. Nelson, G. Nagai, and L. Ching. “Melia azedarach.” Weeds of Hawai’i’s
Pastures and Natural Areas: An Identification and Management Guide. College of Tropical Agriculture
and Human Resources, University of Hawai’i, Manoa, 2003. http://www.ctahr.hawaii.edu/inv-
weed/WeedsHI/W_Melia_azedarach.pdf.
Reemts, Charlotte. “Chinaberry (Melia azedarach).” Weeds Gone Wild: Alien Plant Invaders of Natural
Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/
alien/fact/meaz1.htm.
Scheper, Jack. “Melia azedarach.” Floridata, 1999; updated 2004. http://www.floridata.com/ref/M/
meli_aze.cfm.

n Fire Tree
Also known as: Firetree, fayatree, faya bush, candleberry myrtle, firebush
Scientific name: Morella faya
Synonyms: Myrica faya
Family: Sweet Gale (Myricaceae)
 FIRE TREE n 555

Native Range. Azores, Ma-

deira Islands, Canary Islands.
Distribution in the United
States. All major islands in
Hawai’i. In the late 1980s, it
covered 100,000 ac. (40,470
ha), and it continues to expand
its range.
Description. Fire tree is an
evergreen shrub or small tree,
usually growing 12–26 ft.
(3.7–8 m) tall, although it has
been documented at 50 ft.
(15 m). Stems and branches
are covered with hairs. The
dark green leaves are oblanceo-
late, 2–4 in. (5–10 cm) long
and slightly less than an inch
(2–2.5 cm) wide They are wider
toward the tip than at the base.
Alternate on the stem, the aro-
matic leaves are leathery, shiny,
and smooth, but covered with
inconspicuous brownish glan-
dular dots. Leaf margins are
entire or with shallow serra-
tions toward the apex.
The species is subdioecious,
meaning that both male and
female trees have a few flowers
of the opposite sex. Tiny flow-
ers develop among the current As a nitrogen fixer, fire tree colonizes nutrient-poor cinder and ash
year’s growth of leaves. Flowers deposits on Hawaiian volcanoes, altering the successional process.
have no calyx or corolla and (Native range adapted from USDA GRIN and selected references.
grow in many-branched cat- Introduced range adapted from USDA PLANTS Database, Invasive Plant
kins. Catkins with yellow- Atlas of the United States, and selected references.)
green male flowers, each with four pinkish stamens, occur in small, hanging clusters near
the ends of branches. Female flowers, hanging in clusters of three, grow further down on
the branches. Plants produce an abundance of small edible fruit, bumpy looking like berries
about 0.25 in. (6 mm) in diameter. Fruit is red to blackish-purple when it ripens in June.
Related or Similar Species. None in Hawai’i.
Introduction History. Fire tree was introduced to Hawai’i in the late 1800s by Portuguese
immigrants, probably as an ornamental, for medicinal purposes, or for firewood. They also
made a wine from the berries. In the early 1900s, trees were planted in reforestation proj-
ects, but by 1937, the species was recognized as invasive. The first eradication attempts fol-
lowed in 1944. Fire tree was first discovered in Hawai’i Volcanoes National Park in 1961. By
1977, plants covered approximately 30,000 ac. (12,200 ha) in that park, and approximately
85,000 ac. (34,400 ha) on the island group.
556 n TREES

A. Trees usually grow 25 ft. (7.5 m) tall. B. Leathery leaves are shiny and smooth. C. Male catkins grow near the
ends of branches. D. Fruit are bumpy looking berries. (Forest and Kim Starr.)

Habitat. Occupying a wide variety of habitats, fire tree invades various types of native for-
est, shrublands, agricultural areas, pastures, roadsides, and steep slopes. Because it is a
nitrogen fixer, fire tree is capable of invading newly developed, nutrient-poor volcanic cin-
ders and ash. It grows on recent cinder deposits, thin ash over lava, as well as on well-
developed silty, clay, or loam soils. It forms dense, monotypic stands in both wet and mesic
forests. The most affected ecosystem is the early successional stage of the dry seasonal sub-
montane rainforest. The main infestation in Hawai’i Volcanoes National Park occurs at
4,100 ft. (1,250 m).
Reproduction and Dispersal. Fire tree reproduces solely by seed, which is dispersed
when primarily exotic birds, especially the nonnative Japanese Silver-eye, and feral pigs
(see Volume 1, Vertebrates, Mammals, Feral pig) eat the fruit. Trees reproduce early in life,
produce an abundance of fruit and seeds, and grow rapidly. The average adult plant produ-
ces 40,000–400,000 fruits per year. In one year, 21 original adult trees can expand the infes-
tation to more than 150 mature trees. Seeds remain viable in the soil for a long time. Flowers
are predominantly wind-pollinated but also by introduced honey bees.
Impacts. By creating monospecific stands with a dense canopy that shades out understory
herbaceous plants, fire tree interferes with and prevents natural succession. It invades early
succession sites on lava flows and ash, where the dominant vegetation is evergreen ohia and
koa trees, along with native tree ferns. Ohia is a primary pioneer plant on recently deposited
cinders and ash, which are nitrogen poor. The open-canopied vegetation in ohia forests,
because of the variable substrate, provides good habitat for the initial growth of fire tree.
Birds that perch on ohia disperse fire tree seeds in their droppings. In sites that have both
part shade and part sun, seeds germinate easily and grow rapidly. Because those sites are
deficient in nitrogen, any species, such as fire tree, that can fix nitrogen has a competitive ad-
vantage. Fire tree quadruples the normal amount of nitrogen in the soil, enabling itself and
other invaders to grow better and faster. As the fire tree stands develop a dense canopy, all
understory plants are shaded out, halting the successional process. The amount of leaf litter
also inhibits germination of native plants.
Management. No long-term control methods have been discovered. Because seeds are
spread by nonnative birds and animals, keeping populations of those species under control
will limit fire tree invasions. Although physical means of removal are feasible in crop areas,
 MELALEUCA n 557

they are generally precluded in natural areas because of the destruction that would occur to
the environment. Goats can browse fire tree, but goats pose other problems to Hawai’i’s
indigenous plant life.
Even though it is expensive and labor intensive, chemical control has received the most
attention in research. Undiluted glyphosate is the most effective treatment. Direct injection
of the herbicide into the tree trunk protects nontarget plants that may be affected by sprays.
The search for biological control organisms began in 1955, but because the most promis-
ing insects also attacked mango and avocado trees, the effort was abandoned. Botrytis cin-
erea, a pathogen native to Hawai’i, causes the fruit to rot, both reducing seed viability and
making fruit less attractive to birds that disperse seeds. Adults and larvae of fruit-eating
insects carry the infection from tree to tree. Fruit-eating insects, such as Amorbia emigratella
and Cryptoblabes gnidiella, can be reared in laboratories, inoculated with the pathogen, and
released in fire tree areas. Using native species is preferable to introducing new organisms
to the islands.
A moth (Caloptilia sp. nr. schinella), from the Azores and Madeira Islands, was released in
1991 in Hawaiian fire tree stands. Although not host-specific, the two-spotted leafhopper
(Sophonia rufostachia) has killed fire tree on the island of Hawai’i. A butterfly
(Phyllonorycter myricae) is under investigation. A fungal leaf pathogen (Septoria hodgesii sp.
nov) that attacks southern wax myrtle in the southeastern United States, may also be a
possibility.

Selected References
Benton, Nancy. “Fire Tree.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/
mofa1.htm.
“Fire Tree, Firetree, Faya Bush (Morella faya).” Stopping the Silent Invasion. Hawaii Invasive Species
Partnership, 2008. http://www.hawaiiinvasivespecies.org/pests/firetree.html.
IUCN/SSC Invasive Species Specialist Group (ISSG). “Morella faya (Tree, Shrub).” ISSG Global Invasive
Species Database, 2006. http://www.invasivespecies.net/database/species/ecology.asp?si=100.
Seibold, Ryan. “Controlling Fire Tree (Myrica faya) in Hawaii.” Restoration and Reclamation Review.
Student On-Line Journal (Hort 5015/5071). University of Minnesota, Department of Horticultural
Science, St. Paul, 2001. http://purl.umn.edu/59731.

n Melaleuca
Also known as: Paperbark tree, punk tree, white bottlebrush tree, paperbark tea tree
Scientific name: Melaleuca quinquenervia
Synonyms: Melaleuca leucadendron, M. viridiflora var. angustifolia, M. viridiflora var. rubiflora,
Metrosideros quinquenervia
Family: Myrtle or Eucalyptus (Myrtaceae)
Native Range. Eastern Australia, New Guinea, and New Caledonia. Melaleuca is native to
a narrow zone, 25 mi. (40 km) wide, of seasonally flooded coastal lowlands in northeastern
Australia. Trees are commonly used for park plantings in Australia, and because develop-
ment threatens it in its native habitat, efforts have been made to preserve melaleuca in its
native range.
558 n TREES

Distribution in the United

States. Widespread in southern
Florida, concentrated south-
ward of Lake Okeechobee and
on both coasts, where it is inva-
sive to several million acres.
Also in southern California,
southern Texas, Louisiana, and
Hawai’i.
Description. Melaleuca is a
slim, erect evergreen tree grow-
ing up to 80 ft. (24 m) tall,
occasionally to 108 ft. (33 m),
with a slender, non-spreading
crown. Branches, growing up-
ward, not outward, are irregu-
larly spaced off the trunk. The
white bark is spongy or corky
and paperlike, prone to peeling
in layers similar to a paper birch
tree. The gray-green leaves are
alternate and lance-shaped, as
much as 4–6 in. (10–15 cm)
long, but narrow. Five parallel
longitudinal leaf veins are
prominent. When mature, the
leaves are flat, stiff, and leath-
ery. Because of the oily content,
the leaves release a camphor-
like odor when crushed.
Melaleuca has both a net-
Melaleuca is a serious invader in the Everglades, where it was originally work of roots near the soil sur-
planted to create forests. (Native range adapted from USDA GRIN and face and many vertical roots.
selected references. Introduced range adapted from USDA PLANTS The vertical roots tap the water
Database, Invasive Plant Atlas of the United States, and selected references.) table, allowing the plant to sur-
vive dry conditions.
Although individual creamy white flowers are small, they occur in showy bottle-brush
like spikes at the ends of the branches. The spiky inflorescences are 0.75–2 in. (2–5 cm)
long. Each flower has five petals and 30–50 stamens. Because the flower stem is indetermi-
nate, it continues to grow after flowering. The fruit is a small, woody seed capsule that
resembles a cylindrical or squarish button. Each flower spike can produce 30–70 seed cap-
sules, and each capsule contains 200–350 tiny seeds. Closed seed capsules are stored on the
tree, and one stem or branch may have up to 10 clusters of seed capsules alternating with
leaves because the stem continued to grow and reflower several times.
Related or Similar Species. The genus Melaleuca has many species in its native Australia,
but no others are considered invasive to the United States. The crimson bottlebrush plant, a
common ornamental shrub in the same family as melaleuca, has similar flowers, but they are
distinctly red.
 MELALEUCA n 559

A. Melaleuca trees are slender and straight with a compact crown. (Forest and Kim Starr, U.S. Geological Survey,
Bugwood.org.) B. Bark peels off in layers. (David Nance, USDA Agricultural Research Service, Bugwood.org.)
C. The long narrow leaves are stiff and leathery. (Amy Ferriter, State of Idaho, Bugwood.org.) D. Woody fruit
capsules remain on the tree. (Albert [Bud] Mayfield, Florida Department of Agriculture and Consumer Services,
Bugwood.org.) E. Bottle-brush spikes of flowers grow at ends of branches. (Amy Ferriter, State of Idaho,
Bugwood.org.)

Introduction History. Melaleuca was deliberately introduced to southern Florida as an

ornamental in the late 1800s or early 1900s because of its showy flowers. It was also used
as a means of reclaiming, or drying out, swampland and was planted for windbreaks and
fencerows. In the 1930s, trees were planted for erosion control on canal levees, the south
shore of Lake Okeechobee, and in Big Cypress National Preserve. Seeds were deliberately
scattered over the Everglades to create forests. The trees remain concentrated around the
regions where they were initially introduced. As late as 1970, melaleuca was still considered
a good landscape tree.
Over a million trees were planted for erosion control in Hawai’i.
Habitat. A subtropical tree, melaleuca grows primarily in wet or swampy sites and grows
best in wet areas with frequent fires. It invades several natural habitats, such as sawgrass
marshes, wet prairies, lake margins, pine flatwoods, and cypress stands. Plants do best in
seasonally wet sites, but they can also grow in both standing water and in drained uplands.
Melaleuca will also take over disturbed areas such as roadsides, ditch banks, farmland, and
urban environments, but disturbance is not necessary for invasion. Wet sites are more likely
to be invaded. It is tolerant of brackish water, growing in mangrove swamps in both its
native range and where introduced. Frost limits the northward expansion of melaleuca.
Mature trees are able to survive freezing temperatures, but young trees are susceptible.
Melaleuca grows from sea level to 4,500 ft. (1,400 m) elevation in Hawai’i, on all types of
soils. However, it is not as invasive on the islands because its preferred habitat of wetlands is
less common. Fewer wildfires also limit its competitive advantage.
Reproduction and Dispersal. The reproductive advantage of melaleuca lies in its abun-
dant seed production. Saplings that are three years old, and sometimes only one year old,
will flower and set seed. November to January is the normal bloom season in Florida, but
both flowering and growing seasons will be longer during especially wet years. Because mel-
aleuca can flower up to five times per year, a mature tree can produce a million seeds each
year, with more than 50 million stored on the plant in woody capsules. Seed pods must
dry before releasing seeds, primarily after stress or disturbance such as frost, fire, physical
560 n TREES

injury such as a cut limb, or herbicide application. However, some seeds are released con-
tinually, resulting in a steady but light seed fall. Only 10–20 percent of seeds, most of which
remain within the capsules for at least 10 years, are viable. Viability of seeds that are released
from the capsules is reduced by one-half after eight months in the ground, while others
remain dormant for up to two years. Seeds are dispersed by wind and water. Germination
takes place a few days after soil becomes moistened. Seeds are able to germinate in both
sun and shade and even while submerged.
Trees also reproduce vegetatively, having the ability to resprout from roots or stumps
after a fire.
Impacts. Melaleuca affects not only ecosystems, but also human health. It outcompetes
and displaces native species, eliminates wildlife habitat, and poses a fire hazard. Grass and
sedge marshes are the habitats most affected in southern Florida. Melaleuca shades out
and replaces native vegetation such as sawgrass. Trees frequently form impenetrable thickets
that eliminate all other vegetation, resulting in a monoculture that is little used by native
wildlife. The Everglades is a watery grassland environment, but when melaleuca trees
replace the grass, the area becomes an alien environment to native plants and animals,
thereby limiting diversity in the ecosystem. The trees grow very quickly, 3.3–6.5 ft.
(1–2 m) per year, and a stand can form a grove up to 600 ft. (180 m) in diameter in only
one year. In only 25 years, the species can increase its cover in one square mile (2.6 km2)
from 5 to 95 per cent. In densely occupied areas, as many as 31,000 trees and saplings
can grow on one acre (0.4 ha) of land.
The trees have more biomass than native grasses, providing more fuel for fires. Ground
fires burn hotter and can ignite organic soils. The volatile oils in the leaves cause intense
crown fires, which threaten both buildings and native trees, incurring higher firefighting
costs. Accumulations of leaf litter often build hammocks, or tree islands, above the sawgrass
marsh, perhaps altering hydrology and flow of water.
Pollen from melaleuca is believed to be a mild allergen, worsened by the plant’s propen-
sity to flower several times a year, causing respiratory problems for many Florida
inhabitants.
Along with the Brazilian peppertree (see Trees, Brazilian Peppertree), melaleuca is recog-
nized internationally as the greatest threat to the Everglades ecosystem, which is a World
Heritage Site and International Biosphere Reserve. Control of the species is a major obstacle
to restoration and preservation efforts.
Although melaleuca has economic uses, its shortcomings outweigh its benefits. In 1991,
its economic benefits, such as use in commercial honey production, were estimated at
$15 million. In contrast, assuming the entire Everglades were to become invaded, the poten-
tial loss to ecotourism, because a melaleuca landscape is less attractive and is impenetrable to
hikers and boaters, was estimated to be $168.6 million. As of 1994, control of melaleuca
cost the state of Florida more than $2.2 million each year. From 1991 to 1998, the
Southern Florida Water Management district spent over $13 million on melaleuca control,
while other agencies also spent millions. For 2003 alone, estimated total costs of control in
southern Florida were more than $3 million.
Management. Management and containment of melaleuca stands is expensive and
requires a long-term commitment. Removal of even a moderately thick stand of trees, either
physically or by injecting individual trees with herbicide, may cost hundreds or thousands
of dollars per acre. Aerial spraying with herbicide is less expensive, but also less effective.
Total eradication may be impossible, but spread of melaleuca can be checked, and some
areas can be restored to their natural state. Treatment of dense stands does little good
 MELALEUCA n 561

because any disturbance to the tree triggers either seed release or resprouting, which produ-
ces more trees. However, concentration of efforts on treatment of trees in outlying areas may
prevent new stands from developing.
Physical removal is possible only for isolated trees or small stands. Hand-pulling of seed-
lings may be temporarily successful, but because of the abundant seed production, new
seedlings may develop in the same site if it is close to other trees. Medium-size trees may
be bulldozed over and larger trees cut, but the roots, limbs in contact with moist ground,
or stumps will resprout. Any attempt to manage the stands by fire is ineffective and actually
serves to extend the range of melaleuca. Although young seedlings, as well as native plants,
are killed, fire triggers seed release from mature trees and also provides a nutrient-rich sur-
face for germination.
Chemical applications will cause stressed plants to release seeds, ultimately making the
situation worse. Depending on the method of application, various herbicides succeed in kill-
ing existing trees. A combination of glyphosate and imazapyr is most effective in both foliar
applications and when applied directly to cuts through the basal bark to the cambium layer.
Resprouting may occur when glyphosate is used alone. Either glyphosate or imazapyr is
effective on cut stumps, a choice when it is unwise to leave dead trees standing. Triclopyr
is less effective. Hexazinone applied to the soil at the base of plants is taken up by the roots.
Research is currently being conducted to explore biological controls, specifically insects
from its Australian native habitat. Five species have been introduced to Florida quarantine
for study. Two have been released to the wild and are currently established. Released insects
disperse slowly, and reduction in seed production, not eradication of the plant, is the goal.
The snout beetle (Oxyops vitiosa) eats growing tips, while the melaleuca psylid
(Boreioglycaspis melaleucae) causes foliage and stems to wilt, killing young trees. These two
species and the melaleuca defoliating sawfly (Lophyrotoma zonalis) are host-specific, meaning
that they only affect melaleuca and do not threaten native plants. The sawfly is being tested
for toxicity to wildlife, because some evidence exists that eating the larvae is toxic to animals.
The bud-gall fly (Fergusonina sp.), which creates galls on both flower and leaf buds, retard-
ing growth, may also be host-specific but is still undergoing research tests. In contrast, the
leaf-blotching bug (Eucerocoris suspectus), although effective at defoliating melaleuca, also
attacked other plants and is no longer being considered. Other species are being tested in
Australia for potential import.

Selected References
Geary, T. F., and S. L. Woodall. “Melaleuca.” n.d. http://www.na.fs.fed.us/spfo/pubs/silvics_manual/
volume_2/melaleuca/quinquenervia.htm.
Masterson, J. “Melaleuca quinquenervia.” Smithsonian Marine Station at Fort Pierce, FL, 2007. http://
www.sms.si.edu/IRLspec/Melaleuca_quinquenervia.htm.
Mazzotti, Frank J., Ted D. Center, F. Allen Dray, and Dan Thayer. “Ecological Consequences of
Invasion by Melaleuca quinquenervia in South Florida Wetlands: Paradise Damaged, not Lost.”
Document SSWEC123. University of Florida Institute of Food and Agricultural Sciences (IFAS)
Extension, 2008. http://edis.ifas.ufl.edu/uw123.
Rayamajhi, M. B., et al. “Australian Paperbark Tree (Melaleuca).” In Biological Control of Invasive Plants in
the Eastern United States, by R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-
04, 2002. http://www.invasive.org/biocontrol/8AutralianPaperbarkTree.html.
Swearingen, Jil M. “Melaleuca.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/
mequ1.htm.
562 n TREES

n Paper Mulberry
Also known as: Wauke (Hawaiian)
Scientific name: Broussonetia papyrifera
Synonyms: Papyrius papyriferus
Family: Mulberry (Moraceae)
Native Range. Japan and Taiwan.
Distribution in the United States. Southeastern, northeastern, and midwestern states.
From Texas east to Florida, north to Massachusetts and New York, and west to Kansas and
Oklahoma. Also in California, Hawai’i, and Puerto Rico.
Description. Paper mulberry is a deciduous, low-branching tree that may be 33–66 ft.
(10–20 m) tall. With many sucker stems emerging from the roots, it might look like a large
shrub, with a mounded appearance because the taller stems are in the middle of the clump.
Reddish-brown twigs are densely covered with coarse hairs, and the bark is tan or dark gray,
either smooth or with shallow grooves or furrows. The soft wood is brittle, and the plant has
milky sap. In winter, the tree can be recognized by its conical buds. Leaves are highly varia-
ble in size and shape, 2.5–10 in. (6–25 cm) long and 2–3.5 in. (5–9 cm) wide, on 1–3 in.
(2.3–8 cm) long petioles. They may be alternate, opposite, or whorled around the stem,
and may have 3–5 lobes, none, or be shaped like a mitten. Small leaves are usually not
lobed, medium-size leaves are mitten-shaped, and the largest are deeply lobed, with two
large lobes and two smaller lobes near the leaf base. They have pointed tips and either a
heart-shaped or rounded leaf base. Leaf margins are coarsely toothed. The lower surface of
leaves is densely covered with soft matted hairs, while the upper surface is sparsely covered
with stiff hairs, making the leaves feel slightly rough to the touch. Leaves turn dull yellow in
fall before dropping.
Trees have a taproot as well as a system of matted surface roots, which give rise to
suckers.
Plants are dioecious, meaning that male and female flowers are on separate plants.
Greenish flowers develop in April and May, when the leaves emerge. Male flowers are
arranged in clusters of drooping catkins, each 1.2–3 in. (3–8 cm) long, which contain
many individual small flowers. Female flowers are globular or ball-shaped, 1 in.
(2.5 cm) in diameter. The reddish-purple to orange-red, fleshy fruit matures in summer
and is an aggregate like a berry, 0.75–1.2 in. (1.5–3 cm) in diameter, looking like a spiky
ball.
Related or Similar Species. The nonnative white mulberry has both male and female
flowers on the same plant. Native red mulberry is distinguished by leaves with a distinct
heart-shaped base and only a slight pubescence on the underside. Twigs may be glabrous
or finely pubescent, and the fruits are red and elongated. Leaves of the native sassafras,
which have a spicy odor or taste, may be unlobed, mitten-shaped, or three-lobed, but the
leaf base is distinctly pointed. Yellow-green flowers borne in short clusters are followed by
blue-black berries nestled in red cups. White basswood, also native, has unlobed, distinctly
heart-shaped leaves on long petioles. Pale-yellow or whitish flowers occur in clusters
attached to a long thin bract. Fruit is small, gray, and pea-like.
Introduction History. Paper mulberry has been known in Florida since 1903, and most of
the plants are male. Because of their rapid growth, trees were planted extensively throughout
the southeastern states as ornamentals and for shade around houses. Oceangoing
Polynesians who originally settled Hawai’i brought paper mulberry shoots with them.
 PAPER MULBERRY n 563

Habitat. Paper mulberry has

become a worldwide invasive.
It thrives in several climates,
preferably where rainfall keeps
the soil moist most of the year.
Trees can be found in disturbed
sites at the edges of forests and
fields, and in open ground near
urban areas below 5,000 ft.
(1,500 m) elevation. They grow
best in sun and do not tolerate
heavy shade. Especially prob-
lematic in riparian zones of
floodplain forests, paper mul-
berry is also known for its toler-
ance of 3–4 months of seasonal
drought. The tree prefers light-
textured soils, such as sands or
sandy loams, and can grow in
either well-drained or water-
logged sites. Because the foliage
is not frost tolerant, the tree
often dies back to the ground
in winter in cold climates.
Reproduction and Dispersal.
Paper mulberry reproduces
both sexually and vegetatively.
Long-distance dispersal is
accomplished by birds and
wildlife, which eat the fruit and
expel the seeds elsewhere.
Seeds can germinate one month Paper mulberry trees grow best in climates where soils are moist,
after ripening. Although most of especially the southern states and Hawai’i. (Native range adapted from
the plants in Florida are male, USDA GRIN and selected references. Introduced range adapted from
enough seeds are produced for USDA PLANTS Database, Invasive Plant Atlas of the United States, and
paper mulberry to be invasive selected references.)
in some areas. Flowers and fruit
do not develop in Hawai’i because all plants are male. In all regions, new plants sprout from
the roots to create dense thickets.
Impacts. Paper mulberry grows aggressively to displace native plant species, which neg-
atively affects wildlife that depend on native plants for forage, cover, and nesting sites.
Repeated growth of suckers, which sprout from the surface root system every few feet,
makes the plant hard to contain. Because of its shallow root system, trees are frequently
blown over in high winds. The roots, however, grow aggressively and can damage pipes
and structures. Pollen from flowering plants may cause severe allergic reactions.
Management. Although paper mulberry is still available in nurseries, the best manage-
ment is to avoid planting it. Physical control may be accomplished by pulling small
564 n TREES

A. Trees begin branching close to the base. (Chuck Bargeron, University of Georgia, Bugwood.org.) B. Leaves are
variable but often are deeply lobed. (Chuck Bargeron, University of Georgia, Bugwood.org.) C. Small leaves may
be mitten-shaped or unlobed. (John Ruter, University of Georgia, Bugwood.org.) D. Twigs are very hairy. (James
H. Miller, USDA Forest Service, Bugwood.org.) E. Old trunks are smooth, with shallow grooves. (James H.
Miller, USDA Forest Service, Bugwood.org.)

seedlings. Small trees can be cut to the ground repeatedly as plants resprout, or cut in com-
bination with herbicide applications on new growth.
Chemical treatments, whether applied directly to basal bark or cut-stump or injected into
tree tissue, are all effective. These methods, in contrast to foliar sprays, specifically target
paper mulberry and do no damage to nontarget plants. Triclopyr is the most effective herbi-
cide applied directly to cut tissue. Foliar applications of glyphosate, however, can be used to
kill small trees if they are thoroughly covered with the herbicide. Herbicide treatments
should be done before the fruit and seeds mature. Because of root sprouts, large trees need
repeated treatments.
The plant appears to be free of pests, and no biological agents are available. Research and
data, however, are limited. Several species of fungi and insects are known to attack paper
mulberry in its native range. Of the fungi, Aecidium mori var. broussonetia, Dendryphiella
broussonetiae, and Phomopssis broussonetiae are host-specific.

Selected References
Morgan, Eric C., and William A. Overholt. “Wildland Weeds: Paper Mulberry, Broussonetia papyrifera.”
Publication ENY-702, University of Florida Institute of Food and Agricultural Sciences (IFAS)
Extension, 2004; reviewed 2010. http://edis.ifas.ufl.edu/in498.

Why Is It Called “Paper” Mulberry?

A s long as 2,000 years ago, the Chinese used the strong, fibrous inner bark of
the tree as a source of fiber for making paper, and it is still used today for
high-quality paper in China and Japan. On island environments where fibers such
as cotton and silk were unknown, Polynesians and Hawaiians used the bark to
make tapa cloth. Traditionally used for clothing and bedding, tapa cloth is valuable
because it is warm, water resistant, soft, and washable.
 PRINCESS TREE n 565

“Paper Mulberry, Broussonetia papyrifera.” Excerpted from Circular 1529, Invasive Species Management
Plans for Florida. Institute of Food and agricultural Sciences (IFAS) Extension, 2008. http://
plants.ifas.ufl.edu/node/74.
Swearingen, Jil M. “Paper Mulberry.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2005. http://www.nps.gov/plants/ALIEN/
fact/pdf/brpa1.pdf.
Weaver, Richard. “Botany Weed of the Month, Broussonetia papyrifera, Paper Mulberry.” Florida
Department of Agriculture and Consumer Services, 2010. http://www.doacs.state.fl.us/pi/enpp/
botany/weed-of-the-month/0410-broussonetia-papyrifera.html.
Zheng, Hao, Yun Wu, Jianquing Ding, Denise Binion, Weidong Fu, and Richard Reardon.
“Broussonetia papyrifera, Paper Mulberry.” In Invasive Plants of Asian Origin Established in the US
and Their Natural Enemies, vol. 1, 2004. http://www.invasive.org/weeds/asian/broussonetia.pdf.

n Princess Tree
Also known as: Royal paulownia, empress tree, paulownia, karritree
Scientific name: Paulownia tomentosa
Synonyms: Bignonia tomentosa, P. imperialis
Family: Figwort (Scrophulariaceae)
Native Range. East-central China.
Distribution in the United States. Eastern and midwestern states, from New York and
Massachusetts south to Florida, west to Texas, Missouri, and Illinois. Also in Oregon.
Description. Princess tree is a small- to medium-size tree, growing 30–60 ft. (9–18 m) tall
and as much as 2 ft. (0.6 m) in diameter. The spread of its canopy can match its height, and
branches tend to droop as the tree grows. The stems or twigs are flattened at the juncture of
stems and branches. Pith of stems is hollowed or chambered. The olive-brown to dark-
brown bark on young growth is thin and smooth, with prominent elongated white lenticels.
Twigs are primarily glabrous except for the young tips, areas around the buds, and the upper
edges of leaf scars, which are covered with short hairs. Leaf scars are circular. Older bark is
gray-brown and rough, but mottled with shiny smooth areas. The large leaves, in opposite
pairs on the stem, are broadly oval or heart-shaped, with pointed tips. Margins are usually
entire, or sometimes with three shallow lobes, but they may be toothed on small plants.
Leaves on young saplings may grow in whorls of three. Leaves on mature trees are
6–16 in. (15–40 cm) long and 4–8 in. (10–30 cm) wide, but leaves on stump sprouts may
be twice as large. Leaves are light green, with the lower surface being more pale in color.
Both leaf surfaces are fuzzy or hairy, especially the undersides. Princess tree has many shal-
low, branching horizontal roots, but no strong taproot.
The tree flowers in April and May, before the leaves develop, from leaf buds that formed
the previous summer. Fragrant, showy flowers grow in conspicuous upright terminal
panicles, 6–12 in. (15–30 cm) long, at the ends of the twigs. Bell-shaped pale violet or blue
corollas, with five round, unequal lobes, are 2 in. (5 cm) long. The fruit grow in terminal
clusters of oval, woody capsules, 1.25–1.75 in. (3–4.5 cm) long, on second-year growth.
Initially coated with sticky hairs, the capsules, or seed pods, mature in the fall and become
brown, dry, and woody, resembling pecans. Seed pods have four compartments, which
together contain as many as 2,000 tiny flat, winged seeds. Capsules remain on the tree
throughout winter.
Related or Similar Species. Most members of the figwort family in North America are
herbaceous. Two other species in the same genus, white-flowered paulownia (P. fortunei)
566 n TREES

and elongate paulownia (P.

elongata), are recommended for
timber plantations in the
United States where winter
temperatures do not drop
below 20ºF (−6.5ºC).
Although the native catalpa
tree is similar in size, leaves,
and flowers, it has distinct
differences from princess tree.
The pith of catalpa stems is
solid and whitish. Its leaves are
whorled, with more distinctly
pointed tips. Catalpa flowers
grow on the current year’s
growth, and catalpa fruit are
slender pods, 8–18 in. (20–
46 cm) long.
Introduction History. After
first being taken to Europe in
the 1830s by the Dutch East
India Company, princess tree
was imported to the United
States around 1840 as an orna-
mental landscape tree. The use
of seed pods to cushion porce-
lain cargo shipped from China
may be a second possible mode
of introduction. Princess tree
has been naturalized in the
eastern states for more than
Grown in plantations for timber, princess tree is widely promoted in 150 years and is also grown on
southern states. (Native range adapted from USDA GRIN and selected the West Coast.
references. Introduced range adapted from USDA PLANTS Database, Habitat. Princess tree pri-
Invasive Plant Atlas of the United States, and selected references.) marily invades bare or sterile soil
in disturbed natural areas, such
as burned sites, landslides, and forests defoliated by pests. Roadcuts and construction sites are
also potential habitat. Trees thrive on marginal, even toxic, land. They tolerate infertile or acid
soils but do best on well-drained soil with a high water-holding capacity and a pH of 6–8.
Princess tree can be found along stream banks, on steep rocky slopes, and at the edges of forests
where sunlight is sufficient, but not in shady forest interiors. Because it was planted as an orna-
mental, trees are frequently found around old homesites. Its natural range in China is south of
the 0ºF (−18ºC) January isotherm and where annual precipitation is at least 40 in. (1,015 mm).
Reproduction and Dispersal. Princess tree reproduces both by seed and from root
sprouts. Flowers are insect pollinated in spring. Seed capsules break open to release seed
throughout the following winter and into the spring. One large tree can produce 20 million
seeds, which are transported long distances by wind and water. Seeds can germinate as soon
as they are deposited in a suitable location, which has bare soil, moisture, and direct
 PRINCESS TREE n 567

A. White lenticels are prominent on young twigs. (James H. Miller, USDA Forest Service, Bugwood.org.)
B. Leaves are lighter green and more pubescent on the lower surface. (James H. Miller, USDA Forest Service,
Bugwood.org.) C. Long inflorescences with bell-shaped flowers grow at the ends of twigs. (James R. Allison,
Georgia Department of Natural Resources, Bugwood.org.) D. Fruit are woody capsules that resemble pecans.
(James H. Miller, USDA Forest Service, Bugwood.org.)

sunlight. Seedlings are intolerant of shade and are susceptible to soil fungi, which causes
damping off. Seedlings and small trees grow rapidly and can flower and set seed in
8–10 years. Mature trees, which live for approximately 70 years, become structurally weak.
Root sprouts are abundant and can grow more than 15 ft. (4.5 m) in a year. Root cuttings
of one-year-old plants are used for propagation in plantations.
Impacts. Princess tree grows rapidly and displaces native species by preventing them
from recovering after a disturbance. Although it is a poor competitor with established veg-
etation, princess tree rapidly occupies bare sites where it creates extensive colonies due to
its prolific root sprouts. It often competes with rare plants on rocky cliffs and in scoured
riparian sites. The litter that accumulates from its large leaves alters soil chemistry by
increasing the nitrogen content.
Management. Physical control is possible for young plants. They may be hand-pulled,
providing that all roots are removed because any fragments may sprout new plants. Large
trees may be cut to ground level, which is best done before the onset of flowering in order
to prevent seed development. Resprouts from the roots, however, should be treated with
herbicides. Girdling can be done where herbicide use is impractical. Cutting through the

Can Paulownia Be Useful?

A s early as the third century BC, princess tree had medicinal and ornamental
uses in China, and has been cultivated in Japan for centuries as a timber tree.
The wood is currently valuable in China and Japan for its use in house construction,
in furniture, and as a base for veneers. It is prized for being easily carved into gift
boxes, toys, musical instruments, bowls, and clogs. In the 1970s, wild trees in the
United States were cut for export to Japan. Since the 1990s, trees have been grown
in plantations in the Mid-Atlantic and southeastern United States, with Japan
being the major market. The tree is currently advertised on the Internet as a won-
derful plant that enriches the soil, absorbs soil pollutants from livestock facilities,
and is good for strip mine reclamation. Plantations are also touted as a means of
saving natural forests from being cut for timber.
568 n TREES

Jimmy Carter and Princess Tree

F ormer president Jimmy Carter has planted 10–15 acres of paulownia on his farm
in Plains, Georgia, and is experimenting with turning them into biofuel. A
cabinet that he built from Paulownia elongata wood sold at auction in 2000 for
$230,000.
Source: Paulownia Supply, 2010. http://www.paulowniasupply.com/paulownia_history
.htm.

bark and into and through the cambium layer completely around the tree about 6 in.
(15 cm) above the ground will kill the top parts, but not the roots. Suckers can be controlled
with herbicide.
Chemical applications of glyphosate or triclopyr are effective on seedlings and small trees.
If desirable grasses are present in the infestation, triclopyr is the better choice because it is
selective to broadleaf plants. Cut-stump, basal bark, or hack-and-squirt methods are also
effective. Herbicides should be applied to stumps immediately after cutting or to basal bark
approximately 12–15 in. (30–38 cm) above the ground surface. Hack-and-squirt involves
cutting into the trunk several times about 6–18 in. (15–45 cm) above the ground.
Glyphosate or triclopyr is then injected into the cuts. All three methods can be done all year
if the ground is not frozen.
Princess tree is resistant to insects, and no biological controls are known. Of eight species
of fungi that occur in its native range in China, four (Ascochyta paulownia, Gloeosporium
kawakamii, Mycosphaerella corylea, and Phyllactinia paulownia) may be host-specific. No evi-
dence indicates that any would be a controlling agent.

Selected References
Bonner, F. T. “Royal Paulownia.” n.d. http://na.fs.fed.us/pubs/silvics_manual/volume_2/paulownia/
tomentosa.htm.
Clatterbuck, Wayne K., and Donald G. Hodges. “Paulownia.” Tree Crops for Marginal Farmland.
Revision of University of Tennessee Extension Publication PB 1465 (1992), 2004. http://
na.fs.fed.us/pubs/silvics_manual/volume_2/paulownia/tomentosa.htm.
“Princess Tree.” Southeast Exotic Pest Plant Council (SEEPPC) Invasive Plant Manual, n.d. http://
www.se-eppc.org/manual/princess.html.
Remaley, Tom. “Princess Tree.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/
pato1.htm

n Russian Olive
Also known as: Oleaster
Scientific name: Elaeagnus angustifolia
Synonyms: Elaeagnus hortensis, E. iliensis
Family: Oleaster (Elaeagnaceae)
Native Range. Eastern Europe and western Asia, including western Russia, Mongolia,
northern China, and Southwest Asia from Kashmir west to Iran and the Caucasus.
 RUSSIAN OLIVE n 569

Distribution in the United

States. Almost every state
except the extreme Southeast,
Alaska, and Hawai’i. Especially
invasive in the Intermontane
West and riparian sites on the
Great Plains.
Description. Russian olive is
a perennial small tree or large
shrub as much as 15–35 ft.
(4.5–10.5 m) tall. The tallest
trees are in cultivation. Trees
have a rounded shape, generally
with a single trunk and bra-
nches loosely arranged. Bark
on the trunk is dark brown with
deep fissures. Young branches
are gray, with a scaly pubes-
cence that is lost as they age.
Older branches, covered with
smooth reddish-brown bark,
remain flexible. Stems, buds,
and leaves are densely covered
with silvery or rusty scales.
Twigs are thorny. Narrow
leaves are simple and lance-
shaped, 1–4 in. (2.5–10 cm)
long and 0.4–1.2 in. (1–3 cm)
wide, borne on short petioles
in an alternate arrangement.
Although leaf margins are not
toothed, they may be wavy, Although drought tolerant, Russian olive trees grow primarily in well-
and leaf tips may be either blunt drained riparian sites throughout the United States. (Native range
or pointed. The upper surface adapted from USDA GRIN and selected references. Introduced range
of leaves is dull green and adapted from USDA PLANTS Database, Invasive Plant Atlas of the
covered with star-shaped sil- United States, and selected references.)
very hairs. Lower surfaces are
silvery white and densely covered with scales. Leaves do not change color before dropping
in autumn. The lateral root system is deep and extensive.
Bell-shaped tubular flowers appear in May and June, soon after the plant leafs out, fol-
lowed by fruit from August through October. Flowers, 0.4 in. (1 cm) long, grow in clusters
of 1–10 on short stalks in the leaf axils. Although flowers have no petals, the four yellowish-
greenish sepals, which are fused at the base and flared at the top, resemble petals. Flowers
are very fragrant, silvery, and scaly outside and creamy yellow inside. The hard fruits are
olive-shaped, 0.4–0.8 in. (1–2 cm) long, and remain on the tree all winter or until consumed
by birds or other wildlife. They are mealy, greenish-yellow to brown, sometimes tipped with
red, and covered with silvery scales. Although each fruit contains only one seed, each tree
produces an abundance of fruit.
570 n TREES

A. Russian olive is a shrubby tree. (Norbert Frank, University of West Hungary, Bugwood.org.) B. Narrow lance-
shaped leaves have a silvery or gray cast. (Paul Wray, Ohio State Weed Lab Archive, The Ohio State University,
Bugwood.org.) C. Twigs have stout thorns. (Patrick Breen, Oregon State University, Bugwood.org.) D. Flowers
have four fused sepals. (Joseph Berger, Bugwood.org.) E. Hard fruit each contain one seed. (Patrick Breen,
Oregon State University, Bugwood.org.)

Related or Similar Species. Silverberry is the only species in the genus that is native to
North America. It is a shorter, root-sprouting shrub with dark young stems that are not sil-
very. It grows primarily east of the Rocky Mountains but is also found in Idaho and Utah.
Russian olive may also be confused with the native silver buffaloberry, which is closely
related to Russian olive. Silver buffaloberry grows in the same saline riparian sites, but has
opposite leaves and smaller, red-orange oval berries. It grows slowly into short thickets,
reaching 10 ft. (3 m) tall after 20 years.
The closely related autumn olive (Elaeagnus umbellata) grows in the eastern half of the
United States, from the Atlantic coast west to Nebraska, and in Hawai’i, and is considered
a noxious weed in some states. Both autumn olive and Russian olive can be found growing
together where their ranges overlap. Native to eastern China, Japan, and Korea, autumn
olive was introduced to the United States in the 1830s as an ornamental, for windbreaks
and wildlife habitat, and to restore degraded land such as strip mines. Although it has the
potential of becoming a troublesome weed, seeds are still distributed in some states for wild-
life plantings. A deciduous shrub or small tree reaching 20 ft. (6 m), with a habit more bushy
than tree-like, it thrives in full sun on disturbed sites. The lance-shaped leaves are alternate
and not toothed. The upper surface of the leaves is dark green, with only the underside
covered with silver-white scales. Its branches are not flexible and usually have no thorns.
It has small, light-yellow flowers in early summer. Many round, juicy, reddish berries dotted
with brown to silvery scales mature in autumn, and seeds are dispersed primarily by birds
and small mammals. It does not do well in wet sites or in dense forest, although seedlings
are shade tolerant. Because it is drought tolerant, it invades grasslands and woodlands,
and is typically found in disturbed ground in pastures, fields, and along roadsides.
Methods of control are similar to those used for Russian olive.
Russian olive also resembles silverthorn, also called thorny olive, which is a nonnative
evergreen with brown scaly and hairy twigs. Flowering in late fall, silverthorn matures a
few reddish silver-scaly fruit in spring.
 RUSSIAN OLIVE n 571

Introduction History. Cultivated in Germany since 1736, Russian olive has been a popu-
lar landscaping plant since it was first introduced to North America in the late 1800s or early
1900s. The first records in the West date from 1903 to 1909 in New Mexico, Nevada, and
Arizona. By the 1940s, it was a common ornamental in western cities because of its attractive
silvery foliage. Young, spiny stems made a good barrier hedge. By 1954, it had escaped cul-
tivation in several western states. As recently as 1984, some government agencies promoted
Russian olive for soil stabilization, as windbreaks, and for improvement of wildlife habitat,
even subsidizing distribution of seedlings. It is still recommended and sold in nurseries.
Habitat. Although Russian olive tolerates seasonal drought, it grows predominantly in
wet places, such as riparian sites, where it can withstand both flooding, siltation, and salin-
ity. It often colonizes downstream from dams, where regulated water release changes the
natural cycle of seasonal flooding. The tree also requires good drainage, and may not be
invasive in Southeastern states. Trees are found in disturbed, seasonally moist floodplain
forests and irrigation ditches, but also in drier sites, such as railroad and highway right-of-
ways, fence lines, and grasslands. It is widely used in mixed-species shelterbelts across the
prairie and plains states. Although it tolerates a wide range of soils, from coarse sand to
heavy clay, it grows best in deep sandy or loamy soils. As a nitrogen fixer, it has been used
extensively for mine reclamation on bare mineral substrate and tailings, but it does not tol-
erate acidic soils (pH < 6.0). Seedlings are shade tolerant, but adults are not.
Russian olive can grow in temperature conditions as low as −50ºF (−45ºC) and as high as
115ºF (46ºC), and from sea level to 8,000 ft. (2,500 m) elevation. Although trees need at
least 8 in. (200 mm) of precipitation each year, plants are well adapted to ephemeral water
conditions in riparian zones in dry regions. Russian olive is outcompeted by the invasive salt
cedar (see Trees, Tamarisk) on salty soils.
Reproduction and Dispersal. Reproduction is both sexual and asexual. Trees produce
seed when they are 3–5 years old. Birds feed on the fruit and disperse the seeds. Fruit are
also eaten by coyotes, deer, and raccoons, and seeds are gathered and stockpiled by small
animals. The buoyant seeds may also be transported by water. Seeds are viable for three
years but need 60–90 days of cold conditions to germinate. Russian olive also resprouts from
buds on the root crown after damage to aerial parts, whether natural or human-induced, and
stem cuttings can produce new trees.
Impacts. Russian olive interferes with natural plant succession and alters nutrient cycling.
It also uses more water than native plants and changes hydrology of streams. By forming
dense thickets, plants crowd out and displace native vegetation, particularly riparian forests,
resulting in loss of wildlife habitat.
Native cottonwoods and willows have narrow germination requirements and cannot grow
in the shade. In contrast, germination of Russian olive seeds takes place under a wide range of
conditions, and few seedlings die. Plants grow quickly when young, up to 6 ft. (1.8 m) in one
year, with the result that Russian olive stands replace the native plant community. Both a pio-
neer and climax species, Russian olive persists through successional changes. Beavers’ prefer-
ence for cottonwoods and willows inadvertently favors Russian olive. Russian olive is quick to
displace riparian vegetation after disturbances, such as flooding. Trees may also alter the flood
regime. Thickets stabilize stream banks and alter hydrology, limiting sites for regeneration of
native cottonwoods and changing riparian habitats into dry uplands.
In spite of producing abundant fruits, which are eaten by more than 50 species of birds
and mammals, monospecific stands of Russian olive lower the quality of wildlife habitat.
Birds that require nesting cavities and insect food in native trees are displaced. Waterfowl,
such as ducks, avoid water areas bordered by Russian olive.
572 n TREES

Because it is a nitrogen fixer, Russian olive changes the chemical composition of infertile
soils that native plants may require. Dense thickets also increase the fuel load, resulting in
more damaging fires.
Management. Because mature stands of Russian olive trees are nearly impossible to
totally eradicate, the best management is to destroy small plants as they are discovered. All
control methods should be repeated until the seed bank is exhausted. Any physical control
on mature plants should be done before seeds mature. Small trees less than 3.5 in. (9 cm)
diameter, and their roots, can be pulled out of moist soil. Repeated mowing or burning of
seedlings with stems less than 1 in. (2.5 cm) diameter may control the infestation. Tilling
is effective to renovate pastures because all saplings and roots are churned up with the soil.
However, mowing, cutting, bulldozing, or chaining will remove only the top biomass of
larger trees and provide only short-term effectiveness. Roots should be cut below ground
level and the rest buried. Stumps may also be burned. Girdling and cutting trunks above
ground level is ineffective because of sprouts from the root crown. Any plant debris should
be removed from the site because it does not readily decay and poses a fire hazard. Follow-
ups should be conducted for several years to remove all sprouts and seedlings.
Although chemical control is effective either by foliar spray or basal injection on young
trees or resprouts, the best method for mature trees is application of herbicides to cut
stumps. A systemic, such as glyphosate or triclopyr, can be used during the growing season,
but herbicides such as imazipyr may be more effective when trees are dormant. Trees should
be cut as low as possible, to ground level or below, then treated with herbicide.
No research has been conducted on biological control of Russian olive. It was promoted
as an ornamental because of its resistance to insects or disease. A canker disease
(Tubercularia ulmea) that turns leaves brown can occasionally be found on branches and
trunks, but it is not a means of control. Phomposis canker (Phomposis arnoldiae, P. elaeagni)
may kill seedlings and saplings but is not widespread.

Selected References
“Autumn Olive.” Southeast Exotic Pest Plant Council (SEEPPC) Invasive Plant Manual, 2003. http://
www.invasive.org/eastern/eppc/autolive.html.
Sather, N., Nancy Eckardt, and TunyaLee Martin. “Element Stewardship Abstract, Elaeagnus umbel-
lata.” Global Invasive Species Team, Nature Conservancy, 1987; revised 2009. http://
wiki.bugwood.org/Elaeagnus_umbellata.
Stannard, Mark, Dan Ogle, Larry Holzworth, Joe Scianna, and Emmy Sunleaf. “History, Biology,
Ecology, Suppression, and Revegetation of Russian-Olive Sites (Elaeagnus angustifolia, L.).” Plant
Materials Technical Note No. MT-43. U.S. Department of Agriculture, Natural Resources
Conservation Service, Montana, 2002. ftp://ftp-fc.sc.egov.usda.gov/MT/www/technical/plants/tech
notes/PMC_Tech_Note_43.pdf.
Tu, Mandy. “Element Stewardship Abstract, Elaeagnus angustifolia.” Global Invasive Species Team,
Nature Conservancy, 2003; updated 2009. http://wiki.bugwood.org/Elaeagnus_angustifolia.

n Silk Tree
Also known as: Mimosa, silky acacia, powderpuff tree,
Scientific name: Albizia julibrissin
Synonyms: Albizzia julibrissin, Mimosa julibrissin, Mimosa arborea
Family: Pea (Fabaceae)
Native Range. From northern Iran and Afghanistan through the Himalayas in northern
India and Nebal, and into China and Japan.
 SILK TREE n 573

Distribution in the United

States. Much of the United
States, from California to Flo-
rida, north to New York and
Massachusetts and west to Illi-
nois and Kansas. Absent from
the northern regions of the
Midwest, the Great Plains,
Rocky Mountains, and Pacific
Northwest.
Description. Silk tree is a
small to medium sized decidu-
ous tree, 10–50 ft. (3–15 m)
tall, with a canopy spread of
25–30 ft. (7.5–9 m). It may have
one trunk or multiple trunks.
The leaf canopy, which gives
the tree an umbrella shape, is
generally open, allowing sun-
light to penetrate. The smooth,
thin bark is light brown to gray
and covered with lenticels that
look like corky dots and dashes.
Young stems are lime green,
becoming brown and covered
with lenticels as they age.
Wood is weak and brittle. The
doubly compound leaves, alter-
nate on the stem and 6–20 in.
(15–50 cm) long, are finely di-
vided and almost fern-like or
feathery. The compound leaf Silk tree grows in a variety of environments but is restricted to the
may have 8–24 pairs of pinnae, southern half of the United States by cold winter temperatures. (Native
or branches, with 20–60 dark range adapted from USDA GRIN and selected references. Introduced
green leaflets, each 0.5 in. range adapted from USDA PLANTS Database, Invasive Plant Atlas of the
(1.3 cm) long, on each pinna. United States, and selected references.)
The midrib is off center, closer
to one margin of the leaflet.
The light-pink to dark-pink flowers, showy and sweetly fragrant, emerge from May
through August. Many conspicuous stamen filaments make them resemble small pink pom-
poms or tufts, 1.5–2 in. (3.8–5 cm) long, with white bases. Flowers are sessile and occur in
loosely branched clusters of 15–25 at the ends of the current year’s growth of branches. Flat,
linear, straw-colored seed pods, 6 in. (15 cm) long, contain 5–10 light brown oval seeds,
each 0.5 in. (3.8 cm) long. Seed pods hang in large clusters and turn from light green to dark
brown as they mature in August and September. In winter, the pods are whitish tan.
Although they split to release the seeds, the empty pods remain on the trees all winter.
Related or Similar Species. Woman’s tongue, an introduced tree species native to
southern Asia and northern Australia that is sometimes invasive, is found in California,
574 n TREES

A. Foliage is open and lacy looking. (Rebekah D. Wallace, Bugwood.org.) B. Long leaves are pinnately
compound. (James H. Miller, USDA Forest Service, Bugwood.org.) C. Flowers resemble fluffy pompoms.
(Rebekah D. Wallace, Bugwood.org.) D. The smooth bark is spotted with lenticels. (James H. Miller, USDA
Forest Service, Bugwood.org.) E. Pods each contain 5–10 seeds. (James H. Miller, USDA Forest Service,
Bugwood.org.)

Texas, Florida, Hawai’i, Puerto Rico, and the Virgin Islands. It is distinguished from silk tree
by its creamy or white flowers with numerous long stamens and a longer seed pod, up to
1 ft. (30 in). The seed pods, which do not split when ripe, remain on the tree in winter.
The dried seeds make a rattling sound when the pods are disturbed by wind. Leaflets are
more rounded, and leaves do not appear as feathery.
The crimson bottle brush shrub, a member of the myrtle family and native to Australia,
has similar pompom or bottle brush flowers. It can be distinguished by the red flower color,
lance-shaped entire leaves, and woody capsules or seed pods.
Silk tree, especially seedlings or saplings, may be confused with native pinnately com-
pound legumes. Native to the southeastern United States, littleleaf sensitive-briar is an arch-
ing perennial vine with prickly stems and lavender to pink flowers. Partridge pea, also native
to the Southeast, is a non-woody annual forb with five-part yellow flowers. Honeylocust,
native to the Mississippi River drainage system, can be a very large tree, up to 100 ft.
(30 m) tall. Its pinnate leaves have longer leaflets, 1 in. (2.5 cm), and branches often have
1.2–4 in. (3–10 cm) long thorns. Greenish or whitish flowers are tiny, and the long seed
pods, 6–8 in. (15–20 cm), are slightly curved or twisted.
Introduction History. Silk tree was introduced to the United States in 1745 as an orna-
mental because of its finely divided foliage and its showy, fragrant flowers, which are attrac-
tive to hummingbirds. It also requires little maintenance. Trees have been used to reclaim
disturbed areas in shoreline or riparian sites, and the species is readily available at nurseries
or on the Internet.
Habitat. Silk tree is an invader of disturbed habitats, and can be found along roadsides
and in open vacant lots in urban or suburban areas. It can grow in a variety of soils, includ-
ing clay, loam, and sand, either alkaline or acidic. It is tolerant of drought, wind, salty soils,
and moderate salt spray. Soils can be dry or wet, and plants can be a serious problem in
riparian zones along scoured shorelines. Although the plant prefers full sun, it can grow in
part shade. It is rare under full forest canopy. Limited by cold winters, silk tree grows below
3,000 ft. (900 m) elevation.
Reproduction and Dispersal. Silk tree produces a very large seed crop and also spreads
vegetatively. Most seeds fall around the parent plant, but they may be carried long distances
by animals or by water. Seeds may also be a contaminant in fill dirt. The impermeable seed
coat requires scarification to germinate, enabling seeds to be viable for many years. One
 SILK TREE n 575

study showed that 90 percent of seeds germinated after 5 years, and 33 percent germinated
after 50 years.
Damage to the top portion of the tree will trigger regrowth from the roots. Sprouts can
grow over 3 ft. (1 m) in one season, and multiple root sprouts can result in dense colonies
of trees.
Impacts. Silk tree’s wide tolerance to environmental conditions enables it to outcompete a
wide variety of native trees and shrubs in disturbed areas. The tree is also a strong competi-
tor in open areas and at forest edges. It has a distinct advantage because it grows on several
soil types and reproduces profusely from large seed crops and also has the ability to resprout
after damage. Dense stands or thickets reduce sunlight and nutrients for native plants. As a
nitrogen fixer, it alters soil chemistry. On riparian sites, scoured banks provide open habitat
for germination of seeds, and water currents provide a means of seed dispersal.
Management. Ideal control is to limit planting and to remove existing plants from the
landscape. Because physical control alone is usually inadequate, herbicide application is
appropriate. Young seedlings can be pulled by hand, but care must be taken to remove all
the roots to prevent resprouting. Root sprouts can be repeatedly cut as they emerge or
treated with herbicides. Midsize trees may be cut at ground level, preferably before flowers
mature to prevent seed set. Herbicide should be applied to the resprouts. Girdling may be
necessary for large trees that cannot be cut down or sprayed with herbicides. Cutting
through the bark all around the base of the trunk, about 6 in. (15 cm) off the ground, will
kill the upper part of the tree but not the roots. Resprouts should be treated with a herbicide.
Chemical applications include foliar spray, cut-stump, and injection with either glyph-
osate or triclopyr. Resprouts, small areas, or thickets are appropriate for foliar spraying
where no damage will be done to nontarget native plants. Because it targets broadleaf
plants, triclopyr is a better choice if grasses are present. Cut-stump applications should
be used where sprays might damage desirable natives. Herbicide application should be
done within one minute of cutting the tree. Very large trees can be injected with imazapyr
or triclopyr.
Mimosa wilt (Fusarium oxysporum f. perniciosum), a fungus transferred through the soil, is
a potential biological control. It infects the root system of the tree and can be fatal. Although
currently not used, research is needed to ascertain its effectiveness.

Selected References
“Albizia lebbeck.” In Identification and Biology of Non-Native Plants in Florida’s Natural Areas, edited by
K. A. Langeland and K. Craddock Burks. University of Florida, IFAS, 1998. http://www.fleppc
.org/ID_book/albizia%20lebbeck.pdf.
Miller, James H. “Silktree, Mimosa.” In Nonnative Invasive Plants of Southern Forests: A Field Guide for
Identification and Control. General Technical Report SRS-62. U.S. Department of Agriculture,
Forest Service, Southern Research Station. Asheville, NC, 2003. http://www.invasive.org/eastern/
srs/S_M.html.
“Mimosa.” Southeast Exotic Pest Plant Council (SE-EPPC) Invasive Plant Manual, n.d. http://www
.se-eppc.org/manual/mimosa.html.
“Mimosa Tree, Albizia julibrissin.” Excerpted from University of Florida, IFAS Extension, Circular 1529,
Invasive Species Management Plans for Florida. University of Florida Institute of Food and
Agricultural Sciences (IFAS) Extension, 2008. http://plants.ifas.ufl.edu/node/29.
Remaley, Tom. “Silk Tree.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2005. http://www.nps.gov/plants/alien/fact/
alju1.htm.
576 n TREES

n Strawberry Guava
Also known as: Cattley guava, purple guava, pineapple guava, purple strawberry guava,
cherry guava, waiawi (Hawai’i)
Scientific name: Psidium cattleianum
Synonyms: Psidium littorale var. longipes
Family: Myrtle (Myrtaceae)
Native Range. Atlantic coast of southeastern Brazil. The red-fruited strawberry guava
originates from 3,000 to 4,000 ft. (900–1,200 m) elevation in eastern Brazil, while the
yellow-fruited varieties are native to coastal Brazil.
Distribution in the United States. All major islands of Hawai’i, subtropical Florida, and
Puerto Rico. Also in Arizona.
Description. Strawberry guava
is an evergreen shrub or small
tree usually growing 6–20 ft.
(1.8–6 m) tall, but it can reach
25 ft. (8 m). The trunk is smooth
and pale brown, and multiple
sprouts from the roots may
cause the plant to be shrubby.
Young branches are round
and pubescent. The elliptical to
oblong leaves, arranged oppo-
site on the stems, vary in size,
2–5 in. (5–13 cm) long and
1–2.5 in. (2.5–6.5 cm) wide.
They are glossy dark green,
leathery, and aromatic, with
smooth margins. Lateral veins
are not prominent.
Flowering and fruiting occur
all year. Flowers, 1.2 in.
(2.5 cm) wide, grow singly in
leaf axils. The white petals sur-
round a mass of white and yel-
low stamens. Fruit is a round,
red berry, 1.2–2.4 in. (3–6 cm)
in diameter. The whitish flesh
is translucent and very juicy. It
is sweet when ripe, tasting
somewhat like strawberries.
Two varieties have yellow fruit.
Psidium cattleianum f. lucidum is
a narrow tree, while Psidium cat-
Strawberry guava is considered to be one of the worst invasive pests in
Hawai’i. (Native range adapted from USDA GRIN and selected tleianum var. littorale is more
references. Introduced range adapted from USDA PLANTS Database, substantial. Each fruit contains
Invasive Plant Atlas of the United States, and selected references.) many small, hard, tan seeds.
 STRAWBERRY GUAVA n 577

A. The trunk of a strawberry guava tree is smooth. B. The leathery leaves are opposite on the stem. C. The showy
flowers have many stamens. D. The sweet fruits are round berries. (Forest and Kim Starr.)

Related or Similar Species. The common guava (Psidium guajava) is variable in form
because many cultivated varieties exist. Its leaves are larger, 1.6–6 in. (4–15 cm) long, and
stiff. Leaf cuticles are waxy, and the undersides of leaves are pubescent, with prominently
raised veins. Its bark is smooth, reddish brown, and mottled with reddish brown, gray, or
white patches. The white flowers turn yellow as they mature. Fruit is yellow or pink berries,
1.2–2.4 in. (3–6 cm) in diameter, with many yellow or white seeds.
Introduction History. Strawberry guava was introduced into Florida in the 1880s as both
an ornamental plant and as a fruit crop, and has been planted extensively. Fruit is used in
fruit drinks and in jam. By the 1950s, the plant was considered to be naturalized and grow-
ing wild. It was introduced to Hawai’i in 1825, probably for similar purposes. It is currently
used for firewood in Hawai’i.
Habitat. Strawberry guava grows in a variety of habitats in tropical and subtropical cli-
mates, from dry to moist forests. Its broad tolerance to environmental conditions may be
explained by its genetic variability. Although it is most abundant below 2,600 ft. (800 m),
it grows at elevations as low as 330 ft. (100 m) and as high as 4,265 ft. (1,300 m). In
Hawai’i, the tree grows from sea level to 4,000 ft. (1,200 m) elevation. Yellow-fruit forms
are more common at lower elevations, while red is more common at higher elevations.
Trees grow in a range of annual rainfall from 50 in. (1,250 mm) to 275 in. (7,000 mm).
Although the species grows readily in disturbed sites, such as roadsides and pastures, it
can invade undisturbed healthy forests. It has been found in 23 different vegetation types
in Hawai’i, but is most common in moist forests, both lowland and submontane, especially
sites where feral pigs (see Volume 1, Vertebrates, Mammals, Feral pig) are present. It is quick
to invade wet ohia tree and tree fern rainforest as well as wet ohia tree and koa tree rainfor-
ests. Extremely shade tolerant, seedlings and root sprouts can grow in dense shade under
taller plants, including adult strawberry guava trees. It easily survives on acidic soils and
heavy tree fern litter, both of which are common in natural Hawaiian rainforest. Where
native seedlings may be killed by deep leaf litter, small strawberry guava stems bend under
the weight. They not only survive but send up vigorous shoots.
Mature trees are hardy to about 22ºF (−5.5ºC). Yellow-fruited varieties are somewhat less
tolerant of freezing temperatures.
Reproduction and Dispersal. Strawberry guava reproduces and spreads both sexually
and vegetatively. The number of seeds contained in each fruit can be 25–70. Although seeds
have a short period of viability, they have a high rate of germination. No extensive seed bank
578 n TREES

remains in the soil. The numerous fruits are consumed by birds and feral pigs, and the seeds
are excreted elsewhere. Two nonnative birds, the Common Myna (see Volume 1,
Vertebrates, Birds, Common Myna) and the Japanese White-eye (see Volume 1, Vertebrates,
Birds, Japanese White-eye), are suspected of transporting seeds, but no evidence exists. Feral
pigs play a major role in seed dispersal of strawberry guava. Although scarification is not
required, seeds germinate more quickly after passing through the pig’s digestive tract.
Calculations, extrapolated from a count of the number of seeds found in pig scat, indicate that
in a densely infected area of Kipuhulu Valley on Mau’i, each pig was responsible for dispersing
about eight million seeds each month during the peak fruiting season.
Dense thickets develop when roots sprout stems. Stems from root suckers grow rapidly
and have a good survival rate. Root sprouts also have more leaf area and are more vigorous
because of nutrients stored in the roots. Cuttings will root.
Impacts. Considered by some to be the worst plant pest in Hawai’i, strawberry guava
may affect one-half of the land area of the islands. The tree alters habitat in parks and preserves
and is a major threat to the native flora and fauna. It is present on approximatley 300,000 ac.
(120,000 ha) of natural preserves on the island of Hawai’i alone. It has a competitive advantage
for several reasons. It produces abundant fruit and seeds, and it grows rapidly. It tolerates
shade and leaf litter, sends up sprouts from roots, and may exude alleleopathic compounds.
Strawberry guava has become the dominant tree in several of Hawai’i’s native forests, including
within the boundaries of two national parks. Monospecific thickets of strawberry guava trees
are very dense, shading out native plants in both forests and in formerly open woodlands.
Stem densities in thickets have been measured as high as one every square foot (9 per m2).
A dense network of feeder roots forms a mat at the soil surface, allowing little else to grow.
The tree also threatens groundwater supplies and stream flow. Strawberry guava uses
about 25 percent more water in transpiration than natural rainforest trees require.
Strawberry guava, as well as common guava and Surinam cherry, is a major host for the
nonnative Caribbean fruit fly, which also spreads to commercial crops. Decreased yield
and damaged fruit costs approximately $7.8 million a year for papaya growers alone.
Management. Because they are a major dispersal factor, feral pigs must be controlled
before strawberry guava infestations can be eliminated. Re-infestation is low in intact, pig-
free forests, even when sources are nearby, and recovery potential is good. However, the
absence of pigs will not ensure the absence of strawberry guava. Elimination of pigs must
be followed by physical control, chemical control, or both.
Physical control is only possible on seedlings and saplings that have grown from seed.
After being pulled out, they must be removed from the site, because plants left on the
ground can resprout or grow roots in high-precipitation areas. Because of the extensive root
system, pulling out plants is less effective on root sprouts.
Strawberry guava is sensitive to several types of chemical application. Triclopyr, dicamba,
picloram, and 2,4-D are effective when applied to cut surfaces. Basal bark application of
2,4-D, picloram, and triclopyr also work well. Although glyphosate sprayed on foliage proved
to be ineffective in the long term, it works well when applied directly to slashed trunks.
Long-term management may depend on biological agents, but they must not become a
threat to commercially grown common guava. Of seven potential biological control species
found to attack strawberry guava in its native Brazil, a leaf-galling scale insect (Tectococcus
ovatus) appears to be a good candidate. The formation of galls on new leaves where the insect
feeds saps energy from the plant that would normally be used for growth and the production
of fruit. The insect does not kill the tree but reduces its vigor and limits its ability to spread.
After extensive testing on more than 80 plant species, it was determined that the leaf-galling
 TAMARISK n 579

Resistance to Biological Control

P ublic outcry delayed the proposed release of an insect to control strawberry

guava. Some Hawaiians had several concerns. They were afraid that the insect
would attack commercial guava plantations. They also wanted to continue gather-
ing wild strawberry guava fruit and grow the trees on their own property. They
were also worried that they would have no wood for smoking meat, or that forests
of dead strawberry guava trees would promote forest fires.

insect is host-specific. Neither the common guava nor the closely related ohia tree is suscep-
tible. Fewer fruit will also reduce the number of fruit flies that infest commercial crops.
Because the insect is flightless, it relies on crawling or wind for dispersal, and unless facili-
tated, its spread will be slow. Permits were granted in April 2008, but the release scheduled
for 2009 was delayed.

Selected References
Benton, Nancy. “Strawberry Guava.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/
psca1.htm.
Langeland, K. A., and K. Craddock Burks, eds. “Psidium cattleianum.” In Identification and Biology of
Non-Native Plants in Florida’s Natural Areas. Florida Exotic Pest Plant Council (FLEPPC), 2008.
http://www.fleppc.org/ID_book/psidium%20cattleianum.pdf.
Motooka, P., et al. “Psidium cattleianum.” In: Weeds of Hawai’i’s Pastures and Natural Areas: An
Identification and Management Guide. College of Tropical Agriculture and Human Resources,
University of Hawai’i at Manoa. 2003. http://www.ctahr.hawaii.edu/invweed/WeedsHI/
W_Psidium_cattleianum.pdf
Tunison, Tim. “Element Stewardship Abstract, Psidium cattleianum.” Global Invasives Team, Nature
Conservancy, 1991; updated 2009. http://wiki.bugwood.org/Psidium_cattleianum.
“Waiawi Biocontrol Controversy.” Environment Hawaii 19(1): 2008. http://www.hear.org/articles/envi
ronmenthawaii/eh200807vol19no01/strawberryguavacontroversy200807.pdf.

n Tamarisk
Also known as: Saltcedar, salt cedar, and others; see below.
Five-stamen tamarisk, Chinese tamarisk (Tamarix chinensis); pink-flowered tamarisk
(T. ramosissima); small-flowered tamarisk (T. parviflora); French tamarisk (T. gallica)
Synonyms: Tamarix juniperina, T. petranda, T. pallasii var. brachystachys
Family: Tamarisk family (Tamaricaceae)
Native Range. Arid or semiarid regions of southern Europe and Asia, from the
Mediterranean region east to Korea. Five-stamen tamarisk is native to east-central China.
Pink-flowered tamarisk is from south-central Asia, from the Ukraine east through southern
Russia, Kazakhstan, Mongolia, and western China. This species extends into Southwest
Asia, including Iraq, Iran, Afghanistan, and western Pakistan. Small-flowered tamarisk is
native to the Balkan peninsula and Turkey, while French tamarisk originates in the
Mediterranean Europe regions of Italy, France, and Spain.
580 n TREES

Distribution in the United

States. Tamarisk species are
found primarily in the hot and
arid western, southwestern,
Great Basin, and Great Plains
states. While present in some
of the eastern states, the trees
are less invasive in those more
humid climates. With few
exceptions, five-stamen tama-
risk is limited to the Great
Plains and western states. Pink-
flowered tamarisk has a similar
distribution but is absent from
the Pacific Northwest and
present in the Southeast. Small-
flowered tamarisk is generally
found throughout contigu-
ous United States, with some
absences in the northern and
Great Plains regions. French
tamarisk grows primarily in the
mountain and southwestern
states, but also occurs in the
Southeast.
Description. Four tamarisk
species are considered to be
invasive, none of which is easily
distinguishable from vegetative
characteristics. Although the
worst two, five-stamen tama-
Tamarisk species are distinct in their native ranges, but hybridize in the risk, also called Chinese tama-
United States. (Native range adapted from USDA GRIN and selected risk, and pink-flowered, also
references. Introduced range adapted from USDA PLANTS Database, called saltcedar, are distinct
Invasive Plant Atlas of the United States, and selected references.) species in their native ranges,
hybridization in the United
States confuses identification. All four species, including small-flowered tamarisk and
French tamarisk, are commonly referred to as saltcedar or simply tamarisk.
Tamarisk plants are deciduous shrubs or bushy trees growing 30 ft. (9 m) or more tall.
Normal growth is a tree with one trunk, but with disturbance, the plant may form multiple
sprouts from the base. Although stands may be dense, individuals are loosely branched, giv-
ing the plants an airy look. Bark on young growth is smooth reddish brown, changing to fur-
rowed dark brown or dark purple on older and larger plants. Tiny leaves, 0.06 in. (1.5 mm)
long, are alternate on the stem but are closely packed and scaly, resembling those of cedars.
That characteristic combined with its tendency to grow in salty environments is the basis for
the common name saltcedar. Leaves have salt-secreting glands and are often encrusted with
salt. The drought-tolerant leaves, which are deciduous, have water-storing tissue and
sunken stomata.
 TAMARISK n 581

Salt cedar has an extensive

root system, consisting of both a
deep taproot and lateral roots,
the relative importance of each
type is dependent on soil charac-
teristics and depth to water table.
Lateral roots are 12–16 in. (30–
40 cm) below the soil surface.
The taproot gives the plant
important access to deep under-
ground water sources, enabling
it to survive in very dry habitats.
Flowering generally coincides
with snowmelt and summer
rains, usually from April through
September, but can occur con-
tinuously if water is available.
White, light-pink, or reddish
flowers are small, less than
0.2 in. (5 mm) long. Grouped
in short, dense panicles, 2 in.
long (5 cm), of individual race-
mes at the ends of the current
year’s growth, they appear feath-
ery. Fruit is a capsule containing
many tiny seeds, less than
1 mm long, which may be pro-
duced all summer.
Related Species. A noninva-
sive species also native to
Southwest Asia, athel tamarisk,
is an evergreen species growing Five-stamen tamarisk and pink-flowered tamarisk, primarily found in the
in hot deserts. Although leaves drier western states, are the two most invasive species, (Native range
are not scaly, they conspicu- adapted from USDA GRIN and selected references. Introduced range
ously sheath the stem. Branch- adapted from USDA PLANTS Database, Invasive Plant Atlas of the
lets have a drooping habit. United States, and selected references.)
Introduction History. The
earliest record, 1823, is in New York City, and the trees were available at East Coast nurser-
ies by the 1930s. The plant was brought to the western states sometime in the 1850s. In the
early 1900s, the trees were recommended as ornamentals and for windbreaks, shade for
small livestock, and fuel. As late as 1964, it was still recommended for windbreaks on the
Great Plains. It was intentionally planted along rivers in western states to control bank ero-
sion. Regulation of water flow in reservoirs and diversion channels allowed the tree to rap-
idly spread along drainage ways.
Habitat. The tolerance of tamarisk to many habitats may indicate that plants are actually
several species, ecotypes, or hybrids. They most commonly grow along river systems,
springs, lake shores, reservoirs, floodplains, and irrigation ditches where they have access
to water. The trees are absent, however, from areas susceptible to long-term inundation
582 n TREES

because they cannot survive

more than three months of stand-
ing water. As a phreatophyte,
tamarisk grows its roots down to
the water table but is not depen-
dent on groundwater. It grows
on many substrates but is
most commonly found on fine-
textured soils. Most soils are alka-
line. The plant tolerates extreme
salt concentrations, greater than
50,000 ppm of soluble salts, but
is usually found on soils with
700–15,000 ppm. Tamarisk can
form dense thickets where depth
to groundwater is less than 20 ft.
(6 m), but plants will be more
widely spaced where the water
table is further below the surface.
It grows best in full sun and
is shaded out by tall cotton-
woods or willows.
Bare sites suitable for germi-
nation of seedlings are created
when dams and reservoirs alter
stream flow. Floodplains and
reservoirs subject to receding
water expose bare land to colo-
nization. Flood control also
changes the dynamics of river
flow, preventing or changing
Except for a few northern states, smallflower tamarisk has a wide the season of large flows. Tama-
distribution. French tamarisk is concentrated in the western desert and risk is less common along rivers
mountain states and in the southeast. (Native range adapted from USDA that consistently flood in spring
GRIN and selected references. Introduced range adapted from USDA because the high water scours
PLANTS Database, Invasive Plant Atlas of the United States, and selected the banks free of seeds and
references.)
small plants. When land sup-
porting native riparian vegeta-
tion is converted to agriculture, bare sites become available for tamarisk seed germination.
Plants are found from below sea level to 6,600 ft. (2,000 m) elevation in Death Valley,
California. The tree survives both cold winters and summer temperatures as high as 107ºF
(42ºC). In extremely cold climates, plants die back to the ground in winter.
Reproduction and Dispersal. Reproduction is both sexual and asexual. Tamarisk easily
spreads by means of both seed production and root sprouts. One-year-old plants can flower
and set seed. Tamarisk produces thousands of tiny seeds from each flower, and one mature
plant can produce more than 500,000 seeds in one season. The seeds weigh very little and
have short hairs, which facilitates wind dispersal. Both wind and water can carry the seeds
long distances. They are also easily spread to other water systems via boats, fishermen,
 TAMARISK n 583

A. Trees become shrubby when branches sprout from the base. (Steve Dewey, Utah State University,
Bugwood.org.) B. Leaves of athel tamarisk (left) smoothly sheath the twig, compared with the scaly leaves of
other species (right). (Joseph M. DiTomaso, University of California-Davis, Bugwood.org.) C. Panicles of small
flowers grow on the ends of twigs. (Steve Dewey, Utah State University, Bugwood.org.)

trailers, and water ballast. Seeds are viable for about five weeks and can germinate 24 hours
after moistening. They have no dormancy requirements. Seeds produced later in the season
have a better germination rate. In order to survive their first year, seedlings need saturated
soil for the first few weeks, a high water table, and open, sunny conditions with no competi-
tion. Seedlings establish easiest on sandy soil where water recedes, such as reservoir margins
or stream banks, opening bare areas to colonization. Although tamarisk seedlings will die if
the soil dries within the first four weeks, they are more drought tolerant than willow seed-
lings. The slow rate of growth of tamarisk seedlings, less than 5 in. (12.5 cm) tall after eight
weeks, is offset by a high rate of establishment. Few seeds survive cooler winter tempera-
tures, meaning that no large seed bank exists in the soil.
Tamarisk vigorously reproduces by vegetative means. Both mature plants and roots grow
rapidly. Shoots can grow 10–13 ft. (3–4 m) in one season. In one year, lateral roots can
extend as much as 30 ft. (9 m), and the tap root can grow 16 ft. (5 m). Branches touching
moist soil will root. Burned or cut plants will produce numerous sprouts from the root
crown and roots. Buried stems or broken pieces readily root. Floods frequently damage
plants and distribute the pieces, and fragments may also be carried on boats or trailers.
Impacts. Because of its vigorous growth, which literally crowds out the natives, tamarisk
displaces native riparian species, such as willow, cottonwood, and mesquite. The plant not
only tolerates salty soils, but also increases the salinity. Salt taken up by roots deep in the soil
is exuded onto the leaves of the plant. As leaf litter accumulates, the surface soil becomes too
salty for native species. The plant is less invasive where rivers are not controlled, and displace-
ment of native species is greater where hydrological processes have been altered. Perhaps
because of its greater leaf area, tamarisk uses more water than natives, lowering the water table
and causing surface water to diminish or disappear. A mature tree is capable of using 300 gal.
(1,135 l) per day. Dense stands may also increase local sediment deposition, and it is also
speculated that tamarisk thickets may clog water channels and alter stream flow.
The value of tamarisk for wildlife varies according to many factors, including region or
watershed, density of the stand, and species. In some areas, such as the Lower Colorado
584 n TREES

River in California, biodiversity of birds and small mammals, as well as total populations, are
generally lower in tamarisk thickets than where native plant species exist. However, the trees
do provide nesting sites in areas that previously had no forest. While ground feeders and
seed-eating populations may benefit, fruit-eating birds, cavity nesters, and timber gleaners
generally require native vegetation. Although beaver eat young tamarisk shoots, they prefer
willows and cottonwoods, giving the tamarisk a competitive advantage. The relationship
with other rodents varies with species and location. The scaly leaves, however, are not palat-
able to most grazing animals, either domestic or wild.
Management. Because tamarisk is so widespread and disperses so readily, total eradica-
tion may not be possible except in small areas. Furthermore, the site may be permanently
damaged due to salt buildup or changes in hydrology. All methods are time-consuming,
labor intensive, and expensive. In 1998, complete restoration along a portion of the Rio
Grande in New Mexico ranged from $1,850 to $3,200 per acre ($750 to $1,300 per ha).
Because roots and pieces vigorously resprout, physical means such as cutting or bulldoz-
ing are ineffective if not used in conjunction with other methods. Removal of the aerial parts
of the plants, however, can reduce water consumption by as much as 50 percent. If bulldoz-
ing is used, plants should be plowed up below the root crown. Small sprouts may be pulled
by hand. Burning works best when 25 percent of the biomass is first cut and allowed to dry,
and is most successful in the hottest part of summer, when plants are most water stressed. It
may take three to four years or more to fully eradicate a tamarisk stand. Because tamarisk
fragments resprout vigorously, burning should be followed by spraying new sprouts with
a herbicide. Burning cannot be used where tamarisk is mixed with native cottonwoods
and willows because those species do not resprout and will be killed. Grazing is a poor
option because livestock prefer to graze the native cottonwood and willow seedlings rather
than the tamarisk.
Chemical control can be attempted in several ways, but care must be taken around water-
ways where tamarisk grows. Imazapyr, either alone or in conjunction with glyphosate, is the
most effective herbicide, affording 70–90 percent control. While glyphosate, 2,4-D,

Successful Eradication

C oachella Valley Preserve in southern California, home to 180 species of wildlife,

contains the only undisturbed watershed remaining in that valley. Originally
planted as a windbreak, tamarisk got out of control. Almost three-quarters of the
25 ac. (10 ha) wetland in Thousand Palms Canyon was 80 percent covered by tama-
risk, and the spring had dried up. Although it took several years, 1986–1992, and
5,000 person-hours, tamarisk was successfully eradicated from the site. Using pri-
marily hand labor provided by volunteers and California Conservation Corps crews,
tamarisk trunks were cut close to the ground and immediately sprayed with a her-
bicide. The debris was moved to other areas in the preserve where the wood piles
provided habitat for birds until the native riparian species regenerated. The spring
began flowing soon after the first tamarisk cutting. Seeds of native plants were
hand scattered in the area, and the canyon has been successfully restored.
Occasional tamarisk seedlings from wind-blown seeds are easily pulled out, pre-
venting re-infestation.
Source: Martin, 2001.
 TREE OF HEAVEN n 585

dicamba, or triclopyr will defoliate plants and reduce evapotranspiration and water use, they
do not kill the plants. Leaves will grow back. Aerial spraying of herbicides is useful in mono-
typic stands of tamarisk but cannot be used in areas that include native vegetation.
Herbicides can also be applied to freshly cut stumps or around the base of the tree on the
bark. Basal application, however, requires more herbicide and is less effective than the cut-
stump method. Systemic herbicide application is most effective in fall or winter, when the
plants are becoming dormant and moving resources to their roots.
Biological control may be accomplished by two imported insects. The saltcedar leaf beetle
(Diorhabda elongata) from China, west-central Asia, and the Mediterranean, was released in
several western states. Seven different ecotypes are being studied. Both adults and larvae
eat the leaves and defoliate the plant, and more than one generation develops during a sea-
son, two generations in cooler climates and three to four in warmer climates. More than
1,000 beetles on one plant can completely defoliate it. A mealybug (Trabutina mannipara),
native to Israel, eats tamarisk twigs.

Selected References
Carpenter, Alan T. “Element Stewardship Abstract, Tamarix ramosissima Ledebour, Tamarix pentrandra
Pallas, Tamarix chinensis Loureiro, Tamarix parviflora De Candolle.” Global Invasive Species Team,
Nature Conservancy, 1998. http://www.invasive.org/weedcd/pdfs/tncweeds/tamaram.pdf.
Jacobs, Jim, and Sharlene Sing. “Ecology and management of Saltcedar (Tamarix ramosissima, T. chinen-
sis and T. ramosissima x T. chinensis hybrids).” Invasive Species Technical Note No. MT-13. U.S.
Department of Agriculture, Natural Resources Conservation Service. Bozeman, MT, July 2007.
http://www.mt.nrcs.usda.gov/technical/ecs/invasive/technotes/invasivetechnotemt13/.
Kirk, McDaniel, and Mark Renz. “Saltcedar Information, Biology and Ecology.” Weed Information,
New Mexico State University, Las Cruces, n.d. http://agesvr1.nmsu.edu/saltcedar/Index.htm.
Martin, Tunyalee. “A Success Story: Tamarisk Control at Coachella Valley Preserve, Southern
California.” Wildland Invasive Species Program, Nature Conservancy, January 2001. http://www
.invasive.org/gist/stories/ca003/ca003.PDF.
Zouhar, Kris. “Tamarix spp.” In Fire Effects Information System, U.S. Department of Agriculture, Forest
Service, Rocky Mountain Research Station, Fire Sciences Laboratory, 2003. http://www.fs.fed.us/
database/feis/.

n Tree of Heaven
Also known as: Ailanthus, Chinese sumac, copal-tree, stinking shumac, varnish tree
Scientific name: Ailanthus altissima
Synonyms: Toxicodendron altissima, Albonia peregrina, Ailanthus glandulosa,
Ailanthus peregrina
Family: Quassia (Simaroubaceae)
Native Range. China, except the extremely arid central region.
Distribution in the United States. Common in the East and Midwest, from Maine south
to Florida, west to Texas, and north to Iowa and Wisconsin. Distribution in western and
Great Plains states is discontinuous, often in riparian zones, from New Mexico west to
California and north to Washington State. Also in Idaho and Nevada, east to Nebraska and
Kansas. Plants are often indicative of old mining sites or homesteads in California.
Description. Tree of heaven is a deciduous tree that grows 40–100 ft. (12–30 m) tall. It
has one main trunk and is not shrubby. The smooth gray bark is vertically streaked, becom-
ing fissured with age. The stout twigs form a broad, spreading lacy crown with few branches.
586 n TREES

Shoots are initially covered with

yellow-brown hairs, but old
twigs are hairless. Leaves,
which reach 4 ft. (1.2 m) long,
are alternate and pinnately
compound with 11–41 pointed
leaflets, including a terminal
leaflet. The lance-shaped leaf-
lets are 2–6 in. (5–15 cm) long.
Each leaflet has 2–4 rounded
teeth near its base, while the
rest of the leaflet has a smooth
margin. Each tooth on the leaf
base has a round gland on the
underside. The upper leaf sur-
face is deep green, while the
underside is grayish green.
When leaves fall, they leave a
prominent scar on the stem,
variously shaped like a heart,
horseshoe, or shield. Male flow-
ers and crushed foliage release
an unpleasant odor, variously
described as peanuts, cashews,
or rancid nut oil.
Depending on the soil tex-
ture, trees may have both a deep
tap root and long lateral roots.
The root system is strong and
aggressive.
In late spring or early sum-
Present in most states except the northern Great Plains, tree of heaven mer, April to July depending on
grows well in poor conditions, including nutrient-deficient soils, salt geographic location, small
spray, and drought. (Native range adapted from USDA GRIN and selected greenish star-shaped flowers
references. Introduced range adapted from USDA PLANTS Database, appear on 2.5–5 in. (6–12 cm)
Invasive Plant Atlas of the United States, and selected references.)
long panicles at the ends of new
branches. Flowers have five
overlapping sepals and five petals, 0.08–0.1 in. (2.0–2.5 mm) long. The base of each flower
is slightly hairy. Male and female flowers are produced on different trees. Fruits mature from
August to October and remain on the tree into winter. Fruits go through a color change as they
ripen, first green, then becoming yellow, pink or orange, red, and red-brown. The 1–2 in.
(2.5–5 cm) long dry fruits contain one small seed in the center of two flat wings. The fruit
masses hang down, and trees with no fruit are males.
Related or Similar Species. Several plants native to the eastern United States which have
large, compound leaves may be mistaken for tree of heaven. Native trees have serrated or
toothed leaflet margins, distinct from the smooth leaflet margins of tree of heaven.
Staghorn sumac may further be distinguished from tree of heaven by its fuzzy reddish-
brown branches and red, fuzzy fruits that are borne erect rather than pendant.
 TREE OF HEAVEN n 587

A. Tree of heaven is a tall tree with one main trunk. (Karan A. Rawlins, University of Georgia, Bugwood.org.)
B. The large compound leaves have many leaflets. (Paul Wray, Iowa State University, Bugwood.org.) C. The
trunk becomes fissured with age. (Annemarie Smith, ODNR Divison of Forestry, Bugwood.org.) D. Leaf scars
are prominent, with a distinct shape. (Daniel Herms, Ohio State Weed Lab Archive, The Ohio State University,
Bugwood.org.) E. Small star-shaped flowers grow on long panicles. (Jan Samanek, State Phytosanitary
Administration, Bugwood.org.) F. The base of each leaflet has 2–4 distinct teeth. (James H. Miller, USDA
Forest Service, Bugwood.org.) G. Each winged pod contains one seed. (Annemarie Smith, ODNR Divison of
Forestry, Bugwood.org.)

Introduction History. Tree of heaven has a long history in the United States. A gardener
in Philadelphia imported the plant in 1784, and by 1840, it was commonly available in
nurseries, probably because it was easy to grow under poor conditions. Chinese immigrants
introduced the tree to California during the gold rush in the 1890s.
Habitat. Tree of heaven is a fast-growing pioneer tree that generally needs sunlight, but
may tolerate some shade. The tree thrives in poor soils, including limestone, and tolerates
airborne salt and drought. It predominantly invades wastelands, seminatural areas, or dis-
turbed areas, including grassland, forest openings, riparian locations, and rocky outcrops.
In urban locations, it is commonly seen in disturbed sites such as alleys, sidewalks, parking
lots, and is so frequent that it is almost “natural.”
In rural areas, it is found in fields, fencerows, roadsides, forest edges and clearings. It fre-
quently forms thickets at the edges of forests and will fill in any clearings that develop. Due
to extensive root suckering, it can form dense thickets covering an entire acre. Tree of heaven
is also an agricultural pest. Seedlings by the hundreds can sprout in newly planted fields.
While older trees can resist freezing temperatures, plants younger than six years are sus-
ceptible to frost. Regardless of age, leaves fall at the first frost. In the western states, the tree
grows below 6,600 ft. (2,000 m) elevation.
Reproduction and Dispersal. Tree of heaven reproduces both sexually and vegetatively.
Plants flower early in their life, sometimes only six weeks after germination. Each female tree
may produce 325,000 seeds annually and live for 50 years. Dissemination of seeds is pri-
marily by wind, the wings on the seeds acting like small propellers. Seeds are also spread
by water, birds, vehicles, and machinery. Seeds remain viable for one year. They have a high
germination rate and develop a tap root within three months. If the soil is clay or compacted,
long lateral roots develop instead. Trees grow quickly in full sunlight, as much as 3.3 ft.
(1 m) per year for the first four years.
588 n TREES

When the trunk is cut, suckers resprout vigorously from the stump and from roots. Root
sprouts can emerge 50 ft. (15 m) from the mother plant, even through sidewalk cracks or
other concrete.
Impacts. Tree of heaven thickets displace natural vegetation, especially in riparian habi-
tats, and toxins from leaf litter prevent native plants from reestablishing. Litter is toxic to
35 species of gymnosperms and 10 species of angiosperms, severely limiting competition
and natural succession. In urban areas, the aggressive root system may damage sidewalks,
sewers, and foundations. Sucker growth poses maintenance problems for landscapers.
Management. Because this species vigorously resprouts, the most important management
aspect is to frequently monitor sites for regrowth and pull or treat new sprouts or seedlings
as they occur. Sites should be checked at least once a year, preferably more often. Targeting
large female trees will help to reduce seed production. Any physical means of removal must
involve the whole plant, because any root fragments will sprout. Small plants, especially seed-
lings, can be pulled or dug up. Although it is possible to remove entire seedlings, including
roots, removing the root from root suckers is not as simple. Chopping, cutting, or mowing
plants, either manually or mechanically, encourages sprouts. Cutting should be done in early
summer or just before flowering when root reserves are lowest. Burning the trunks or stems
with a flamethrower, which can be done during wet weather and snow conditions to avoid
wildfires, can be used for spot treatment. However, plants will resprout. Grazing will remove
tops of seedlings and sprouts, thereby preventing flowering. With repetition over several years,
grazing, cutting, or burning may deplete root storage and weaken plants.
Although no studies have been specific to tree of heaven, chemical applications provide
the most effective control. Herbicides may be applied to leaves, basal bark, or cut stumps.
Chemicals are especially necessary on cut stumps to prevent resprouting. It is easy to kill
the above-ground parts of the plants, but roots are much more difficult. Both glyphosate
and triclopyr are systemics, meaning that the chemicals are translocated to the roots, with
no residue in the soil. Because glyphosate is nonselective, it cannot be sprayed onto foliage
in mixed stands where native vegetation would also be damaged. In contrast, triclopyr is
selective; it will affect broadleaf and woody plants but will not harm grasses. Basal bark
application, which works well on trees less than 6 in. (15 cm) in diameter, is the easiest
method because no cutting is involved. It is most effective when done in late winter or early
spring. During late spring or early summer, plants are moving fluids upward to grow, not
down into the roots. An alternative is hack-and-squirt. After making down-angled cuts into
the trunk, squirt triclopyr into the cut. Girdling the trunk is not recommended without
follow-up herbicide treatment because it triggers a survival response of vigorous sprouting
and suckering in the plant. After a trunk is cut to a stump, herbicide must be applied within

Beneficial Uses of Tree of Heaven

B ecause of its tolerance for poor soils, tree of heaven has been used for reveg-
etating acidic mine spoils where the pH is less than 4.1. It is ideal for urban
and industrial areas because it tolerates pollution, including particulates like
cement dust and gases from the coke and coal-tar industry. The leaves absorb sul-
phur and mercury, and the plant is resistant to ozone. The book A Tree Grows in
Brooklyn, by Betty Smith in 1943, is based on Ailanthus.
 VELVET TREE n 589

5–15 minutes, before the plant seals off the area. It is labor intensive to cut trees before treat-
ment, unless the area must be cleared anyway.
Little research has been conducted on biological control of tree of heaven. However, a
potential exists in fungal pathogens (Verticillium dahliae and Fusarium oxysporum) found on
dead or dying plants in New York and Virginia. A possibility also exists in the use of the
zonate leafspot (Cristulariella pyramidalis) for defoliation.

Selected References
Hoshovsky, Marc C. “Element Stewardship Abstract, Ailanthus.” Global Invasive Species Team, Nature
Conservancy, 1988. http://invasive.org/gist/esadocs/documents/ailaalt.pdf.
Swearingen, Jil M., and Phil Pannill. “Tree of Heaven.” Weeds Gone Wild: Alien Plant Invaders of
Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2005. http://
www.invasive.org/gist/esadocs/documnts/ailaalt.pdf.
“Tree-of-Heaven.” Invasive Species Identification Sheet, U.S. Department of Agriculture Natural
Resources Conservation Service. Tolland, CT, 2002. http://www.ct.nrcs.usda.gov/tree-of
-heaven.html.

n Velvet Tree
Also known as: Miconia, purple plague, bush currant
Scientific name: Miconia calvescens
Synonyms: Cyanophyllum magnificum, Miconia magnifica
Family: Melastome (Melastomataceae)
Native Range. Central and South America, from southern Mexico at 20º N latitude to
northern Argentina and southern Brazil at 25º S latitude. It grows as high as 6,000 ft.
(1,830 m) elevation in Ecuador. The form with the very large leaves and purple undersides
is native only to Mexico and Central America, including Guatemala, Belize, and Costa Rica.
Distribution in the United States. The major islands of Hawai’i. Some reports of naturali-
zation in Puerto Rico.
Description. Velvet tree is a shrubby evergreen tree, with many slender branched stems
that grow vertically. The tree can reach 45 ft. (13.5 m) tall. The oval leaves, opposite on
the stem, can be very large, up to 3 ft. (1 m) long. Although they may appear velvety, leaves
have no hairs. In contrast to the dark-green upper surface of the leaf, the underside is an iri-
descent purple. Leaves are further distinguished by three prominent veins running from the
base to the tip, one in the center of the leaf and one closer to each leaf margin. These veins
are connected by many smaller veins running crosswise on the leaf. The root system is
shallow.
The inflorescence is a large erect panicle with 1,000–3,000 small white to pink or
tan flowers. The sweetly scented flowers each live for only 12–24 hours. The small,
0.25–0.5 in. (6–13 mm) diameter, dark-blue or purple fruit is sweet and attractive to birds.
Each fruit contains many small seeds, each about the size of a grain of sand.
Related or Similar Species. Eight additional members of the Melastome family are
present in Hawai’i, the worst being Koster’s curse or soap bush (see Shrubs, Koster’s
Curse), cane ti or cane tibouchina, and bristletips. Both cane ti and bristletips are much
smaller plants. Cane ti, native to Brazil, Paraguay, and Uruguay, is a branched upright shrub,
as tall as 9 ft. (2.7 m). Leaves are 3 in. (7.5 cm) long and hairy, with 5–7 prominent longitu-
dinal veins. The four-petaled flowers are pink with bright yellow anthers. A second shrub
590 n TREES

species of Tibouchina, glorybush

or princess flower, which is also
native to Brazil, has purple
flowers with five petals. Glory-
bush is less invasive because it
does not seem to spread by
seed. Native to the Himalayas
and southwestern China, bris-
tletips is a shrub 6.5–13 ft. (2–
4 m) tall with drooping
branches. Leaves are 3–6 in.
(8–16 cm) long with five paral-
lel veins. Leaves are generally
glabrous, but veins on the lower
surface are covered with stellate
hairs. Pinkish-purple flowers,
0.75 in. (2 cm) long, grow in
pendulous clusters.
Introduction History. Velvet
tree was deliberately brought to
the Wahiawa Botanical Garden
in central O’ahu in 1961, and
also to Waimea Botanical
Garden in northwestern O’ahu.
Both of those plantings were in
seasonally dry areas of the
island, with 59–65 in. (1,500–
1,650 mm) annual rainfall. A
tree was planted at Harold L.
Lyon Arboretum near Hono-
lulu in 1964, and another
Velvet tree is a very destructive invader of tropical rainforest habitats on tree was planted near the same
Pacific islands, including Hawai’i. (Native range adapted from USDA area in the late 1970s or early
GRIN and selected references. Introduced range adapted from USDA 1980s. In spite of the fact that
PLANTS Database, Invasive Plant Atlas of the United States, and selected the original trees were destro-
references.)
yed in the 1990s when the inva-
sive nature of the tree was real-
ized, velvet tree has become naturalized on O’ahu. A plant was introduced to a private
estate on the island of Hawai’i in the early 1960s, and a specimen was taken to a private
nursery and botanical garden on Mau’i in the early 1970s. Infestations on Kaua’i stem from
a single seedling imported from O’ahu. Velvet tree was sold in nurseries because of its attrac-
tive foliage until it was declared a Hawai’i noxious weed in 1992.
Habitat. Capable of growing in several habitats, including coastland, scrubland, and
riparian sites, velvet tree occupies the tropical montane zone in Hawai’i, where the forests
receive 70–80 in. (1,800–2,000 mm) of annual precipitation. Usually found between
1,000 and 5,900 ft. (300–1,800 m) elevation, it can grow as high as 6,500 ft. (2,000 m)
elevation, as well as in lower elevations, and is a threat to habitats up to the upper limit of
forest growth. It is quick to invade following deforestation due to fire, overgrazing, or other
 VELVET TREE n 591

A. Velvet tree has destroyed much of the natural forest in Tahiti. (The Nature Conservancy Archive, Bugwood.org.)
B. The large leaves have distinctive veins. (Forest and Kim Starr.) C. The inflorescence is a large panicle. (Forest and
Kim Starr.) D. Veins are prominent on the purple undersides of leaves. (Forest and Kim Starr.)

disturbances, but can also invade intact, healthy forests. It is especially invasive on fallow
land from abandoned sugarcane plantations, from where it can spread into adjacent upland
forests. In Tahiti, it rapidly invaded forests damaged by hurricanes. The tree prefers to grow
on mineral soil, dead tree trunks, and dead fern trunks, and is very shade tolerant.
Reproduction and Dispersal. Plants produce flowers and fruit when they are 4–5 years
old and about 12 ft. (3.7 m) tall. Each mature tree, taller than 26 ft. (8 m), can flower three
times a year, With 220 inflorescences, 200 fruits per inflorescence, and 50–200 seeds per
fruit, trees can produce 2–3 million seeds each time they flower. The soil seedbank is huge
and may be more than 5,000 seeds per sq. ft. (50,000 per m2). Seeds remain viable for 5–
10 years. Seeds are dispersed predominantly by birds, and somewhat by rodents, which eat
the fruit. Birds can carry thousands of seeds long distances, as far as 1.25 mi. (2 km). Seeds
are dispersed predominantly by nonnative bird species, such as the Japanese White-eye,
Common Myna (see Volume 1, Vertebrates, Birds, Common Myna), and the Red-billed
Leiothrix. Seeds are also spread by human activities, by adhering to shoes or clothing, machi-
nery, and vehicles, or being transported in soil or garden waste. Seeds can also cling to
hooves of livestock. Seeds may be spread on native tree fern logs that are transported between
islands. Seedlings can germinate in either sun or deep shade, with as little as 0.02 percent of
available sunlight. Moisture, however, appears to be a limiting factor. Seeds lose viability
when they become dry. Under good conditions, juvenile trees can grow 5 ft. (1.5 m) per year.
Impacts. Velvet tree is considered to be one of the most destructive invaders to tropical
rainforest habitats on the Pacific Islands, including Hawai’i. It is successful because it grows
rapidly, reaches maturity early, and produces abundant seeds that are widely dispersed. If
not stopped, it will threaten almost all the mesic and wet forests in Hawai’i that receive more
than 60 in. (1,500 mm) of annual rainfall. It alters habitat, changes hydrology, and may con-
tribute to soil erosion. Velvet tree forms dense monospecific stands that shade out and displace
native plants, decreasing the biodiversity of the ecosystem, threatening endangered species,
including plants, birds, and invertebrates. Velvet tree thickets threaten to replace ohia and
koa forests, endangering rare native Hawaiian birds and insects, which depend on that habitat.
592 n TREES

Velvet tree alters the hydrology. Dense thickets create an umbrella-like canopy over the
watershed, reducing infiltration and groundwater recharge, resulting in more runoff.
Because its roots are shallow, landslides and erosion are more likely where velvet tree displa-
ces native plants. With less infiltration of rainfall, more runoff increases erosion, which in
turn increases siltation, changing the quality of surface water. Estimates of costs due
to reduced groundwater recharge and to changes to water quality on O’ahu approach
$145 million per year. Loss of topsoil through soil erosion reduces nutrients and organic
matter, which in turn lowers agricultural productivity. Economic losses for the worst-case
scenario of damage to water and erosion on all the Hawaiian Islands are potentially
$380 million per year.
Management. Control or eradication of velvet tree requires a sustained effort. Because
seeds are easily carried in clothing and equipment, people should change clothing and
shoes after working in an infested area. All equipment, including anything that might carry
seeds, should be thoroughly cleaned. Physical control is possible with small trees or saplings,
less than 10 ft. (3 m) tall, that can be pulled or uprooted. Although it is unusual, uprooted
trees left lying on the soil may grow adventitious roots. Cutting down trees, or removing
the canopy, triggers germination of seedlings, which can be 40–100 per sq. ft. (400–1,000
per m2), all less than 2.3 ft. (0.7 m) tall, after 18 months of growth. These smaller trees
may then be sprayed with a herbicide, repeated in 2–3 years until the seed bank is exhausted.
Chemical control is the most effective. Triclopyr applied to foliage, cut stump, or basal
bark is best. When applied to cut stumps, triclopyr + 2,4-D has resulted in few resprouts.
In inaccessible areas such as steep slopes or rugged lava flows, aerial spraying of herbicides
has had 70 percent success on fruiting trees, but it is expensive and threatens nontarget
plant species.
A range of biological agents from Latin America that damage velvet tree, including fungi,
weevils, beetles, nematodes, wasps, butterflies, and moths, is being studied. Because Hawai’i
has no native plants in the Melastome family, it is more likely that introduced pests will be
host-specific to velvet tree. Chinese rose beetle (Adoretus sinicus) causes 50 percent defolia-
tion but does not kill the tree. A defoliating sawfly (Atomacera petroa), which attacks leaves,
has good potential because it is highly host-specific. A leaf spot disease caused by Cocostroma
myconae is also a possibility. A gall wasp (Allorhogas sp.) and a beetle (Apion sp.) that both
feed on the fruit are being evaluated in Brazil for host preferences. A fungus
(Colletotrichum gloesporioides f. sp. miconiae), which causes leaf spotting and leaf drop, is
now present on Hawai’i and Mau’i.

Velvet Tree in Tahiti

V elvet tree was introduced to Tahiti in 1937, possibly from Sri Lanka by the bot-
anist J. F. Rock who worked extensively in botanical gardens in Asia. It now
dominates over 65 percent of the island’s forest cover, and has displaced native spe-
cies due to its deep shade-producing foliage. Tahiti and Hawai’i are similar in
topography and climate, both being volcanic islands with heavy rainfall at a similar
distance from the equator. If not checked, velvet tree has the potential to severely
and irreparably alter Hawai’i’s botanical landscape as it has done to Tahiti.
 VELVET TREE n 593

Selected References
“HNIS Report for Miconia calvescens.” A product of the Hawaiian Ecosystems at Risk Project (HEAR),
1997. http://www.hear.org/hnis/reports/HNIS-MicCal.pdf.
Medeiros, A. C., L. L. Loope, P. Conant, and S. McElvaney. “Status, Ecology, and Management of the
Invasive Plant, Miconia calvescens DC (Melastomataceae) in the Hawaiian Islands.” Records of the
Hawaii Biological Survey for 1996. Bishop Museum Occasional Papers 48: 23–36, 1997. http://
hbs.bishopmuseum.org/pdf/melastome97.pdf.
Motooka, P., L. Castro, D. Nelson, G. Nagai, and L. Ching. “Miconia calvescens.” In Weeds of Hawai’i’s
Pastures and Natural Areas: An Identification and Management Guide. College of Tropical Agriculture
and Human Resources, University of Hawai’i, Manoa, 2003. http://www.ctahr.hawaii.edu/invweed/
WeedsHI/W_Miconia_calvescens.pdf.
National Biological Information Infrastructure (NBII) & IUCN/SSC Invasive Species Specialist Group
(ISSG). “Miconia calvescens (Tree).” ISSG Global Invasive Species Database, 2006. http://www.issg
.org/database/species/ecology.asp?fr=1&si=2.
Wise, Andrea, and Robert E. Lyons. “Velvet Tree.” Weeds Gone Wild: Alien Plant Invaders of Natural
Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/
alien/fact/mica1.htm.
n Vines
n Chocolate Vine
Also known as: Five-leaf akebia
Scientific name: Akebia quinata
Synonyms: Rajania quinata
Family: Lardizabala (Lardizabalaceae)
Native Range. East-Central China to Korea and Japan.
Distribution in the United States. Eastern and midwestern states, from New York and
Massachusetts south to Georgia, west to Louisiana, and north to Missouri, Illinois, and
Michigan.
Description. Chocolate vine is a vigorous trailing ground cover or climbing woody vine
that grows fast. The vines climb by twisting or twining stems around the host, and their
height is restricted only by the structures or plants they climb. Plants are deciduous in cooler
climates, but evergreen where winters are warmer. The slender round stems are green when
young, but mature to grayish brown, with small, round lenticels. Bud scales are light reddish
brown. Palmately compound leaves are alternate on the stem. Five or fewer oval leaflets,
1.5–3 in. (4–7.5 cm) long and slightly notched at the tip, grow on small stalks that merge
in the center of the compound leaf. Leaflets are dull blue-green with a purplish tinge that
turns to blue-green when the leaves mature.
Flowering occurs from late March to early April. The reddish to chocolate-purple color
flowers, 1 in. (2.5 cm) in diameter, are fragrant, with a vanilla or chocolate scent. Both male
and female flowers grow on racemes from leaf axils. The two slightly smaller flowers at the
end of the racemes are male, while the remainder are female. The ball-shaped buds unfold
three broad sepals to form a cup holding the inner flower parts. Fruit ripens from late
September to early October. The purple-violet seed pods resemble 2.5–4 in. (6–10 cm) long,
flattened sausages. Pods split when mature, revealing a whitish pulpy interior with many
tiny black or brownish seeds arranged in irregular rows.
Related or Similar Species. A related species with three leaflets, three-leaf akebia, is
also native to China. A hybrid occurring in Massachusetts, which is a cross between
five-leaf akebia and three-leaf akebia, has 3–5 leaflets.
Two species of Aristolochia common in the southern and eastern part of the United States
are also climbing vines. Both pipevine and wooly Dutchman’s pipe are easily distinguished
by having simple (not compound) heart-shaped leaves and flowers that resemble a pendu-
lant curved pipe.
Introduction History. Chocolate vine was introduced to the United States in 1845 as an
ornamental, for its attractive purplish foliage. Because it rarely sets seed, its escape from cultiva-
tion, especially over long distances, was probably aided by humans. Both species and the
hybrid, as well as cultivars with different color flowers, are available through Internet nurseries.
Habitat. Chocolate vine grows in many habitats, including riparian, urban, and wetland sites,
but it also tolerates drought. Plants grow best in partial shade, especially on northern aspects, but
they can also do well in full sun. Because it is shade tolerant, the vine can invade healthy forests. It
 CHOCOLATE VINE n 595

grows on various soil textures,

from light to heavy, including
sandy, loamy, and clay. Plants
prefer well-drained but moist
soils that may be either acid or
alkaline. Although dormant
plants are hardy to −4ºF (−20ºC),
young spring growth will be
killed by frost. In its native
China, chocolate vine grows
from 1,000 to 5,000 ft. (300–
1,500 m) elevation.
Reproduction and Dispersal.
Reproduction of chocolate vine
is primarily vegetative because
vines do not self-pollinate well.
Both fruit and flowers are
uncommon outside of cultiva-
tion. Vines, which root where
they touch the ground, can
grow as much as 20–40 ft. (6–
12 m) in one growing season.
Birds eat the fruit and disperse
the seeds, but most spread of
chocolate vine is done by hu-
mans. Seeds germinate within
1–3 months at temperatures of
59ºF (15ºC). Plants require five
years to mature to flowering
and fruiting stage, but the life
span is unknown.
Impacts. Chocolate vine nat- This shade-tolerant vine, which thrives in a variety of habitats, can in-
uralizes easily and can overrun vade healthy forests, especially in the mid-Atlantic states. (Native range
and smother ground-level adapted from USDA GRIN and selected references. Introduced range
herbs, seedlings, saplings, and adapted from USDA PLANTS Database, Invasive Plant Atlas of
the United States, and selected references.)
understory shrubs because of
its rapid growth. Its dense
growth, a mat of impenetrable vines, prevents the germination and establishment of native
plants by depriving them of light, water, nutrients, and space. If left unchecked, plants can
even kill canopy trees by overtopping and smothering them.
Management. Because chocolate vine will sprout from its roots, any control requires
follow-up to eliminate resprouts. Physical control may be accomplished by repeatedly cutting
the stems during the growing season to exhaust nutrient reserves in the roots. If available per-
sonnel and money allow only one cutting to be done, plants should be cut to the ground at the
end of summer. Groundcover vines may be either left on site or disposed of elsewhere.
Detached vines will not root and grow. Vines that climb trees or buildings may also be severed
from the roots. Remaining roots from either growth type must be either dug out or treated with
herbicides to prevent resprouting. Small plants, including the root, can be totally removed.
596 n VINES

A. Vines climb into the canopy of trees. (Chris Evans, River to River CWMA, Bugwood.org.) B. Compound leaves
have five oval leaflets. (James H. Miller, USDA Forest Service, Bugwood.org.) C. Small flowers have three sepals.
(Nancy Loewenstein, Auburn University, Bugwood.org.) D. Young vines are slender. (James H. Miller, USDA
Forest Service, Bugwood.org.) E. Older vines are thicker, with lenticels. (James H. Miller, USDA Forest Service,
Bugwood.org.) F. Seed pods resemble small sausages. (James H. Miller, USDA Forest Service, Bugwood.org.)

Mulching is effective for small areas of trailing chocolate vine on the ground or where
herbicides cannot be used. Plants can be covered with several inches of any biodegradable
material, such as grass clippings, hay, or wood chips. Hay and grass clippings, however,
may be contaminated with other weed seeds. An additional covering of cardboard over the
mulch will impede its decay and prolong its life. Mulching should be done for two growing
seasons, refreshing the cover as necessary. Mulching can also be done after herbicide treat-
ment as an extra precaution against regrowth.
Chemical application of herbicides by spraying the foliage is practical for large stands
where soil disturbance caused by uprooting plants would be too disruptive to the site.
Triclopyr ester is a more effective foliar spray than is glyphosate, and is also absorbed
through any bark it reaches. To avoid damage to native species, spraying should be done
before native wildflowers emerge. Triclopyr ester applied to basal bark, after vines have been
stripped away to allow access, affords good control, but care should be taken to avoid drip-
ping the herbicide on the bark of the host tree. Stems of climbing vines may be cut close to
the ground and again slightly higher to ensure a complete separation of roots from the
climbing vines. The severed vines will die, rot, and fall off the tree or structure. Apply triclo-
pyr amine or glyphosate to the cut stems to protect against sprouts.
No biological controls are currently available for chocolate vine. Four species of fungus
(Microsphaera akebiae, Microsphaera penicillata, Muyocopron smilacis, and Aecidium akabiae)
are associated with chocolate vine in its native China. Three moths (Evecliptopera decurrens,
Ophideres fullonica, and Archips asiaticus) can damage the plant. The most common, however,
is Ophideres fullonica, a fruit-piercing moth, which is a serious pest in orchards.

Selected References
National Biological Information Infrastructure (NBII) & IUCN/SSC Invasive Species Specialist Group
(ISSG). “Akebia quinata (Vine, Climber).” ISSG Global Invasive Species Database. 2005. http://
www.issg.org/database/species/ecology.asp?si=188&fr=1&sts.
 CLIMBING FERNS n 597

Uses of Chocolate Vine

C hocolate vine, including stems, roots, and fruit, is an ancient traditional medici-
nal plant in China. The sweet, edible fruit is used in wine making, and the soft,
young shoots may be eaten either pickled or raw in salads. Leaves are steeped for a
type of tea. When ingested, some plant parts control bacterial and fungal infec-
tions, such as those of the urinary tract. The plant is also used to induce menstrua-
tion and lactation and for reducing fever. Oil from the seed is used for making
soap.

Swearingen, Jil M., Adrienne Reese, and Robert E. Lyons. “Fiveleaf Akebia.” Weeds Gone Wild: Alien
Plant Invaders of Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2006.
http://www.nps.gov/plants/alien/fact/pdf/akqu1.pdf.
Zheng, Hao, Yum Wu, Jianqing Ding, Denise Binion, Weidong Fu, and Richard Reardon. “Akebia qui-
nata.” In Invasive Plants of Asian Origin Established in the United States and Their Natural Enemies. U.S.
Department of Agriculture, Forest Service. FHTET-2004-05, 2004. http://www.invasive.org/weeds/
asian/akebia.pdf.

n Climbing Ferns
Japanese Climbing Fern
Also known as: No other names
Scientific name: Lygodium japonicum
Synonyms: Ophioglossum japonicum
Old World Climbing Fern
Also known as: Small-leaf climbing fern
Scientific name: Lygodium microphyllum
Synonyms: Lygodium scandens, Ugena microphylla
Family: Climbing Fern (Lygodiaceae)
Native Range. Japanese climbing fern is native to eastern Asia, from India east through
Southeast Asia, eastern China, Japan, Korea, the Philippines, and Indonesia east to New
Guinea. Old World climbing fern is from the Old World Tropics, sub-Saharan Africa,
India, and thoughout Southeast Asia, southeastern China, the Philippines, Indonesia east
to New Guinea, northern Australia, and some western Pacific islands.
Distribution in the United States. Japanese climbing fern is found in the southeastern
states, from Texas east to Florida, and north to North Carolina. Also in Pennsylvania,
Hawai’i, and Puerto Rico. Old World climbing fern is limited to southern Florida.
Description. Although both species are ferns, with distinct terminology for their vegeta-
tive parts, their overall structure resembles stems and leaves. Both climbing fern species
are vine-like perennials that climb onto and over shrubs, trees, or structures. The slender
twining stems, which may be 90 ft. (30 m) long, are green, orange, or black. All stems are
wiry and difficult to break. Although both species have light-green compound leaves, oppo-
site on the stem, the leaves are different.
Japanese Climbing Fern: Leaves of Japanese climbing fern are doubly compound and vary in
appearance. The overall shape of the compound leaf is triangular, 3–6 in. (8–15 cm) long and
598 n VINES

2–3 in. (5–8 cm) wide. Leaflets

grow on stalks and are lobed.
Terminal lobes of the leaflets
are often very finely dissected,
appearing lacy or feathery, while
the basal lobes on the same leaf-
let may be either dissected or
irregularly lobed. The lower sur-
face of the leaves is pubescent,
with short curved hairs. In cli-
mates where temperatures drop
below freezing, leaflets die in
winter, but they may remain
green in sheltered places. The
dry, brown fronds and stems
remain in place, providing a trel-
lis for new growth the following
season. Slender, creeping rhizo-
mes, which are actually the fern
stems, are dark brown or black.
They form a mat, 0.4–1.2 in.
(1–3 cm) below the surface.
While leaflets on the lower
part of the stems are sterile and
toothed, leaflets higher on the
stem become successively more
fertile as the vine grows longer.
The fertile leaves, or fronds, are
usually smaller. The leaf mar-
gins of fertile fronds have lobe-
like projections containing two
Japanese climbing fern, not considered a wetland species, commonly rows of sporangia (structures
invades open forests and roadsides in the southeastern states. (Native enclosing spores). Slightly roll-
range adapted from USDA GRIN and selected references. Introduced ed leaf margins partially cover
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the the sporangia. The nearly mic-
United States, and selected references.) roscopic spores are dispersed
during late summer and early
fall.
Old World Climbing Fern: Leaves of Old World climbing fern are once compound, 2–5 in.
(5–12 cm) long, with an oblong outline. Leaflets are leathery, usually not lobed, and gla-
brous or waxy on the lower surface. When the leaflet is detached, the wiry stalk remains
attached to the stem. Restricted to warmer climates, Old World climbing fern does not die
back in winter, continuing to produce vegetative growth and fertile leaflets. A dense mat of
rhizomes may be 3 ft. (1 m) or more thick on the top of the soil. Fertile leaflets are the same
size as other leaflets, with a tiny lobed fringe of slightly folded leaf tissue covering sporangia.
Sterile fronds have entire margins.
Related or Similar Species. Growing in wetlands throughout most states east of the
Mississippi River, the rare American climbing fern is the only Lygodium species native to
 CLIMBING FERNS n 599

North America, where it is threat-

ened or endangered. Its dis-
tinctly palm-like leaflets are
easily recognized by the 5–7
fingerlike lobes.
Introduction History. Both
species have been cultivated in
Florida as garden plants. A spe-
cies of climbing fern, advertised
as Old World but probably
Japanese, was offered for sale
in a nursery catalog as early as
1888. Japanese climbing fern
was known to be naturalized in
Georgia by 1903. The species
is still recommended as a gar-
den ornamental and sold on
the Internet. Although the date
and manner of introduction of
Old World climbing fern is not
known, the first naturalized
specimen was collected in
Florida in 1965. Since that
time, the plant has spread rap-
idly to cover thousands of acres,
including several counties and
the Everglades, and is expand-
ing its range northward.
Habitat. Both plants are lim-
ited to warm temperate or tropi-
cal areas, usually growing on
damp or disturbed sites, in Old World climbing fern, restricted to warm climates, is a major threat to
either sun or shade. Little is the Everglades ecosystem. (Native range approximated from USDA GRIN
known regarding specific habi- and selected references. Introduced range adapted from USDA PLANTS
tat requirements. Neither spe- Database, Invasive Plant Atlas of the United States, and selected
cies grows in very dry soils or references.)
under long-term flood condi-
tions, but both tolerate fluctuating water levels. Infestations in remote environments in the
Everglades indicate that neither species requires human disturbance to become established.
Although both are found in somewhat similar environments, Japanese climbing fern is
not considered to be a wetland species. It is commonly found in yards and along roadsides,
but also in moderately disturbed areas in open forests and at the margins of wetlands, such
as swamps, lakes, and creeks. It may also be scattered in tree plantations.
Old World climbing fern is found in both wetland and mesic habitats and is common in
bald cypress swamps, hammocks, pine flatwoods, wet prairies, mangrove communities, and
disturbed sites. Populations of Old World climbing fern in sawgrass marshes grow on
slightly elevated mounds or on tree trunks or rotting logs. This species seems to be limited
in its northern extent by minimum winter temperatures of 39–44ºF (4–6.5ºC). The potential
600 n VINES

A. Infestation of Old World climbing fern. (Amy Ferriter, State of Idaho, Bugwood.org.) B. Foliage of Japanese
climbing fern. (Chris Evans, River to River CWMA, Bugwood.org.) C. The ferns climb by twining their stems
around objects. (Richard Old, XID Services, Inc., Bugwood.org.) D. Fertile frond of Japanese climbing fern.
(Richard Old, XID Services, Inc., Bugwood.org.) E. Pieces can be dispersed in pine straw bales collected from
areas where Japanese climbing fern exists. (Dennis Teague, U.S. Air Force, Bugwood.org.) F. Fertile frond of
Old World climbing fern. (Amy Ferriter, State of Idaho, Bugwood.org.)

for spores to be transported northward along the moderate climate regions of the Atlantic
coast and westward along the Gulf coastline would greatly increase its range.
Reproduction and Dispersal. Both climbing ferns reproduce by wind-dispersed spores.
Spores may also be carried long distances as a component of dust on vehicles and other
objects. Although spores may germinate in 6–7 days, dried spores are able to germinate after
two years. The young fern initially resembles a small liverwort before it grows leafy fronds.
The creeping rhizomes enable the plant to spread locally. Plants resprout after winter frosts
and after vines are burned.
Impacts. Both species grow as either individual plants or tangled masses of stems and
leaves that cover the ground, shrubs, and even tall trees. They form dense mats of old and
new stems and fronds that smother native vegetation, lowering the biodiversity of the com-
munity. The dense canopy blocks sunlight, with mats as thick as 10 ft. (1 m) that do not
allow native plants to grow through. Hammocks in the Everglades may be so engulfed by
climbing fern that the plants beneath cannot be seen.
Both species intensify fires due to the added fuel load and, because of their climbing
habit, carry fire into the tree canopy. Native bromeliads growing on tree trunks are
destroyed. The dry leaflets of Japanese climbing fern and wind-blown burning fragments
are additional fire hazards.
Japanese Climbing Fern: Japanese climbing fern threatens three species on the Florida
endangered list, a perennial shrub called Georgia bully, wooly Dutchman’s pipe (a vine),
and branched tearthumb (a forb). Although the effect on wildlife is believed to be signifi-
cant, no data are yet available.
Old World Climbing Fern: Because Old World climbing fern is restricted to warmer cli-
mates and does not die back in winter, the accumulation of its stems and leaves can grow
to massive proportions. It can form a monoculture in cypress swamps and other wet areas,
where it displaces all the native plant and animal species, completely disrupting the eco-
system. Several rare plants, such as tropical curlygrass fern, giant air plant (also called
giant wild pine), and other bromeliads, are severely threatened as Old World climbing
fern invades their last remaining habitats. Because of its potential to spread, not just in
Florida but to other subtropical areas of the Gulf coast and into Mexico and Central
America, Old World climbing fern is considered to be a serious danger to subtropical
ecosystems.
 CLIMBING FERNS n 601

Old World climbing fern is a serious problem in Florida pine plantations, where it not only
complicates harvesting, but may also become a contaminant in pine straw. The use of pine straw,
which is freshly fallen pine needles, in landscaping may inadvertently introduce both spores and
fragments of Old World climbing fern into home gardens and new regions. In one investigation,
approximately 25 percent of the pine straw bales sold in nurseries were contaminated.
Economic costs of eradicating the ferns from natural areas or managed lands are high.
Control costs in 2000 were $135–500 per acre ($330–1,250 per ha), but one application
in a remote location cost $1,520 per acre ($3,750 per ha). The climbing ferns, however,
grew again, and the area had to be retreated in 2001.
Management. Areas should be consistently monitored to prevent new infestations. In
existing sites, prevention of spore formation and dispersal is important to minimize the
spread of climbing ferns into new areas. Because the tiny spores are easily transported, con-
trol activities should be undertaken at times of the year when spores are not being produced
or dispersed. If work must be done at that time, avoid driving equipment through the fern
area. To reduce the risk of spreading spores or plant pieces, workers should not go to other
areas on the same day.
Physical efforts on small plants or small areas may be done by hand-pulling or cutting,
but new vines will grow from the rhizomes. Machines can be used to pull large mats off veg-
etation, but doing so may damage the understory soil and vegetation. Fire is a poor option,
not only because new stems will grow from the rhizomes, but the climbing vines will pull
the fire into tree canopies and do more damage.
Little research has been conducted on chemical controls. Glyphosate, imazapyr, and met-
sulfuron methyl are all effective, either in combination or alone. Imazapyr causes the greatest
damage to nontarget species, including large hardwood trees. Herbicide applications are
best done during July through October.
Because only one species of Lygodium is native to the United States, and in a different cli-
mate zone, the possibility is good of finding a biological agent that is host-specific. Searches
have been conducted in most of the native ranges of the alien climbing ferns. A West Indies
Lygodium species, from Cuba, the Dominican Republic, and Haiti, may be an additional
source. Most studies have been done on Old World climbing fern, including testing for pos-
sible damage to 40 species of threatened or endangered ferns in Florida. A South American
rust (Puccinia lygodii), which has been found on Japanese climbing fern in northern Florida,
has been studied in greenhouse settings, but it is unknown whether the rust occurs in the
region infested by Old World climbing fern. The moth Neomusotima fuscolinealis is a native pest
on Japanese climbing fern in Japan, but because it also attacks American climbing fern, it is
unsuitable. A related tropical moth from northern Australia and Southeast Asia
(Neomusotima conspurcatalis) defoliates plants but does not pose a threat to American climbing
fern because it cannot tolerate cooler temperatures. It was cleared for release in 2007. In 2004–
2005, a defoliating moth (Austromusotima camptozonale) from Australia was released in several
locations in Florida, but did not become established. Additional possibilities include a gall
mite (Floracarus perrepae), lygodium saw fly (Neostrombocerus sp.), flea beetles (Manobia sp.),
and stem borers.

Selected References
MacDonald, Greg, Brent Sellers, Ken Langeland, Tina Duperron-Bond, and Eileen Ketterer-Guest.
“Japanese Climbing Fern.” Center for Aquatic and Invasive Plants, University of Florida, 2009.
http://plants.ifas.ufl.edu/node/639.
602 n VINES

The Genus Lygopodium

T he genus Lygopodium is ancient, its ferny fronds easily recognizable in the fossil
record from the Upper Cretaceous geologic time period, contemporaneous
with dinosaurs.

Miller, James H. “Japanese Climbing Fern.” In: Nonnative invasive plants of southern forests: a field guide
for identification and control. Gen. Tech. Rep. SRS-62, U.S. Department of Agriculture, Forest
Service, Southern Research Station, Asheville, NC, 2003. http://www.invasive.org/eastern/srs/
JCF.html.
Minogue, Patrick, Stella Jones, Kimberly K. Bohn, and Rick L. Williams. “Biology and Control of
Japanese Climbing Fern (Lygodium japonicum).” Publication #FOR 218. University of Florida IFAS
Extension, n.d. http://edis.ifas.ufl.edu/fr280.
Pemberton, R. W., J. A. Goolsby, and T. Wright. “Old World Climbing Fern.” In Biological Control of
Invasive Plants in the Eastern United States, by R. Van Driesche et al. USDA Forest Service
Publication FHTET-2002-04, 2002. http://wiki.bugwood.org/Archive:BCIPEUS/Old_World
_Climbing_Fern.

n English Ivy
Also known as: No other names
Scientific name: Hedera helix
Synonyms: None
Family: Ginseng (Araliacaeae)
Native Range. Western Europe, from Scandinavia south to the Mediterranean Sea, and
east to the Ukraine, Turkey, and the Caucasus region. Possibly the Mediterranean countries
of northern Africa and the Canary Islands and Madeira Islands.
Distribution in the United States. Western states, Arizona, Utah, and Idaho and the
Pacific coast. Midwestern, southern, and eastern states, from Texas and Oklahoma east to
Florida, north to Maine, and west to Wisconsin, Illinois, and Missouri. Also in Hawai’i.
Description. English ivy has two growth forms. In the juvenile stage, it is an evergreen,
perennial vine growing as long as 100 ft. (30 m), or as tall as the structures or trees on which
it climbs. When mature and able to produce flowers and fruit, after about 10 years, English
ivy becomes a woody shrub. Without something to climb on, the young stems form a trailing
ground cover, rooting at leaf nodes. Leaf shape is variable, depending on the age of the plant,
with the three-lobed form being most commonly recognized. The simple, alternate leaves
are a deep, glossy green, with a waxy or almost leathery cuticle. They exude an odor
when crushed. Leaves on young vines are palmately three- to five-lobed and heart-shaped,
1.5–4 in. (4–10 cm) long and similar in width. The terminal lobe is the largest, while the
two at the base are smaller or even absent. Climbing stems have more lobes and lighter-
colored veins. Plants become adults when they reach the reproductive phase. They grow a
woody, erect stem from either the ground or from a climbing vine and become a shrub.
Leaves on shrubby stems are a lighter green, oval, and not lobed. Young shoots and leaves
are hairy and scaly, but leaves of older plants are glabrous. In both stages, the petioles are
as long as the leaf. Plants grow fastest in the vine stage and more slowly in the shrub stage.
 ENGLISH IVY n 603

Roots in the soil are shallow

and not extensive. Adventitious
rootlets at leaf nodes enable the
plant to climb vertical surfaces,
such as walls and tree trunks.
However, the rootlets do not
damage the host. They exude a
glue-like substance that allows
them to adhere to, but not pen-
etrate, surfaces. English ivy
vines grow up buildings, other
structures, and trees, but do
not strangle trees and are not
parasitic.
Plants will flower and set
seed if they receive enough
light. Bloom time is autumn.
Flowers emerge in umbel clus-
ters, either solitary or on race-
mes, on the ends of branches
that extend at right angles from
the climbing vines. Flowers are
radial, 0.2–0.3 in. (5–7 mm),
with five white or yellowish-
green petals. Fruit is a berry-
like drupe, dark red or purple
to black, 0.25–0.35 in. (6–
9 mm). Each contains 1–5 hard,
stone-like seeds. If not eaten by
birds, fruit may remain on the
vine throughout the winter.
Because it is evergreen, English With the exception of wet areas, English ivy thrives in most
ivy grows year-round, although environments, including open forests and meadows. It is absent from
more slowly during the cooler most of the mountain and northern Great Plains states. (Native range
winter months. approximated from USDA GRIN and selected references. Introduced
Related or Similar Species. range adapted from USDA PLANTS Database, Invasive Plant Atlas of the
United States, and selected references.)
English ivy has hundreds of
horticultural varieties. Native
climbers that may appear similar include wild grape and Dutchman’s pipe vine. Wild grape
plants are woody vines with climbing tendrils and large three-lobed leaves with toothed
margins. Dutchman’s pipe is a deciduous woody vine that has heart-shaped leaves with
smooth margins.
Nonnative, sometimes invasive, species may appear similar to English ivy. Plants of the
cinnamon vine, also called Chinese yam, climb by twining, and its leaves are opposite and
arrow-shaped. Porcelainberry (see Vines, Porcelainberry) is a woody, deciduous vine that
climbs with tendrils. Although its leaves are alternate and deeply lobed, they are usually
toothed, resembling leaves of wild grape. Boston ivy also resembles English Ivy, but it is a
deciduous vine. Atlantic ivy, also called Irish ivy, has leaves as large as 4 in. (10 cm) wide,
604 n VINES

A. English ivy can cover and replace all other vegetation. (David J. Moorhead, University of Georgia,
Bugwood.org.) B. Roots at leaf nodes enable the plant to climb. (Chuck Bargeron, University of Georgia,
Bugwood.org.) C. Vines can smother the host tree. (James H. Miller, USDA Forest Service, Bugwood.org.)
D. When they become a shrub, plants are mature and able to produce flowers. (Jan Samanek, State
Phytosanitary Administration, Bugwood.org.) E. Fruit are small, berrylike drupes. (Forest and Kim Starr, U.S.
Geological Survey, Bugwood.org.)

covered with yellowish-brown or rusty-brown hairs. Leaves of colchis ivy, also called
Persian ivy, may be wider than 4 in. (10 cm). They are dull green on the upper surface
and have scale-like hairs. Poison ivy has some similar characteristics with English ivy, but
it is deciduous, has compound leaves with three leaflets, and whitish berry clusters.
Distinguishing characteristics of English ivy are that the evergreen leaves are palmate and
dark green with a waxy cuticle.
Introduction History. English ivy was probably introduced into North America as an
ornamental in early colonial times. It was, and remains, a popular ornamental because it is
a dependable, evergreen groundcover requiring little maintenance. It is very cold hardy
and pest free, and is still widely sold in plant nurseries. It was also used for erosion control
because of its propensity to root at leaf nodes, although its shallow root system renders it
ineffective. It kills other ground-growing plants, leaving a bare soil surface that offers little
resistance to water flow.
Habitat. Widespread in woods and rocky locations in its native range, English ivy grows
from sea level to 3,300 ft. (1,000 m). In North America, it invades open forests and forest
edges, fields, hedgerows, and the margins of coastal salt marshes, often after some type of
natural or human-induced disturbance. It prefers moist deciduous forest, but can grow
almost anywhere, in a variety of exposures, moisture regimes, and slopes. Although it is tol-
erant of various conditions, it does not grow where the water table is high or in wet areas. It
grows in both alkaline and acid soils but prefers slight acidity, a pH of 6.5. Although it is tol-
erant of shade, where it will grow and spread, it grows more vigorously and will flower in
full sunlight.
Reproduction and Dispersal. English ivy reproduction is primarily vegetative, but it also
produces abundant seeds. Plants produce many adventitious roots, from each leaf node, and
any stem fragment in contact with soil will sprout. Flowers are pollinated by flies and bees in
the fall, when few other plants are flowering. Birds eat and disperse the seeds, of which
approximately 70 percent are viable. The seeds have a hard coat that must be scarified, usu-
ally in the bird’s digestive tract. However, the toxic glycosides in the seeds often cause the
birds to vomit the seeds.
 ENGLISH IVY n 605

Impacts. English ivy is a serious problem in some areas on the West Coast and in wood-
lands on the East Coast, especially from Virginia to New York. It is very common near urban
areas. By covering plants and trees, it alters or halts normal succession and threatens all
layers in forests and in open areas, both ground plants and canopy. It inhibits generation
of understory native plants by shading the soil and preventing herbs and seedlings from
growing, with the result that the forest ecosystem cannot maintain itself. When English
ivy grows up tree trunks, especially in deciduous forests where more light is available for
winter growth, it shades out the tree canopy, killing the tree within a few years. In addition,
the weight of the vines can increase storm damage during periods of high wind, rain, snow,
or ice. Trees heavy with English ivy also may pose a hazard to people or structures. Wildlife
is affected by the altered ecosystem, and the leaf litter changes the nutrient content of
the soil.
Toxic compounds, glycosides, in the berries and leaves cause vomiting, diarrhea, nervous
disorders, and dermatitis in sensitive individuals. Some symptoms are more serious, includ-
ing breathing difficulty, coma, fever, muscular weakness, and lack of coordination.
English ivy is host for a pathogen, bacterial leaf scorch (Xylalla fastidiosa), which affects
native and ornamental trees, such as elms, oaks, and maples.
Management. Most research regarding English ivy has been in the development of culti-
vars, not in its control as an invasive species. The best control advice is to not plant it.
Pruning and hand-pulling up vines is an effective physical control method, especially for
small populations. Workers should take precautions to protect themselves from the plant’s
potentially toxic effects. Hand-pulling or cutting can be effective in areas where herbicide
use is impractical or impossible. The entire plant, including roots and runners, must be
removed. It is especially important to remove the vines from trees because the flowers and
seeds grow on the upright parts of the vine. Because stem fragments can sprout if they con-
tact soil, cut portions should be bagged and properly disposed of. After the base of the plant
is cut, the vines in the tree may remain a couple of years, but will eventually die. To exhaust
nutrient supplies in the roots, new sprouts should be cut as they appear. Any method
attempted should minimize disturbance to the soil, and native plants should be reintro-
duced to prevent re-infestation of English ivy. Selectively burning plants with a blow torch
may eventually use up the plant’s reserves. The potential effects of grazing are unknown.
Covering ground infestations with several inches of wood chips, grass clippings, or hay
may be effective. Mulch coverings should be applied for at least two growing seasons,
replenishing as needed.
Because English ivy is tolerant of preemergent herbicides, and the waxy cuticle on the
leaves makes it resistant to postemergent herbicides as well, chemical control is not always
effective. Glyphosate may be useful on young plants, and triclopyr may be effective when
most of the top growth of leaves is removed. Cutting the stems prior to herbicide application
facilitates chemical control. Glyphosate or triclopyr can be applied to cut stumps or cut
vines. Systemic foliar sprays are best applied on the evergreen ivy when the deciduous plants

Old Plants

E nglish ivy is noted for having a long life. Some vines are 1 ft. (0.3 m) in diameter,
and the oldest reported plant is 433 years old.
606 n VINES

are dormant, any time temperatures are above 55–60ºF (13–16°C), but before spring herbs
and wildflowers emerge. Winter treatments are less effective because the plant is growing
more slowly.
No biological controls are known, and attempts are unlikely because English ivy is an
important landscape plant in many parts of the United States.

Selected References
“English Ivy.” Southeast Exotic Pest Plant Council (SEEPPC) Invasive Plant Manual, 2003. http://
www.se-eppc.org/manual/HEHE.html.
“English Ivy Hedera helix.” Weed of the Week. U.S. Department of Agriculture, Forest Service, Invasive
Plants, 2006. http://www.na.fs.fed.us/fhp/invasive_plants.
Swearingen, Jil M., and Sandra Diedrich. “Hedera helix.” Weeds Gone Wild: Plant Conservation
Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/pdf/hehe1.pdf.

n Field Bindweed
Also known as: Creeping Jenny, European bindweed, small-flowered morning glory,
creeping Charlie, field morning glory, devil’s guts vine, corn bind, and others
Scientific name: Convolvulus arvensis
Synonyms: Convolvulus ambigens, C. incanus, Strophocaulos arvensis
Family: Morning Glory (Convolvulaceae)
Native Range. With the exception of the far northern latitudes, Europe, Asia, and
northern Africa.
Distribution in the United States. All states except Alaska.
Description. Field bindweed is an herbaceous perennial vine with slender stems that
sprawl on the ground or twine around objects. The branching stems, 1–6.5 ft. (0.3–2 m)
long, are normally glabrous but may be pubescent. The stem surface appears ridged or cor-
rugated. Prostrate plants can form tangled mats on the ground 2 in. (5 cm) thick. Leaves,
0.4–4 in. (1–10 cm) long and 0.1–2.4 in. (0.25–6 cm) wide, are alternate. They are usually
arrowhead in shape, with the lobes at the base pointing outward, often at right angles from
the leaf. Leaves are variable, however, and sometimes are almost oval. The leaves, which are
sometimes sparsely hairy, are supported by glabrous or pubescent petioles approximately
1.2 in. (3 cm) long. Vines are deciduous, dying back to the root crown in winter.
The fleshy, pale root system includes a perennial taproot, which may extend 2–10 ft.
(0.6–3 m) deep. Many creeping lateral roots, 2–10 ft. (0.6–3 m) long, grow from buds on
the taproot. Occurring in the top 2 ft. (0.6 m) of soil, rhizomes associated with the lateral
roots can create independent plants. One plant may have a root system 20 ft. (6 m) in diam-
eter and 30 ft. (9 m) deep.
From May to September, small clusters of 1–3 flowers grow on a 2.5 in. (6 cm) long
peduncle extending from the leaf axis. The peduncle has a pair of small linear bracts about
midway down. The funnel- or trumpet-shaped flowers, which open only in the morning,
are white, pink, tinged with pink, or rarely, red. The flowers are small, 0.5–1 in. (1.5–
2.5 cm) both long and wide. Both the five petals and the five green sepals are fused. The
sepals are glabrous, but the petals may be slightly pubescent. Flowering is indeterminate,
meaning that flowers will continue to develop as the stems grow until killed by frost.
 FIELD BINDWEED n 607

Fruit, which matures from

June to September, is a glabrous
cone-shaped capsule, 0.25 in.
(0.6 cm) long, with four parts
and four dark-brown or black
egg-shaped seeds.
Related or Similar Species.
Western morning glory, a per-
ennial native to the western
states, is also called chaparral
false bindweed or bush morn-
ing glory. Although growing
primarily in chaparral or pine
forests on dry slopes in Cali-
fornia, it can also be a weedy
pest in agricultural fields. Its
flowers are 1–1.5 in. (2.5–
4 cm) long. The basal lobes of
the leaves are squared or
slightly indented with two
lobes.
Hedge false bindweed, whi-
ch is also known by several
other names, is another invasive
vine native to Europe. It can be
distinguished by larger vegeta-
tive parts. Leaves are 2–4.5 in.
(5–12 cm) long. Flowers, 1.5–
3 in. (4–7 cm) long, are solitary,
not in clusters, and the pedicels
are 2–6 in. (5–15 cm) long, and
leaves are 2–4.5 in. (5–12 cm) Present in every state except Alaska, field bindweed has been considered a
long. weed in the United States since 1875. (Native range approximated from
Mallow bindweed or holly- USDA GRIN and selected references. Introduced range adapted from
hock bindweed, from the Med- USDA PLANTS Database, Invasive Plant Atlas of the United States, and
iterranean region of Europe, selected references.)
has become naturalized in the
foothills of some California mountains and in the southwestern states. It is a showy peren-
nial with purple to deep-pink flowers. Some of the upper leaves are distinguished by having
deeply divided lobes.
Field bindweed is difficult to distinguish from Ipomoea species, which are also referred to
as morning glories. The styles are different, and the sepals of Ipomoea may not be fused.
Dioscorea species, in the yam family, are also twining vines with heart-shaped leaves.
Because they are monocots, the leaves have parallel veins with no secondary vein network
and the flowers have three parts. Chinese yam, also called air potato, is an invasive vine. It
can be distinguished by the leaf venation, inconspicuous flowers, and the bulbils, or tubers,
which hang from the vine.
608 n VINES

A. Stems of field bindweed sprawl on the ground. (Richard Old, XID Services, Inc., Bugwood.org.) B. Leaves are
supported by long petioles; flowers are on long peduncles. (Steve Dewey, Utah State University, Bugwood.org.)
C. Flowers are funnel-shaped, with fused petals. (Joseph M. DiTomaso, University of California-Davis,
Bugwood.org.) D. Leaf form is variable, but usually arrow-shaped. (Ohio State Weed Lab Archive, The Ohio
State University, Bugwood.org.) E. Plants often clog machinery. (John D. Byrd, Mississippi State University,
Bugwood.org.)

Introduction History. Field bindweed was introduced to the United States in the 1730s
for use in medicine and as an ornamental. It was considered a weed in European gardens
as early as 1633 and was recognized as a weed in North America in 1875.
Habitat. Field bindweed grows in a wide range of climates, including temperate,
Mediterranean, and tropical regions. It is most abundant in slightly disturbed sites, such as
rangeland, pastures, abandoned fields, roadsides and railroad right-of-ways, and waste pla-
ces, but it is also found in cultivated fields, orchards, vineyards, and gardens. It is common
along irrigation drip lines in vineyards and along fencerows. Field bindweed grows best on
moist, fertile soils, but not in wet sites or standing water. It may also be found on poor,
dry, gravelly soils. It grows best in open, sunny locations but can invade forests if the canopy
is not too dense.
Reproduction and Dispersal. Field bindweed reproduces both by seed and vegetatively.
Seed production is variable. More seeds are produced under dry, sunny conditions from
plants growing on calcareous soils. Rain and heavy wet soils reduce seed output. Plants usu-
ally do not flower and set seed until their second season. One plant can produce as many as
500 seeds. Field bindweed is not self-compatible and must be pollinated by honey bees,
bumblebees, butterflies, and moths. Seed pods are dehiscent. Most seeds fall near the parent
plant, but some are dispersed over a long distance by water, birds and other animals, and
agricultural activity. Seeds or root pieces may be contaminants in soil, mulch, crop seed,
or hay, and may be also transported by farming equipment. The hard-coated seeds can sur-
vive as long as six days in the digestive tract of birds and animals. Although some seeds are
able to germinate just 10–15 days after the flower is pollinated, 80 percent develop an
impermeable coat that requires scarification. Seeds remain viable for 20–50 years, and the
seed bank is extensive. If moisture is available, germination may take place throughout the
growing season, but the peak occurs in mid-spring to early summer. A cold period of six
weeks in winter, with temperatures dropping to approximately 41ºF (5ºC), aids germina-
tion. Light is not required.
Most new shoots from rhizomes emerge in early spring. Root pieces as small as 2 in.
(5 cm) can produce new growth. Patches of field bindweed can expand their radius as much
 FIELD BINDWEED n 609

as 33 ft. (10 m) in one season. Although rhizomes can survive low winter temperatures, they
will die if the ground freezes.
Impacts. Growing in large patches and difficult to control, field bindweed is one of the
worst weeds in agricultural fields. It is especially a problem in cereal, bean, and potato crops.
It can reduce the harvest of cereal grains by 30–40 percent or more and outcompete vegeta-
ble seedlings. The plant competes with crops for nutrients and soil moisture and can
increase irrigation costs. Although the taproot accesses deep soil moisture, enabling the
plant to withstand drought, plants may also become dormant in summer, resprouting with
fall rains. Its tangled vines can interfere with crop harvesting.
Field bindweed can be an alternate host for viruses that affect crops, such as potato x dis-
ease, tomato spotted wilt, and vaccinium false bottom. Alkaloids in the foliage are mildly
toxic to grazing animals and may cause intestinal problems in horses.
In natural areas, the weight of the vines can break or uproot native species. By outcom-
peting native plants for sun, moisture, and nutrients, bindweed crowds out grasses and
forbs, decreasing biodiversity in the habitat.
Management. Control and prevention vary according to site, whether it is natural or agri-
cultural. Maintenance of a good vegetative cover can prevent infestations. Once established,
field bindweed is extremely difficult to eradicate. Physical control, including tilling, plowing,
or disking, must be carefully planned to avoid eliminating competitors, which would encour-
age more root sprouts from field bindweed. To prevent spreading seeds or rhizome pieces in
fields of actively growing crops, avoid tilling the field around the bindweed infestation. A year
of deep cultivation, followed by planting a competitive crop such as winter wheat or alfalfa,
may control bindweed in agricultural fields. The field should be cultivated before plants flower
and repeated each time new shoots appear. Tilling every two weeks should deplete root
reserves, but may also increase the potential for soil erosion. If done within three weeks of
emergence, cultivation to a depth of at least 4 in. (10 cm) will destroy new seedlings.
Small plants or new infestations, including all roots, can be pulled out by hand. Because
field bindweed does not tolerate water-logged soils, flooding the area, 6–10 in. (15–25 cm)
deep, for 60–90 days will kill plants. Although sheep and cattle avoid using it as forage, it
can be fed to livestock as fodder. Hogs and chickens will eat the stems, leaves, and exposed
roots or rhizomes. Mowing will reduce the seed crop. Shading plants by covering with
paper, straw, wood chips, or black plastic may kill them. In orchards or vineyards, a cover
crop of ryegrass, hairy vetch, and red clover will shade plants and can reduce the density
and biomass of bindweed.
Chemical applications repeated over several years may not eliminate field bindweed but
may suppress it enough to minimize the problem. Different biotypes have varying resistance
to herbicides. Glyphosate or 2,4-D, which should be applied before seed sets, is less likely to
damage crops. Although herbicides are difficult to use in infested fields of broadleaf crops, a
broadleaf selective herbicide can be used in grain crops. Spot treatment may prevent plants
from spreading. Herbicides are less effective when plants are drought-stressed, and fall
applications may be more effective than those applied in spring.
Two biological control agents have been released. A bindweed gall mite (Aceria malher-
bae), native to Europe and North Africa, causes galls to form on the leaves, distorting growth
and reducing seed production. Stems with galls can be picked and taken to other infested
sites to disperse the mites. The gall mite also attacks hedge bindweed and some native
Calystegia species of morning glory. A bindweed defoliating moth (Tyta luctuosa), native to
Europe, North Africa, and Eurasia, feeds on flowers and leaves. The insect was not yet estab-
lished in 2005, and the extent of damage is not known.
610 n VINES

Uses for Bindweed

B indweed has been considered a medicinal plant in Europe and Arabic-speaking

countries for centuries. It is used to stop bleeding and heal wounds, to reduce
fever, and to induce vomiting. Native Americans adopted the plant as early as
1890, using it as a laxative. Individuals of the Navajo Nation may use it on spider
bites and as an intestinal stimulant. The Pomo culture uses it to facilitate menstru-
ation and childbirth.

Selected References
Burnham, R. J. “Plant Diversity Website, Convolvulus arvensis.” CLIMBERS Website, 2010. http://www
-personal.umich.edu/~rburnham/SpeciesAccountspdfs/ConvarveCONVFINAL.pdf.
“Convolvulus arvensis, Field Bindweed.” Texas Invasives, 2008. http://www.texasinvasives.org/plant
_database/detail.php?symbol=COAR4.
“Dioscorea oppositifolia, Chinese Yam.” Texas Invasives, 2008. http://www.texasinvasives.org/plant
_database/detail.php?symbol=DIOP.
Jacobs, Jim. Ecology and management of field bindweed (Convolvulus arvensis L.). Invasive Species
Technical Note No. MT-9. U.S. Department of Agriculture, Natural Resources Conservation
Service, 2007. http://www.msuextension.org/ruralliving/Dream/PDF/Weed/bindweed.pdf.

n Japanese Dodder
Also known as: Giant Asian dodder, Tu Si Zi, devil’s hair, devil’s gut, strangleweed
Scientific name: Cuscuta japonica
Synonyms: Monogynella japonica
Family: Morning Glory (Convulvulaceae) or Dodder (Cuscutaceae)
Native Range. Temperate eastern Asia, including Mongolia, China, Manchuria, Korea,
Japan, Ryukyu Islands, Taiwan, and tropical Vietnam.
Distribution in the United States. Southeastern states, Texas, Florida, and South
Carolina. Recently found in 2004 in northern California. The northernmost plant, in
Redding, California, died naturally over the winter.
Description. Japanese dodder is an annual holoparasitic vine, meaning that it can only
obtain food and water from a host plant and will die without one. Usually an annual, it
may overwinter on host plants from tissue embedded in the host. In warm climates, the
plant may grow year-round. The much-branched stems are round, somewhat fleshy, and
smooth. Although the vine appears leafless, the stem is lined with tiny scale-like leaves.
The stems lack chlorophyll and range in color from pale green to vibrant yellow-green or
gold, sometimes with red spots or striations. Stems are thick, similar in size to and resem-
bling intertwining strands of cooked spaghetti. The plant coils around and attaches to its
host with peg-like modified roots called haustoria, which dissolve and penetrate the bark.
The haustoria extract everything needed for life from the host plant. The tangled mass of
stems can blanket the host plant and drape down to the ground.
The plant produces many inconspicuous pale-yellow flowers, supported by short stalks,
in leaf axils. The fruit is a two-cell capsule containing four seeds. Seeds are tiny, less than
0.1 in. (3 mm) long, pale straw to black in color, and irregularly dented.
 JAPANESE DODDER n 611

Related or Similar Species.

The term dodder is used to refer
to more than 150 parasitic, yel-
low twining plants that appear
to be nothing but leafless stems.
Stems of the many native dod-
der plants, which are present
in all states except Alaska, are
thin, resembling string or
thread. Native dodders usually
parasitize herbaceous plants
and small shrubs, enveloping
them in a large, spreading web,
and can be pests in orchards,
field crops, and gardens.
Common dodder, also called
scaldweed or swamp dodder,
usually grows in low, damp
locations. The stem is yellow to
dull orange, and the tiny bell-
shaped flowers are white.
Alfalfa dodder, a bright yellow
species native to Africa and
Eurasia, can be a problem in
alfalfa fields and vegetable
crops.
Introduction History. Japan-
ese dodder was accidentally
introduced three times, on
kudzu plants, to San Antonio,
Texas, Quincy, Florida, and
the Clemson University cam- The horticultural and agricultural industries of the United States may be
pus, South Carolina. It was seriously harmed if Japanese dodder spreads from its presently localized
believed to have been eradi- sites. (Native range adapted from USDA GRIN and selected references.
cated in all three locations. In Introduced range adapted from USDA PLANTS Database, Invasive Plant
2001, the parasitic vine was Atlas of the United States, and selected references.)
newly discovered in several
locations in Houston, Texas, and in 2004 in Redding, California. By 2010, Japanese dodder
was recorded at more than 245 sites in California, and more are expected. Dodder seeds are
a common contaminant of commercial seed, spice, baggage, and straw. Any shipment found
to contain whole dodder seeds of any species is denied entry into the United States. The
shipment is either returned to the country of origin or the seed is sterilized. Heat-treated,
sterile seeds may be legally imported, but seeds declared to be sterile are often not and will
sprout. In spite of such precautions, seeds are also intentionally imported and distributed
as a medicinal herb, probably by Asian immigrants or visitors. Because crushed or ground
seeds are not viable, they are not a threat.
Habitat. Japanese dodder will attach itself to both herbaceous and woody host plants.
In Japan, the plant is known to parasitize 7 types of crops and 32 different species of wild
612 n VINES

A. Vines completely cover this privet bush and drape to the ground. (Kim Camilli, Texas Forest Service,
Bugwood.org.) B. Vines look like a tangled mass of cooked spaghetti. (© Barry Rice, sarracenia.com.) C.
Haustoria from the white stem are beginning to penetrate the bark of this lemon tree. (W. Thomas Lanini,
University of California-Davis.) D. Stems climb by twining around the host plant. (© Barry Rice,
sarracenia.com.) E. Infected plants must be completely removed, as in this street scene in Houston, Texas.
(Kim Camilli, Texas Forest Service, Bugwood.org.)

plants in several families and genera. Japanese dodder has been seen on 20 species of
plants in south Texas, both herbaceous and woody, including ornamental crape myrtles,
native live oaks, and introduced privets. Crops in the United States that may be affected
include orchard trees, eggplant, potatoes, onions, pumpkins, tobacco, and soybeans, as
well as others. It is also commonly found on two introduced plants, kudzu (see Vines,
Kudzu), and purple osier willow. It can invade natural areas as well as roadsides and
unimproved property. In California, Japanese dodder may be found as ornamentals in res-
idential areas, particularly in Southeast Asian neighborhoods, where it is used for medici-
nal purposes. It may also grow in riparian areas, where it affects important native plants
such as willows, California live oak, and California buckeye. Loss of nesting sites may
affect Least Bell’s Vireo and other endangered birds. Japanese dodder does not grow in
hot, dry deserts, or at high elevations. Because the plant grows in several temperate cli-
mates in Asia, the potential is high for it to extend its range to similar climates in North
America.
Reproduction and Dispersal. Dodder seeds are carried over long distances as a contami-
nant in crop seed, and more locally by moving water, machinery, and soil disturbances such
as erosion. Although most germinate within a year, seeds remain viable for 10–20 years.
Plants in South Carolina produce viable seeds, but none have been seen in California or
Texas. Seeds may germinate either in soil or on the bark of the host plant, and seedlings
are both rootless and leafless. Although they may survive several weeks, using the soil as
an anchor, seedlings will die unless they find a host. As the seedling stem grows longer, its
tip twines toward other plants and seems to “sense” a potential host. Stems can grow several
inches a day.
Stem pieces are distributed locally by water, birds, animals, and human garden activities,
such as pruning, composting, and improper disposal.
Impacts. Because it is a parasite that takes nutrients and water from the host plant,
Japanese dodder negatively affects growth and the yield of infested plants. It can kill a large
plant in 2–3 years. It is capable of completely destroying a crop, and is a serious threat to
agricultural and horticultural industries. The loss of native plants in riparian areas alters
food chains, eliminates shade and nesting habitats, and may contribute to erosion.
 JAPANESE DODDER n 613

Although tall trees may be resistant to the effects of Japanese dodder, the understory is more
susceptible.
Potential impacts of more widespread infestations of Japanese dodder are serious.
The export market for ornamentals and crop seeds may be threatened. Other countries
may deny entry of seed exported from the United States, especially crop seed, if it may
be contaminated with dodder. Ornamental plants grown for export would have to be
more closely inspected. Maintenance costs of both private landscapes and public parks
would increase as infested plants are removed and replaced. The agriculture industry
would also bear the expense and loss in removing infected fruit and nut trees in
orchards.
Japanese dodder is a host for several serious citrus viruses that may be transmitted to
orchard crops.
Management. Physical control can be accomplished in small infestations by cutting the
infected host plant to the roots, then bagging all host and dodder fragments and disposing
of them in a deep commercial landfill. If possible, remove plants within 10 ft. (3 m) of the
infected plant. Because of the risk of re-infestation, neither host nor dodder should be com-
posted. Widespread infestations in fields can be eliminated with frequent tilling, burning, or
herbicide applications. Burning will destroy not only the dodder and its seeds, but also the
host plant. Host plants must be removed because haustoria embedded in the plant will
sprout.
Chemical applications are effective. Preemergent herbicides will prevent germination of
seeds and kill seedlings that have no host. Postemergents will kill both the dodder and host,
allowing all plant debris to be removed without threat of fragments resprouting. Sites can be
treated with both a soil fumigant and a preemergent. Systemics may be injected into large
trees.
No biological controls are known.

Selected References
Camilli, Kim. “Giant Asian Dodder: A New Invasive Plant Detected in Texas.” Invasive Plants of the
Eastern United States. U.S. Department of Agriculture, Forest Health Technology Enterprise
Team. Morgantown, WV, 2003. http://dnr.state.il.us/stewardship/cd/other/texas.html.
“Japanese Dodder (Cuscuta japonica).” Plant Division, Noxious Weed Control. Oregon Department of
Agriculture, n.d. http://www.oregon.gov/ODA/PLANT/WEEDS/profile_japanese dodder.shtml.

Medicinal Uses for Japanese Dodder

S eeds and other plant parts of Cuscuta have been used for centuries by Chinese
herbalists for various ailments, including jaundice, kidney and digestive prob-
lems, gallbladder disorders, and sore eyes. The plant is used as a male aphrodisiac
and is a major ingredient in currently sold herbal products for male enhancement.
Cuscuta is still used by contemporary Chinese herbalists for treating such wide-
ranging conditions as sperm leakage, dry eyes or blurred vision, lower back pain,
frequent urination, and ringing in the ears. Products are widely available in health
food stores and on the Internet.
614 n VINES

Markmann, C., and R. Marushia. “Summary of Dodder (Cuscuta japonica) Biology, Concerns, and
Management.” Plant Pest and Health Prevention Services (PHHPS). California Department of
Food and Agriculture, 2006. http://www.cdfa.ca.gov/phpps/ipc/noxweedinfo/pdfs/jdodder
_summary.pdf.
Miller, Arthur. “Japanese Dodder.” Invasive Plants of the Eastern United States. U.S. Department of
Agriculture, Forest Health Technology Enterprise Team, Morgantown, WV, 2003. http://dnr.state
.il.us/stewardship/cd/other/jdodder.html.

n Japanese Honeysuckle
Also known as: woodbine, Chinese honeysuckle
Scientific name: Lonicera japonica
Synonyms: Nintooa japonica
Family: Honeysuckle (Caprifoliaceae)
Native Range. Eastern China, Korea, and Japan.
Distribution in the United States. Most of the United States except for the northern Great
Plains. Scattered in California and on the Hawaiian Islands.
Description. Japanese honeysuckle is a perennial woody vine or shrub with a trailing or
twining habit. By twisting or coiling its stems around limbs and trunks of small trees, which
are typically less than 6 in. (15 cm) in diameter, it can climb vertical structures. Encircling
the tree trunk in closely spaced spirals gives the vine support for itself after the host tree
has died. Vines, usually 6.5–10 ft. (2–3 m) long, can be 80 ft. (24 m). Pubescence is variable,
but reddish-brown to light-brown young stems are often finely hairy. Older stems are
glabrous and hollow, with brownish bark peeling in long strips. In the southern and Mid-
Atlantic states, Japanese honeysuckle remains evergreen. In the colder North, it is semiever-
green, dropping its leaves after a long cold spell. Leaves, 1.5–3 in. (4–8 cm), are opposite on
the stem and have short petioles. Leaf shape is variable, oblong to oval. Leaves on new spring
shoots may be pinnately lobed. Leaf petioles and the midribs on both sides of the leaves may
also be finely hairy.
Roots are vigorous and widespread in the soil. One root system was recorded at almost
10 ft. (3 m) across and 3.3 ft. (1 m) deep.
Bloom time varies with location; April through December in the South, but only June in
Michigan. The very fragrant flowers grow in the leaf axils, usually in pairs but occasionally
singly. They are tubular, with five fused white petals, tinged with pink or purple, but they
turn yellowish as they age. Corollas are distinctly bilabiate. Fruit, inconspicuous small
round berries, 0.25 in. (6 mm), that become glossy black when ripe, develop on 1.2 in.
(3 cm) stalks in the fall. Each berry contains 2–3 oval seeds.
Related or Similar Species. Two cultivars are also invasive. Hall’s honeysuckle is more
aggressive and has pure white flowers. Chinese honeysuckle is less common, with purple,
glabrous leaves and red flowers.
Several Eurasian bush honeysuckles (see Shrubs, Exotic Bush Honeysuckles), such as
Amur honeysuckle, Morrow’s honeysuckle, Tartarian honeysuckle, and European fly honey-
suckle, have also been introduced to North America for ornamental plantings, wildlife cover,
and erosion control. They are generally deciduous shrubs ranging 6–15 ft. (1.8–4.5 m) tall
and do not have climbing, coiling vines. Because they have variable tolerances, bush
honeysuckles are found in habitats similar to those occupied by Japanese honeysuckle,
especially on disturbed sites. Opposite leaves, 2–2.5 in. (5–6 cm) long, are oval or
 JAPANESE HONEYSUCKLE n 615

egg-shaped. Older stems of

Japanese honeysuckle are usually
hollow, compared with native
honeysuckle stems that are solid.
The native coral honeysuckle
is distinguished from Japanese
honeysuckle by several features.
Flowers are red with yellow
interiors, and berries are orange
to red. Young leaves are gla-
brous, not hairy, and the upper-
most leaves on the vine are not
separate, but fused, forming
one leaf through which the vine
grows.
Introduction History. Japanese
honeysuckle was introduced to
Long Island in 1806 as an orna-
mental because of its fragrant
flowers and rapid growth.
A second introduction occurred
in 1862, also in New York. By
1919, the plant had naturalized
from Massachusetts to the Gulf
of Mexico and was described as
an invasive problem. It contin-
ues to be sold in nurseries as
an attractive landscape plant
and is still planted along high-
ways for erosion control and
bank stabilization. Wildlife offi-
cials in some states have pro- Japanese honeysuckle, an attractive ornamental with fragrant flowers, is
moted its use as winter forage still planted along highways for erosion control and is found in most
for deer. states. (Native range adapted from USDA GRIN and selected references.
Habitat. Japanese honey- Introduced range adapted from USDA PLANTS Database, Invasive Plant
Atlas of the United States, and selected references.)
suckle grows best in full sun
with rich soil, but is tolerant of
many conditions. It invades a variety of habitats, primarily where the natural ecosystem
has been disturbed, including fields, forests, wetlands, floodplains, southern pine
stands, and abandoned homesites. Thickets are often dense at forest margins, along
roadsides, and in open woodlands, prairies, and old fields. Although it is rarely found
in deep shade, it is somewhat shade tolerant and can invade mature forests, both dry
and mesic, by first growing in forest openings or in partial shade. Japanese honeysuckle
can go unnoticed in healthy forests until wind, disease, ice, landslide, or human activity
causes an opening to develop, after which plants take advantage of better conditions and
grow vigorously.
Cold winters may limit its northward expansion, while drier conditions, which affect
seedlings, may limit its western spread. Although the northern limit is currently where high
616 n VINES

A. Vines can overtop and completely enclose other plants. (Chris Evans, River to River CWMA, Bugwood.org.)
B. Oblong leaves are opposite on the stem. (Karan A. Rawlins, University of Georgia, Bugwood.org.) C. Lower
leaves on new spring growth may be pinnately lobed. (Ted Bodner, Southern Weed Science Society,
Bugwood.org.) D. Tubular, bilabiate flowers grow in pairs from leaf axils. (Chuck Bargeron, University of
Georgia, Bugwood.org.) E. Fruit are small berries. (Karan A. Rawlins, University of Georgia, Bugwood.org.)
F. Bark on old stems is stringy and peeling. (James H. Miller, USDA Forest Service, Bugwood.org.)

temperatures in winter average −13ºF (−25ºC), Japanese honeysuckle continues to spread

north due either to increased tolerance or to warmer winters. Plants on the northern edge
are smaller due to the shorter growing season and rarely produce flowers or seeds. It grows
most vigorously and is the biggest pest where annual precipitation is 40–47 in. (1,000–
1200 mm) and winter low temperatures average 5 to −17.5ºF (−8 to −15ºC).
Reproduction and Dispersal. Although Japanese honeysuckle reproduces both vegeta-
tively and sexually, its runners and rhizomes allow plants to overrun sites. New shoots grow
from the roots, and vines will develop roots at nodes when in contact with moist soil. Stem
cuttings will also root. Because the bloom season lasts several months, many fruits and seeds
are produced. Fruit are eaten by deer, rabbits, turkey, and quail, but long-distance seed dis-
persal is primarily by birds, which deposit seeds in new locations. Little research has been
conducted on seeds, but the number produced is related to soil chemistry. A large seed bank
remains in the soil, ready to sprout when the site is disturbed. Dormancy and germination
requirements are variable.
Impacts. A pest in both managed forests and natural areas, Japanese honeysuckle spreads
and outcompetes and replaces native plants in several ways. By branching frequently and
rooting at nodes, the plant forms dense interlaced canopies, arbors, or thickets that can com-
pletely cover other vegetation. Forming a dense blanket over existing plants, it eliminates
their light and prevents them from photosynthesizing. Shrubs and trees that provide support
are either killed by girdling or break under the weight of the vines. The root system is vigo-
rous, depriving native plants of necessary nutrients and rendering them less healthy. It is a
rapid grower, and its evergreen to semi-evergreen nature gives it an advantage over decidu-
ous species.
Plants change forest structure by altering or stopping natural forest succession by
eliminating the understory, either by shading or girdling. As an evergreen, Japanese honey-
suckle casts shade in early spring, preventing the germination of ephemeral spring herbs. As
 JAPANESE HONEYSUCKLE n 617

understory herbs, shrubs, and young trees are covered and killed, gaps are created within
the forest. With more light, Japanese honeysuckle grows even more vigorously, and the
openings provide places for other invaders, such as kudzu and English ivy, to take hold.
Changes in the understory of forests alters wildlife habitat and eliminates nesting sites for
song birds. In contrast, dense thickets provide cover for other birds and small mammals,
while leaves and seeds provide food for cottontail rabbits and birds. White-tailed deer eat
the evergreen leaves, providing important winter forage.
Japanese honeysuckle is a temporary host for agricultural pests, such as a spider mite
(Tetranychus urticae) that attacks corn and peanuts, a cicadallid cotton pest (Empoasca bigut-
tula), and the tobacco budworm.
Management. Because of the difficulty of controlling Japanese honeysuckle, the goal
should be 100 percent mortality. Infestations can be prevented if plants are removed as soon
as they are noticed, preventing their spread. Plants, however, are hard to find when small.
Regardless of what method is used to control Japanese honeysuckle invasions, sites should
be monitored frequently for new growth. Methods should also minimize soil disturbance,
which creates habitat for germination of not only Japanese honeysuckle, but other invasive
species as well.
Physical methods are generally ineffective in the long term because vines form roots at
nodes and existing plants resprout from root buds. Cutting, pulling, or burning will weaken
the plant but not kill it. Small stands or seedlings can be hand-pulled, but roots must also be
removed. Although burning fails to kill the roots of mature vines, which will resprout, fires
are more likely to kill seedlings and young plants. Mowing at least twice a year will keep
vines and new shoots from spreading, but will not kill plants. Mowing, however, may also
stimulate growth. Twining vines should be cut off of trees and shrubs, and all pieces must
be properly disposed of because cut vines and branches will grow roots. Disking may be
effective, but is very destructive and may also stimulate germination of the seed bank.
Although goats will browse on Japanese honeysuckle, they would be invasive themselves if
not confined.
Chemical methods are the most effective means of control. Application is best done in the
fall, when Japanese honeysuckle is still growing but native plants are going dormant. Large
areas can be mowed to remove biomass, then treated with herbicides. Effective herbicides
include glyphosate, 2,4-D + picloram, tebuthiuron, dicamba, and sulfometuron. A combina-
tion of burning in fall and winter followed by spraying new sprouts with herbicide is
effective.
Japanese honeysuckle has few natural enemies in North America. Although biological
efforts are necessary for regional control, little research has been done. It is susceptible to
honeysuckle latent virus and to tobacco leaf curl (Begomovirus).

How Fast Does Japanese Honeysuckle Grow?

A growth rate of about 5 ft. (1.5 m) per year is typical, but in one location, it
took vines only one year to cover a 14.75 ft. (4.5 m) tree. Each vine has many
runners or canes, which also branch. The combined length from one sprout for
one year’s growth can be more than 49 ft. (15 m).
Source: Nuzzo, 1997.
618 n VINES

Selected References
Bravo, Melissa A. “Japanese Honeysuckle.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas.
Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/
fact/pdf/loja1.pdf.
Nuzzo, Victoria. “Element Stewardship Abstract, Lonicera japonica.” Global Invasive Species Team,
Nature Conservancy, 1997; updated 2009. http://wiki.bugwood.org/Lonicera_japonica.
Swearingen, J., K. Reshetiloff, B. Slattery, and S. Zwicker. “Exotic Bush Honeysuckles.” Plant Invaders
of Mid-Atlantic Natural Areas. National Park Service and U.S. Fish and Wildlife Service, 2002;
updated 2003. http://www.invasive.org/eastern/midatlantic/loni.html.

n Japanese Hops
Also known as: Japanese Hop
Scientific name: Humulus japonicus
Synonyms: Humulus scandens
Family: Hemp (Cannabidaceae)
Native Range. Temperate Asia, including southern and eastern China, Manchuria, the
Amur region of Siberia, Korea, Japan, and the Ryukyu Islands. Also in tropical Vietnam.
Distribution in the United States. Scattered in the eastern United States, from North
Dakota east to Maine, south to Georgia, and west to Kansas. Currently most common in
the northeastern United States, Japanese hops continues to expand its range.
Description. Japanese hops is usually an annual vine, but it can be weakly perennial in
warmer climates. The plant has no tendrils and climbs structures and other plants by twin-
ing its stems around the host. Without something on which to climb, plants can be trailing
or prostrate. Stems, occasionally branching, may be 8–35 ft. (2.5–10.5 m) long by late
summer. Stems are fairly stout, light green to reddish purple, and longitudinally ridged.
Short, sharp, downward pointing prickles, which help the plant grasp host plants, line the
stem ridges. The opposite leaves, both 2–6 in. (5–15 cm) long and wide, are palmate, usu-
ally with five deep lobes that taper to a point. Leaves on the top parts of the vine often have
3–5 lobes, while others may have 7–9 lobes. Leaf margins are coarsely toothed, with the ser-
rations pointing forward. The upper leaf surface has a sparse cover of short, rough hairs,
while stiff prickly hairs line the major vines on the underside. The stout, light-green leaf
petioles, also covered with prickly hairs, are as long as or longer than the leaves, with a pair
of small bracts, curved slightly downward, where the petiole joins the stem.
Plants are dioecious, with male and female flowers on different plants, and bloom in July
and August. Both types of inflorescences grow in leaf axils. Male inflorescences are erect
spreading panicles, 6–10 in. (15–25 cm) long and as wide as 5 in. (12.5 cm). The panicle
branches are slightly hairy and light green to pale red, with a pair of small bracts at the base.
The yellow-green male flowers are very small, approximately 0.1 in. (3 mm), with five
spreading light-green to pale-red sepals and no petals. Male flowers droop downward from
the panicle branches. The female plant inflorescence is a short, catkin-like drooping spike of
flowers and bracts, 0.25–1.5 in. (0.5–4 cm) in diameter, which becomes globular with age.
A pair of small, inconspicuous female flowers, which have no petals, occurs at the base of
each of the bracts. The cluster of pale-green, overlapping bracts, densely hairy at the mar-
gins, somewhat resembles a small pine cone. Initially, the bracts are narrow triangles, but
they become broader as they age, with pointed, slightly recurved tips. The female flower
inflorescence is called the hops. Seeds, one from each female flower, mature in September.
 JAPANESE HOPS n 619

The yellowish-brown achenes,

with dark specks, are small,
approximately 0.1–0.2 in.
(3–5 mm), slightly oval and flat-
tened, with a blunt tip.
Related or Similar Species.
The only other species in the
genus is common hops, which
is used in beer making. Com-
mon hops has five varieties, three
of which are native to the United
States, one native to Europe and
West Asia, and one native to
East Asia. Common hops leaves
usually have three lobes or none,
with rounded tips. The petiole is
shorter than the length of the
leaf. The male inflorescence is
pale yellow, lacking the reddish
tint of the sepals and branches
of Japanese hops. The female
flower spikes are oval and hair-
less, and the bracts, or scales,
are blunt, either straight or curv-
ing slightly inward. Japanese
hops does not contain the ele-
ments in common hops that are
essential for brewing beer.
Two native cucumber spe-
cies may be confused with
Japanese hops. The palmately
veined leaves of the native bur Growing best in sunny habitats, Japanese hops frequently invades
cucumber are alternate, with disturbed areas, such as roadsides and construction sites, in the eastern
3–5 very shallow lobes that are half of the United States. (Native range adapted from USDA GRIN and
pointed at the tips. Although it selected references. Introduced range adapted from USDA PLANTS
Database, Invasive Plant Atlas of the United States, and selected references.)
is pubescent on the stem and
underside of leaves, the plant
has no prickles. It uses tendrils to climb. The monoecious plants have showy flowers,
0.5 in. (1.3 cm) in diameter, with white petals fused at the base. Fruits occur in clusters
and are covered with white prickles which are approximately 0.5 in. (1.0–1.5) cm long.
Wild cucumber has alternate, glabrous leaves and hairless stems. The thin leaves, with
five deep, pointed lobes, have smooth margins. Wild cucumber also climbs with tendrils.
The showy white flowers, 0.5–0.75 in. (1.3–2 cm) in diameter, each have six narrow,
loose white petals. The fruit, occurring individually, resembles a small cucumber, about
2 in. (5 cm) long. Native Virginia creeper has compound leaves with five leaflets and no
prickles.
Introduction History. Japanese hops was introduced into the United States in the late
1800s for ornamental purposes and for use as a tonic in Asian medicine. It is an attractive
620 n VINES

A. Climbing plants form thick masses of vines. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology, University
of Connecticut, Bugwood.org). B. Leaves are palmately lobed. (Chris Evans, River to River CWMA, Bugwood.org).
C. Female flowers are the “hops.” (Leslie J. Mehrhoff, Ecology and Evolutionary Biology, University of
Connecticut, Bugwood.org.) D. Downward pointing prickles help the plant grasp its host. (Chris Evans, River
to River CWMA, Bugwood.org.)

annual in gardens, especially the variegated forms, and is useful for garden screens. It is still
sold in nurseries.
Habitat. Although capable of growing in part shade, Japanese hops prefers bright sunlight.
It is tolerant of any type of soil, including, sandy, loamy, clay, acid, neutral, or basic, as long as
it is moist. Plants are most common in riparian and floodplain locations, ditches, and creek
banks, but also threaten open woodlands, fields, and prairies. Because seeds prefer bare soil
for germination, the plant is often found on disturbed sites, such as roadsides, fence lines,
old fields, hayfields, construction sites, weedy meadows, and other non-crop areas. The plant
may also spread rapidly into urban locations and managed areas such as parks.
Reproduction and Dispersal. Flowers are wind-pollinated. Seeds mature in the autumn,
when growth of the vine slows. After plants are killed by the first hard frost, vines disinte-
grate and decay. Seeds may be carried long distances by animals, people, machinery, and
waterways. Seeds begin to germinate in early spring, and continue to do so all summer as
long as conditions are sunny and moist. Seedlings can remain dormant in the cotyledon
stage for several weeks. When the weather turns hot, seedlings rapidly grow into long vines.
Seeds remain viable for three years.
Impacts. With several thousands of plants per acre, Japanese hops can blanket large areas
of open ground or low vegetation. Its rapid growth in summer enables it to grow long stems
that overtop other plants, such as understory shrubs and small trees in woodlands. It can
create dense mats that are several feet thick. Its weight can cause host plants to topple over
or break, and the twining habit can also girdle plants. It displaces native species by blocking
light to existing plants and preventing seedling growth, especially in riverbank and flood-
plain locations.
The prickles are irritating, and contact with plants may cause dermatitis or blistering in
some people. In China and Korea, pollen concentrations cause symptoms of asthma and
allergic rhinitis, a problem that could be increased if Japanese hops continues to increase
its range in the United States.
Management. Because both hops species are common garden or commercial plants, little
information is available regarding their control. Any eradication efforts should be continued
 JAPANESE HOPS n 621

for at least three years to ensure the elimination of all seeds in the seed bank. Because
Japanese hops requires bright light, maintenance of a closed tree canopy will prevent its
invasion. Vines use weeds and low branches as “ladders” to reach shrubs and small trees.
If appropriate to the site, weeds and low branches can be removed. Although Japanese hops
will grow and overtop grasses once started, they do not germinate well in sod-covered grass-
lands. Maintenance of grassland pastures might prevent infestations. Any machinery used
for eradication should be thoroughly cleaned.
Because it is slow and labor intensive, physical removal is appropriate only for small
stands. The annual plants can be pulled any time of year before seed develops. Although
the root system is not deep or extensive, allowing plants to be easily pulled out, the roots
must be removed because buds will sprout. Physical removal should be done early in spring,
when the roots are small and before vines become entangled in other vegetation. Because
seeds germinate all summer, removal should be done monthly. Repeatedly cutting or mow-
ing plants close to the ground surface until they die in the autumn is effective because it pre-
vents seed production. All root and stem pieces should be removed from the site. Burning is
inappropriate because the vines would carry the fire into the tree crowns.
Chemical applications of both preemergents and postemergents are effective. Application
of preemergents can be done around young trees in plantations or restoration projects, but
not in natural woodland. When applied in spring, sulfometuron methyl proved to be the
most long-lasting preemergent. Postemergent herbicides should be applied in April and
May, after most Japanese hops seeds have germinated, but before the vines grow enough
to cover vegetation and before new seeds mature in August. Metsulfuron methyl and glyph-
osate offer the best control, but plants may resprout from the roots if they are insufficiently
damaged.
Although some insects and pathogens attack or feed on Japanese hops, the introduction
of biological control agents is unlikely because of the importance of common hops as a crop.
Although Japanese hops is susceptible to hop latent carlavirus and Humulus japonicus ilarvi-
rus, both transmitted by aphids, these viruses also infect common hops. Two moths
(Epirrhoe sepergressa and Chytonix segregata) and a fungus (Pseudocercospora humuli), natural
enemies of Japanese hops in Asia, are being studied. The Japanese beetle (Popilla japonica)
feeds on hops but does little damage.

Selected References
Hilty, John. “Japanese Hops (Humulus japonicus).” Weedy Wildflowers of Illinois, 2010. http://
www.illinoiswildflowers.info/weeds/plants/jp_hops.htm.
Pannill, Philip D., Aaron Cook, Anne Hairston-Strang, and Jil M. Swearingen. “Japanese Hop.” Weeds
Gone Wild: Alien Plant Invaders of natural areas. Plant Conservation Alliance’s Alien Plant Working
Group, 2009. http://www.nps.gov/plants/alien/fact/pdf/huja1.pdf.
Renz, Mark. “Japanese Hops, a New Weed Species to Be Aware of in Wisconsin.” Wisconsin Crop
Manager Newsletter. Integrated Pest and Crop Management (IPCM), University of Wisconsin,
2008. http://ipcm.wisc.edu/WCMNews/tabid/53/EntryId/623/Japanese-hops-a-new-weed-species
-to-be-aware-of-in-Wisconsin.aspx.
Steffen, Brad, and Bob Edgin. “Japanese Hops (Humulus japonicus Sieb. & Zucc.) Species Character.”
Vegetation Management Guideline 1(40): 2007. Illinois Nature Preserves Commission (INPC),
Illinois Department of Natural Resources (DNR). http://dnr.state.il.us/inpc/pdf/VMG%20Japanese
%20hops%20original%202007.pdf.
622 n VINES

n Kudzu
Also known as: Japanese arrowroot
Scientific name: Pueraria montana
Synonyms: Dolichos lobatus, Dolichos hirsutus, Pueraria hirsuta, P. lobata, P. pseudohirsuta,
P. Thunbergiana, P. triloba
Family: Pea (Fabaceae)
Native Range. Eastern Asia, including China, Korea, Japan, Taiwan, and Southeast Asia.
Distribution in the United States. Widespread in the eastern states, as far north as
Illinois, Michigan, and Maine, west to Texas, Oklahoma, Kansas, and Nebraska. Plants have
also been reported in Portland,
Oregon, and in Washington
State. Also in Hawai’i. The
worst infestations are in the
southeastern states.
Description. Kudzu is a per-
ennial twining or climbing
woody vine. It produces exten-
sive mats that cover all other
vegetation, including tall trees.
New growth vines are pubescent
and as much as 0.5 in. (1.3 cm)
in diameter. Vines become
fibrous as they age, and can be
as much as 6 in. (15 cm) in
diameter and as long as 100 ft.
(30 m). Very old stems may be
12 in. (30 cm) in diameter.
Leaves, supported by petioles
2–4 in. (5–10 cm) long, are
alternate and compound. Each
of the three heart-shaped leaflets
is 3–7 in. (7.5–18 cm) long and
can be either entire or two- or
three-lobed. The herbaceous
stems, petioles, and underside
of leaves are covered with tan
or brown hairs. The plant is
deciduous and not frost tolerant,
dropping its leaves at the first
frost in autumn. Although the
leaves and new growth die, the
dried leaves remain on the vine
Because kudzu was formerly promoted by the federal government for
erosion control, the plant is widespread in southern states. (Native as a dead mat. New growth
range adapted from USDA GRIN and selected references. Introduced begins in May.
range adapted from USDA PLANTS Database, Invasive Plant Atlas of the Kudzu produces large, fleshy
United States, and selected references.) taproots or tubers, up to 7 in.
 KUDZU n 623

(18 cm) in diameter and 6 ft. (2 m) or more long. Roots are typically 3.3–10 ft. (1–3 m)
deep, but some can extend down as far as 16 ft. (5 m). Tubers, which can weigh up to
400 lb. (180 kg), store an abundance of carbohydrates to sustain the plants.
Plants in full sun bloom from July to October. The fragrant flowers, which emit a
grape-like aroma, are bright red-pink or purple pea-like florets, 0.5–1 in. (1.3–2.5 cm) long,
clustered on 4–12 in. (10–30 cm) long racemes. They are common on upright or draped
climbing vines but rarely occur on plants covering flat ground or other horizontal surfaces.
Clusters of 20–30 bean-shaped hairy pods ripen in the fall but remain on the plant until
January. Pods are 3 in. (8 cm) long and contain 3–10 hard flattened, kidney-shaped seeds
up to 0.3 in. (8 mm) wide.
Related or Similar Species. Four native legumes that also have three-lobed leaves may
appear similar. Prostrate ticktrefoil, also called dollar leaf plant, is a forb, with leaflets that
are glabrous and almost round. Leaves of American hogpeanut, which can be either a forb
or a vine, are less pubescent. Its flowers are white or pale lilac, and the fruit is only 0.2 in.
(5 mm) long. Leaves of amberique bean, also called trailing wild bean, are sparsely hairy on
the lower surface and smaller. The plant can also be either a vine or a forb. The flowers form
in small clusters, and the seed pods resemble green beans. Poison ivy, which can be a shrub,
forb, or vine, climbs with adventitious roots and its leaflets are more distinctly lobed.
Introduction History. Kudzu was introduced to the United States at the 1876 Centennial
Exposition in Philadelphia and promoted as an ornamental because of its fragrant flowers.
By 1900, rooted cuttings were available by mail order from an enthusiastic Florida farmer
who planted 35 acres for hay and considered kudzu an ideal plant for livestock forage.
From 1935 to the mid-1950s, it was promoted by the federal government for use in erosion
control, especially in the impoverished soils of the Appalachian piedmont in Alabama,
Georgia, and Mississippi. The Soil Conservation Service distributed 85 million seedlings.
Farmers were paid up to $8 per acre to plant kudzu, and more than 1.2 million acres were
planted. During the 1940s, kudzu clubs formed in honor of the vine. In the 1930s and
1940s, kudzu was also promoted as a high-nitrogen forage crop for livestock, but it proved
hard to bale and cattle trampled the vines. The federal government removed kudzu from its
list of acceptable cover plants in 1953, and in 1972, the U.S. Department of Agriculture
declared it a noxious weed.
Habitat. Kudzu grows under a wide range of environmental conditions and is present in
a variety of ecosystems, from dry flatwoods to riparian zones. It can grow where the water
table is high, but cannot tolerate flooded soils. Although it is found on many types of sub-
strate, it grows best on well drained, acid to neutral soil, with a pH of 4.5–7. As a nitrogen
fixer, however, it can thrive on poor soils. It prefers full sun but can exist on the floor of a
closed-canopy forest because the vines climb trees to get enough light. The plant can invade
agricultural land, forests, plantations, range, grassland, riparian zones, scrub, and urban
areas as well as disturbed sites, such as abandoned fields and roadsides. Infestations seen
around abandoned homesteads are probably from ornamental plantings gone wild.
Although it is unable to root on healthy grass cover, the vines can overrun grass areas such
as pasture. It grows best in temperate regions with mild winters, 40–60ºF (4.5–15.5ºC),
summers above 80°F (26.5°C), and 40 in. (1,000 mm) or more of rainfall. Kudzu can sur-
vive both drought and cold winters because of its massive root reserves.
Reproduction and Dispersal. Kudzu spreads almost exclusively by vegetative means, by
runners and rhizomes that root at nodes. Pieces of the plant can be carried and subsequently
deposited by vehicles, machinery, or flowing water. Although it grows best in wet climates,
it is drought tolerant, resprouting from its massive root crowns. One root can have up to 30
624 n VINES

A. Kudzu can rapidly overtop and smother even tall trees. (James H. Miller, USDA Forest Service, Bugwood.org.)
B. The three leaflets may have entire margins. (James H. Miller, USDA Forest Service, Bugwood.org.) C. In the
variety lobata, leaflets are lobed. (Ted Bodner, Southern Weed Science Society, Bugwood.org.) D. Pea-shaped
flowers are clustered on long racemes. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.) E. The large
tubers store carbohydrates for the vines. (Forest and Kim Starr, U.S. Geological Survey, Bugwood.org.) F. Vines
climb by twining around other plants. (Rebekah D. Wallace, Bugwood.org.) G. Seed pods are hairy. (Ted Bodner,
Southern Weed Science Society, Bugwood.org.)

vines growing from it. Wherever nodes touch the soil, the vine will root and produce new
root crowns, as many as one in every 1–2 sq. ft. (5–10/m2). The result can be thousands
of plants in a dense stand. Vines can grow more than 1 ft. (0.3 m) in a day, and 65–100 ft.
(20–30 m) in a year.
Flowers appear in the late summer or early fall of the plant’s third year of growth.
Although kudzu is capable of producing abundant seeds, seed counts vary from zero to
180 per sq. ft. (0–1,800/m2) of soil. Insect damage may be responsible for low seed counts.
Seeds are dispersed by mammals and birds. Although few seeds are viable, they remain so
for many years and are probably responsible for long-distance migration. Seeds do not easily
germinate and require scarification. Seedlings are poor competitors.
Impacts. Kudzu kills trees and other vegetation by smothering the plants and blocking all
light. Vines girdle tree trunks, and the weight of the plant breaks branches and uproots trees,
disrupting entire ecosystems and rendering the land useless. The weight of the vines can also
cause power lines and buildings to collapse. Kudzu is a problem on more than seven million
acres (2.8 million ha.) in the southeastern United States, and it is estimated by the
Congressional Office of Technology that $50 million is lost in land productivity every year.
Costs of controlling kudzu may exceed the productivity value of the land it covers.
Expenditures of power companies to clear vines off utility lines are estimated at $1.5 million
per year.
Recent studies suggest that kudzu increases ozone pollution. As a nitrogen fixer, kudzu
alters the nitrogen content of soils and doubles soil emissions of nitric oxide. The plant also
emits large amounts of isoprene, a volatile organic compound (VOC). Nitric oxides and
VOCs in the atmosphere chemically react with sunlight to create ozone, a major component
of smog. Atmospheric modeling predicts a 50 percent increase in the number of unhealthy
high-ozone days attributed to kudzu infestations.
 KUDZU n 625

Are There Uses for Kudzu?

K udzu has traditional culinary and herbal uses in Japan and in China, and it has
also been used to reduce hypertension. It currently is being studied as treat-
ment for alcoholism because it appears to reduce the craving. Kudzu can produce
forage for livestock, with a carrying capacity of one cow per acre. Goats will also
feed on the vines.

Management. Preventing the spread of kudzu depends on proper cleaning of all vehicles
and equipment that come into contact with the plant, because pieces will root and may be
carried in soil and on vehicles. Successful eradication of kudzu requires that the root system
be destroyed, a process which requires persistent treatment and may take 10 years.
Physical means of control that remove the vines may not be effective without follow-up
applications of herbicides on root crowns. Digging out plants is difficult because of the large
tubers, and is best used only for small, initial infestations. Repeated grazing by cattle, or
more effectively by goats, over several years may remove the foliage, causing depletion of
the energy stored in the roots. Mowing or cutting may do the same. Cut vines can be used
as livestock feed, burned, or placed in plastic bags in a landfill. Burning the upper growth
promotes seed germination, and new sprouts must be eliminated.
Chemical control can be achieved by applying systemic herbicides, either glyphosate or
triclopyr. Spraying of foliage can be used in large populations where saving native plants is
not possible, but the root crown must also be treated. After locating the tuber by following
the stems, workers should cut into the root crown and apply the herbicide to not only the
root crown but also to any runners in the soil. Where it is desirable to preserve native vegeta-
tion in a mixed forest, kudzu can be selectively cut to stumps 2 in. (5 cm) above ground
level, for immediate application of the herbicide. With either method, new sprouts must also
be sprayed soon after emergence.
Biological control is being investigated and surveys are being conducted for insects in
kudzu’s native range. Some potential candidates have been rejected because they would also
feed on crops. Possibilities include leaf-feeding beetles, sawflies, stem-boring weevils, and a
large beetle that lays eggs on the main vines and roots. A fungal pathogen (Synchytrium puer-
ariae) from southern China may have potential. Three pathogens native to North America, a
bacterium (Pseudomonas syringas pv. Phaseolicola), a fungus (Myrothecium verrucaria), and
Colletotrichum gloeosporioides, have also been studied. The fungus, from the sicklepod plant
(Senna obtusifolia), has effectively killed kudzu in both greenhouse and field studies.

Selected References
Bergmann, Carole, and Jil M. Swearingen. “Kudzu.” Weeds Gone Wild: Alien Plant Invaders of Natural
Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/
alien/fact/pumo1.htm.
Britton, Kerry O., David Orr, and Jianghua Sun. “Kudzu.” In Biological Control of Invasive Plants in the
Eastern United States, by R. Van Driesche et al. USDA Forest Service Publication FHTET-2002-04,
2002. http://www.invasive.org/eastern/biocontrol/25Kudzu.html.
“Kudzu.” Southeast Exotic Pest Plant Council Invasive Plant Manual, n.d. http://www.se-eppc.org/
manual/kudzu.html.
626 n VINES

“Kudzu (Pueraria lobata).” Washington State Noxious Weed Control Board, March 2007. http://
www.nwcb.wa.gov/weed_info/Pueraria_lobata.html.
Samarrai, Fariss. “Invasive Kudzu Is Major Factor in Surface Ozone Pollution, Study Shows.” Top
News from the University of Virginia, 2010. http://www.virginia.edu/uvatoday/newsRelease.php
?id=11862.

n Mile-A-Minute
Also known as: Asiatic tear-thumb, devil’s tail tear-thumb, mile-a-minute weed
Scientific name: Persicaria perfoliata
Synonyms: Polygonum perfoliatum; Polygonum arifolium var. perfoliatum
Family: Buckwheat (Polygonaceae)
Native Range. Southern and eastern Asia, from India east to China, and north to far
eastern Russia and Korea. South to Vietnam, Thailand, and the Malay Peninsula. Also
Japan, the Ryukyu Islands, Taiwan, and the Philippines.
Distribution in the United States. Mile-a-minute is currently localized in the eastern
states, from New York and Massachusetts south to North Carolina, centered on the intro-
duction site in southern Pennsylvania.
Description. Mile-a-minute is an annual, herbaceous, trailing vine with green stems that
become reddish with age. The plant’s height varies with the site. In open areas, it becomes
a dense mat, but vines will climb as high as 26 ft. (8 m) on trees at forest edges. Except where
they become woody at the base, the elongated and branched stems are slender and delicate.
Stems, leaf veins, and petioles are covered in 0.04–0.08 in. (1–2 mm) curved barb-like
spines. The pale-green leaves are thin, alternate on the stem, and form distinct equilateral
triangles, 1.5–2.8 in. (4–7 cm) long and 2–3.5 in. (5–9 cm) wide. Cup-shaped structures,
called ocreas, form sheaths, 0.4–0.8 in. (1–2 cm) in diameter, which encircle the stems at
the nodes. Vines die back with the first fall frost. Roots are shallow and fibrous.
Flower buds, and subsequently the fruit, emerge from inside the ocreas, either on the
ends of stems or in the axils of the upper leaves. Inflorescences are spike-like clusters of
10–15 tiny, inconspicuous white flowers, 0.1–0.15 in. (3–4 mm). After bloom, the calyx
thickens to produce a metallic blue segmented berry-like fruit 0.2 in. (5 mm) in diameter.
Each segment holds a single shiny black seed, 3 mm in diameter.
Related or Similar Species. Many species of Persicaria, Polygonum and other genera in the
buckwheat family occur in North America, both native and introduced, but none resembles
mile-a-minute.
Introduction History. Mile-a-minute was first recorded in the United States in 1890, from
an herbarium specimen in Portland, Oregon, but the plant did not become established. It
was accidentally brought to a plant nursery in southern Pennsylvania in the 1930s, when
imported holly seeds were planted. It was also introduced into a garden in Maryland, where

World’s Worst 100?

T his plant, Persicaria perfoliata, is not the same as mile-a-minute (Mikania

micrantha), which is in the Asteraceae and on the list of 100 of the World’s
Worst Invasive Alien Species.
 MILE-A-MINUTE n 627

it has since been eradicated.

Interested by the new plant,
the nursery owner in Pennsy-
lvania allowed it to grow. It has
since escaped cultivation and
expanded its range approxi-
mately 300 mi. (480 km) in
every direction, and continues
to spread.
Habitat. Because mile-a-
minute is an early successional
species in its native range, it
most often invades bare ground
and disturbed areas, such
as roadsides, edges of wood-
land, forest openings, orchards,
riparian sites, drainage ditches,
and fallow fields—any open,
sunny site. It is quick to invade
sites that have been treated with
herbicides to clear brush, such
as kudzu sites and utility corri-
dors. Although it can grow on
drier sites, it prefers moist soil
and even tolerates very wet sites
with poor soil structure. Plants
also grow in urban locations,
recreational areas, and plant
nurseries. Although it tolerates
some shade, it satisfies its light
requirement by climbing on
top of existing vegetation. Although mile-a-minute vine has thus far been contained relatively close
As a temperate or subtropi- to its point of introduction in Pennsylvania and Maryland, it has the
cal species, it can invade cli- potential to greatly increase its range. (Native range adapted from USDA
mates with moderately cold GRIN and selected references. Introduced range adapted from USDA
winters. Eight weeks of temper- PLANTS Database, Invasive Plant Atlas of the United States, and selected
references.)
atures 50ºF (10ºC) or below are
necessary for seed germination.
After the cold period, seeds germinate in both spring and summer at temperatures ranging
from 40º to 68ºF (4.5–20ºC). Although it was formerly believed to persist as a perennial
in mild climates that lack frost, that possibility has been disproved.
Reproduction and Dispersal. Mile-a-minute plants have a long flowering season, June to
October, and flowers are self-pollinating. They are prolific seed producers. Fruit usually
develops from August to the date of the first frost, usually October, but, depending on site
conditions, can develop as early as June. Seeds are dispersed by birds, insects, and small
mammals. Birds disseminate seeds under utility lines, fence lines, and other perching loca-
tions. Ants have been seen carrying seeds short distances, and chipmunks, squirrels, and
deer also eat the seeds. Because vines frequently overhang streams, water provides a
628 n VINES

A. Climbing vines can overtop other vegetation. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology University
of Connecticut, Bugwood.org.) B. Plants, with distinctly triangular leaves, may form a dense mat in open areas.
(Britt Slattery, U.S. Fish and Wildlife Service, Bugwood.org.) C. The ocrea completely encircles the stem.
(Leslie J. Mehrhoff, Ecology and Evolutionary Biology University of Connecticut, Bugwood.org.) D. Flowers
and fruit emerge from the ocrea. (Richard Old, XID Services, Inc.). E. Stems are covered with barb-like spines.
(Richard Old, XID Services, Inc.)

significant means of transportation in riparian locations, especially during storm events.

Fruits remain buoyant for 7–9 days. The population in North Carolina may have originated
from seeds floating down stream. Although newly discovered in summer of 2010, this new
infestation may already be too large and dispersed to eradicate. Most seeds germinate in full
sun in early spring.
Impacts. This rapidly growing vine smothers herbs, shrubs, and small trees. The inter-
tangled mat of vines covers all vegetation, blocking their foliage from light, thereby killing
them because they cannot photosynthesize. Mile–a–minute causes a reduction in plant
diversity, resulting in less food and shelter for wildlife. The vine has an economic impact
because it is a threat to pine plantations, reforestation efforts, and Christmas tree farms,
quickly smothering seedlings. It can also overrun plants in commercial nurseries and
orchards. The densely spiny plant is a hazard in urban and recreational areas. Plants are also
considered a weed harmful to agriculture in some parts of its native range.
Management. The best way to prevent infestations of mile-a-minute is to maintain the
stability of the native plant communities and avoid creating gaps where it can invade.
Intact vegetative cover along streams, roadsides, and other corridors provides no bare
ground for the plant to gain a foothold. Because the root system is weak and not persistent,
small vines can be removed by physical means. Hand-pulling should be done while plants
are still young, before the curved spines harden, and before seeds mature. Pulling vines that
contain fruit may result in a dispersal of more seeds. Repeated mowing or cutting the vines
close to the ground will reduce or prevent seed production. Because they do not resprout
or root, severed vines can be rolled up and left to dry.
The most effective chemical control is glyphosate, which kills existing plants, followed by
a preemergent, such as imazapyr or atrizine, early the following spring to prevent germina-
tion of seeds. Because seeds are persistent in the soil bank, remaining viable for as long as
six years, several applications of herbicide applications are necessary.
Although some insects feed on mile-a-minute, no native species does significant damage.
Of approximately 100 natural enemies in China, a few show promise for effective biological
control. A weevil (Rhinoncomimus latipes), which is host-specific to mile-a-minute, was
 ORIENTAL BITTERSWEET n 629

Preventing Spread

A lthough mile-a-minute is currently concentrated around the area where it was

first introduced, it is present in only 20 percent of its potential range. Several
other states provide suitable climate conditions. The mature plant, an annual with
a shallow root system, is readily killed by herbicides and will not resprout. Its seeds
have no means of extremely long-distance dispersal to widely disjunct locations
across the United States except by human activity. If people are diligent about
eliminating the plant when it first appears and are extremely careful not to trans-
port seeds, the spread of this invasive species can be halted.

released into affected eastern states at various times beginning in 2004 and continuing to the
present, with planned releases for 2011. This weevil produces several generations from
spring to fall, with adults feeding on foliage, and larvae boring into the plant stems.
Reports of a second weevil (Homorosoma chinensis) were subsequently found to be a misiden-
tification of R. latipes. Two geometrid moths (Timandra griseata and T. convectaria) feed on
young leaves and buds, defoliating the plants. Adults and larvae of a bug (Cletus schmidti)
eat the skin of the fruit, exposing and damaging the developing seed. Although some of these
insects will attack other species in the buckwheat family, their feeding preference is mile-a-
minute. A sawfly (Allantus nigrocaeruleus) and fungal pathogens are also undergoing
research.

Selected References
Okay, Judith A. Gerlach, Judith Hough-Goldstein, and Jil M. Swearingen. “Mile-A-Minute, Persicaria
perfoliata.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant Conservation
Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/pdf/pepe1.pdf.
“Pest Alert, Mile-A-Minute Weed (Persicaria perfoliata).” U.S. Department of Agriculture, Forest
Service, Northeastern Area, State and Private Forestry. Newtown Square, PA, 2009. http://
www.na.fs.fed.us.
Wu, Yun, C. Readon, and Ding Jian-qing. “Mile-a-Minute Weed.” In Biological Control of Invasive Plants
in the Eastern United States, by R. Van Driesche et al. USDA Forest Service Publication FHTET-2002
-04, 2002. http://www.invasive.org/eastern/biocontrol/26MileAMinute.html.

n Oriental Bittersweet
Also known as: Round-leaved bittersweet, Asian bittersweet, Asiatic bittersweet,
climbing spindleberry
Scientific name: Celastrus orbiculatus
Synonyms: Celastrus articulatus
Family: Staff-tree (Celastraceae)
Native Range. Temperate East Asia, including eastern Mongolia, northeastern China,
Korea, and Japan.
Distribution in the United States. The eastern half of the United States, from Minnesota
south to Louisiana, east to Georgia and South Carolina, and north to Maine. It is cultivated
in the Pacific Coast states, but has not yet been reported as naturalized in that region.
630 n VINES

Description. Oriental bitter-

sweet is a deciduous, woody
perennial vine or trailing shrub.
With no tendrils or adventitious
roots, it climbs vegetation and
structures by twining around
the object for support. Its light-
or dark-brown twigs are round
and glabrous, with noticeable
lenticels that appear as raised
whitish corky dots. Older
branches become olive drab or
pale gray-brown with elongated
lenticels and a scaly or shred-
ded appearance. Branches can
climb as high as 60 ft. (18 m)
and can be as much as 5 in.
(13 cm) in diameter. Leaves are
variable in size and shape, but
are generally broadly oblong or
almost round, 1–5 in. (2.5–
12.5 cm) long and 0.5–3 in.
(1.5–8 cm) wide. Leaf bases are
generally obtuse, while leaf tips
are acute to rounded. The
glossy, finely toothed leaves,
alternate on the stem, are sup-
ported on light-green petioles,
0.4–1.2 in. (1–3 cm) long. In
fall, leaves turn lemon- to
golden-yellow, enabling plants
Although Oriental bittersweet is cultivated as an ornamental in the to be seen after most other
western states, it has only become naturalized and invasive in the deciduous leaves have fallen.
Midwest and East. (Native range adapted from USDA GRIN and selected The outer surfaces of the roots
references. Introduced range adapted from USDA PLANTS Database, are a bright orange.
Invasive Plant Atlas of the United States, and selected references.)
Oriental bittersweet can be
either dioecious or monoecious.
In May and June, clusters of 3–7 small, inconspicuous flowers grow from the leaf axils. The
greenish-white or greenish-yellow flowers initially have both male and female parts, but one
type is aborted or reduced, resulting in unisexual flowers and unisexual plants. A few vines,
however, retain both types of flowers. Flowers have five petals and, if male, five stamens.
Many clusters of fruit, which are usually shorter than the leaves, develop at many points along
the stem. The fruit is a globular capsule, approximately 0.25 in (7 mm) in diameter, with an
outer covering that changes from green to bright yellow as it ages. The mature yellow capsule
splits open to reveal three red-orange fleshy arils, each with 1–2 brown or yellow seeds. The
three parts of the capsule become folded back, looking like three yellow wings behind the arils.
Related or Similar Species. The native American bittersweet, which grows in similar hab-
itats, is differentiated by flowers and fruit. Plants have a single panicle of flowers at the tips of
 ORIENTAL BITTERSWEET n 631

A. Vines climb by twining around objects. (James H. Miller, USDA Forest Service, Bugwood.org.) B. Older and
larger vines can girdle and kill their hosts. (Chris Evans, River to River CWMA, Bugwood.org.) C. Leaves are
usually oblong or round, with pointed tips. (James H. Miller, USDA Forest Service, Bugwood.org.) D. Fruit are
round capsules. (James R. Allison, Georgia Department of Natural Resources, Bugwood.org.) E. Clusters of
inconspicuous flowers grow from leaf axils. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology University of
Connecticut, Bugwood.org). F. The three parts of the capsule fold backward to look like wings. (James R.
Allison, Georgia Department of Natural Resources, Bugwood.org.)

the stems, and the clusters of flowers and fruit are approximately the same length as the
leaves. The plant has fewer fruit, but in larger clusters, and the fruit is orange when ripe.
Although identification by leaf shape and size is not reliable, leaves of American bittersweet
tend to be twice as long as they are wide, and tapered at each end. Hybrids occur between
American bittersweet and Oriental bittersweet.
Introduction History. Oriental bittersweet was bought to the United States in the 1860s
as an ornamental. It is often associated with old homesites, from where it has escaped to
natural areas and has become naturalized in several states. Oriental bittersweet itself, or a
plant mistaken for the American bittersweet, was also used extensively as highway landscape
plantings and for erosion control. It has been promoted as enhancement for wildlife, provid-
ing food and cover. The fruit is showy and attractive, and fruiting branches are frequently
used in floral arrangements. It is still available in the horticultural trade, often mislabeled
as American bittersweet.
Habitat. Oriental bittersweet has wide environmental tolerances. It grows at the edges of
forests and marshes, in woodlands and coastal areas, along hedgerows, and in fields in the
early successional stages. It is especially prominent in disturbed sites in meadows, thickets,
beaches, and young forests. It grows best in open, sunny sites, but is tolerant of shade and
can invade dense canopied forests. It can grow at elevations as high as 7,250 ft. (2200 m).
Reproduction and Dispersal. Oriental bittersweet reproduces both by seed and by root
suckering. Flowers are pollinated by bees and by wind. Fruit remains on the vine through-
out the winter, providing food for several birds, such as Black-capped Chickadees,
Mockingbirds, Blue Jays, and European Starlings, as well as for small mammals, all of which
distribute the seeds. Human use of the fruiting stems in dried flower arrangements also con-
tributes to dispersal, as old branches are disposed of in compost or discarded in brush.
Seeds germinate in late spring.
Oriental bittersweet produces copious root suckers from rhizomes, especially when the
plant is damaged or cut. Clones or thickets can evolve from only one or a few plants.
632 n VINES

Impacts. Because it grows quickly and sprouts from its roots, Oriental bittersweet com-
monly creates monotypic stands. This aggressive vine threatens both understory and canopy
plants of forested and open areas. By climbing and overtopping plants, it kills them by gir-
dling and preventing photosynthesis. The weight of the vine can cause host plants to become
uprooted during high wind or heavy snows. Although the plant is most noticeable along
roadsides, its greatest impact is in forests, where it resembles kudzu in the way it blankets
existing vegetation.
It outcompetes and displaces the native American bittersweet. The germination rate of
Oriental bittersweet averages 70 percent, compared with 20 percent for American bitter-
sweet, especially at low light intensities. Evidence exists that Oriental bittersweet is able to
recover more quickly from grazing than native bittersweet does, giving it a further advan-
tage. Hybridization between the two species threatens the genetic identity of American
bittersweet, a plant that is becoming increasingly rare.
Because the plant can grow in pure sand, Oriental bittersweet vines may spread into bird
nesting areas on beaches or alter dune formation. By doing so, it may threaten the breeding
behavior of the Piping Plover, a federally threatened species, on Long Island Sound.
Fruit of the Oriental bittersweet is toxic to humans.
Management. A combination of physical and chemical methods works most effectively. It
is questionable, however, whether infested natural areas can recover after removal of
Oriental bittersweet because of too many vegetative changes and disruptions to the natural
ecosystem. Clonal thickets are especially hard to kill. After a 250 sq. ft. (25 m2) plot was
treated with triclopyr in 1986, the area recovered to produce over 50 sprouts each sub-
sequent year. It took six years to exhaust the seed bank. Re-infestation of any area is possible
from other sources through long-distance dispersal of seeds.
Physical control can be accomplished by hand-pulling small plants, including the entire
root system, before fruit matures. If fruit is present, plants should be bagged and disposed
of, to prevent seed dispersal. An alternative method is to leave the bag in the sun to bake,
which will kill any seeds. On climbing vines, the best method is to sever the vines and treat
the rooted base with herbicide. Cutting dense, low-growing patches close to the ground in
early spring, followed by spraying new growth after about a month, can completely kill the
plants.
For large infestations, especially where pulling or uprooting vines would do too much
damage, chemical applications of glyphosate or triclopyr are effective. Foliar sprays, which
are also absorbed through the basal bark of the vines, are best done in fall after nontarget
plants become dormant. Foliage of new seedlings may also be sprayed. Although herbicides
may be applied directly to basal bark of the plants, chemical treatments work best when
applied to cut stems. Application should be done in fall and winter, before ephemerals
emerge in spring, to avoid damaging native plants. Repeated treatments may be necessary.
Because it targets only broadleaf plants, triclopyr is preferred where nontarget vegetation
includes grasses, sedges, and members of the lily family.
No biological controls are known.

Selected References
Burnham, R. J. “Plant Diversity Website, Celastrus orbiculatus.” CLIMBERS Website, 2010. http://www
-personal.umich.edu/~rburnham/SpeciesAccountspdfs/CelascanCELAFINAL.pdf
Dreyer, Glenn D. “Element Stewardship Abstract, Celastrus orbiculatus.” Global Invasive Species Team,
Nature Conservancy, 1994; modified 2009. http://wiki.bugwood.org/Celastrus_orbiculatus.
 PORCELAINBERRY n 633

Swearingen, Jil M. “Oriental Bittersweet.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas.
Plant Conservation Alliance’s Alien Plant Working Group, 2006. http://www.nps.gov/plants/alien/
fact/pdf/ceor1.pdf.

n Porcelainberry
Also known as: Porcelain vine, porcelain ampelopsis, creeper, wild grape,
Amur peppervine
Scientific name: Ampelopsis glandulosa var. brevipedunculata
Synonyms: Ampelopsis brevipedunculata, Cissus brevipedunculata, Vitis brevipedunculata,
Ampelopsis heterophylla
Family: Grape (Vitaceae)
Native Range. Extreme northeastern China.
Distribution in the United States. Eastern states, from Wisconsin and Iowa south to
Alabama and Georgia, and north to New York and New Hampshire. Especially invasive in
the northeastern states.
Description. Porcelainberry is a woody perennial vine. It climbs by using twining, nonad-
hesive tendrils, which grow opposite the leaves. Although vines are mature at approximately
10 ft. (3 m) long, they can climb as high as 20 ft. (6 m). Plants running along the ground
have an upright, spreading habit, and may form a dense matted ground cover. Vines need
support in order to climb. The thin green twigs grow in a slightly zigzag pattern for the first
year and become brown and semi-woody in their second season. The white stem pith is con-
tinuous across the stem nodes. The bark has lenticels and does not peel. Leaves, alternate on
the stem, are broadly oval and palmate in shape, 5 in. (12.5 cm) long. Leaves may be either
shallowly or prominently cleft, forming 3–5 coarsely toothed lobes. Plants with the deeper
lobes are known as the variety Ampelopsis glandulosa var. heterophylla (formerly A. brevipe-
dunculata var. maximowczii). The leaf base is heart-shaped, and the tip may be either
rounded or pointed. Leaves are shiny, ranging from medium to dark green to blue-green.
Leaves may have delicate hairs along the veins, and young twigs and petioles are hairy.
Porcelainberry is deciduous, but leaves do not change color in fall. The taproot is large
and vigorous.
The small, inconspicuous creamy-green flowers appear June through September on the
current season’s growth. They are densely arranged in cymes opposite the leaves. The fruit
are hard berries, hanging in small, globular clusters. They change color as they mature and
can be white, yellow, green, lilac, purple, turquoise, sky blue, or bright blue. Dots give the
fruit a speckled or marbled appearance. Because of the long flowering season, all colors of
fruit may occur on the vine, or even in one cluster. Each small fruit, 0.25 in. (0.6 cm), which
matures in September or October, contains 2–4 seeds.
Related or Similar Species. A cultivar, known as ‘Elegans,’ is a deeply lobed and vari-
egated plant available in nurseries. It has smaller leaves that are mottled with white and
sometimes pink coloration, and is less vigorous and invasive than the species. Boston ivy,
also known as Japanese creeper, is a deciduous, climbing vine native to Asia. It has simple
lobed leaves with serrated edges and hairy leaf veins. It can be distinguished from porcelain-
berry by adhesive tips on the ends of the tendrils, blue fruit on red pedicels, and foliage that
turns red in the fall.
Two native species of Ampelopsis are widespread in the southeastern United States.
Peppervine is a ground cover or climbing vine that has few tendrils. It is invasive in some
634 n VINES

regions. This plant is easily dis-

tinguished by its bi- or tripin-
nately compound foliage,
which turns pale yellow in the
fall. The inedible berries are
bluish purple, becoming black
and shiny as they ripen. Heart-
leaf peppervine uses tendrils,
including twining flower stems,
to climb trees in riparian sites,
but it is not invasive. The
broadly oval leaves, as much as
5 in. (12.5 cm) long and 4 in.
(10 cm) wide, are not lobed
and have a heart-shaped base.
Berries are purple.
Although porcelainberry
resembles native grapes, grape
vines have distinguishing char-
acteristics. Grape pith is dark
brown and is not continuous
across stem nodes. The bark
peels and has no lenticels, and
flowers occur in panicles.
Introduction History. Porce-
lainberry was introduced to the
United States in the 1870s as a
landscape plant because of its
colorful berries. It also makes a
good ground cover or trellis-
climbing vine and was used as
Found in the upper midwestern and eastern states, porcelainberry prefers a bedding plant and screening
moist soils but is tolerant of drought (Native range adapted from USDA plant. The variety ‘Elegans’ was
GRIN and selected references. Introduced range adapted from USDA brought to the United States
PLANTS Database, Invasive Plant Atlas of the United States, and selected before 1847, either by direct
references.)
plantings or mixed in bird seed.
Pest resistant and tolerant of
adverse conditions, porcelainberry plants are still available in the horticultural trade.
Habitat. Porcelainberry grows in several habitats, including disturbed sites, such as aban-
doned fields, vacant lots, and road, railroad, and utility right-of-ways. It can also be found at
forest edges, pond margins, along stream banks, in pastures, and in successional or planted
forests. It grows on most types of soil, rich to poor, and with various levels of acidity or alka-
linity. It prefers moist soil, but does not tolerate sites that are permanently wet. Plants can
withstand both drought and heat. Although it can be found in full sun to partial shade, por-
celainberry is less tolerant of heavy shade and absent from mature forest interiors. Elevation
limits are 500–2,000 ft. (150–600 m).
Reproduction and Dispersal. Porcelainberry reproduces both by seeds and vegetatively.
Birds and small mammals, such as squirrels, eat the berries and expel the seeds. Uneaten
 PORCELAINBERRY n 635

A. Vines climb into tree canopies. (Steven Manning, Invasiveplantcontrol.com.) B. and C. Depending on the
variety, leaves may be deeply or shallowly lobed. (James H. Miller, USDA Forest Service, Bugwood.org.)
D. Tiny flowers grow in cymes. (James H. Miller, USDA Forest Service, Bugwood.org.) E. Berries hang in small
clusters. (James H. Miller, USDA Forest Service, Bugwood.org.) F. Old stems become semi-woody. (James H.
Miller, USDA Forest Service, Bugwood.org.)

fruits split to release the seeds. Infestations that develop downstream from existing sites pro-
vide evidence that seeds may also be dispersed by water. Seeds germinate readily the follow-
ing year, but some remain viable in the soil for several years. Plants resprout when the top
portion is damaged or cut.
Impacts. Porcelainberry invades both open and wooded habitats. Seeds grow well after a
disturbance, either natural or human-caused. Its thick mat of vines cover and shade out
native shrubs and small trees. The weight of the vines makes the host plant susceptible to
wind and ice damage. When provided with high to moderate light and plenty of water, the
plant spreads fast, and vines can grow 15 ft. (4.5 m) in one season.
Management Any management program requires monitoring and follow-up. Some meth-
ods must be repeated, not only during the growing season but for several years. The major goal
is to prevent flowering, fruiting, and production of mature seeds. Physical control includes
hand-pulling vines in either fall or spring to prevent flowers the following summer. To avoid
seed dispersal, vines should be pulled up before fruit matures. If the plant is in fruit, cuttings
should be bagged and disposed of in a landfill. Large vines may be cut close to the ground and
monitored for resprouts, pulling them soon after they emerge. Cutting vines in the fall will not
prevent flower development because flowering occurs on the current season’s growth.
Chemical treatment with herbicides may be necessary. Triclopyr or glyphosate may be
applied to large vines that have been cut at ground level. Triclopyr is the most effective,
either as a foliar spray or applied to cut plants. Applications of triclopyr ester to sections of
stem 2–3 ft. (.6–1 m) long work well.
Although 9 species of fungi and 13 species of arthropods are natural enemies of porce-
lainberry in its native range, no biological controls are available. Japanese beetles (Popillia
japonica) attack landscape plants.

Selected References
“Ampelopsis brevipedunculata.” The Ohio State University, n.d. http://www.hcs.ohio-state.edu/hcs/TMI/
Plantlist/am_ulata.html.
“Porcelainberry, Amur Peppervine.” Invasive Plant Atlas of New England (IPANE), University of
Connecticut, 2009. http://www.invasive.org/weedcd/pdfs/ipane/Ampelopsisbrevipedunculata.pdf.
636 n VINES

Swearingen, J., K. Reshetiloff, B. Slattery, and S. Zwicker. “Porcelainberry.” Plant Invaders of Mid-
Atlantic Natural Areas. National Park Service and U.S. Fish and Wildlife Service, 2002. http://
www.invasive.org/eastern/midatlantic/ambr.html.
Young, Jamie. “Porcelain-berry.” Weeds Gone Wild: Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2005. http://www.nps.gov/plants/alien/fact/
pdf/ambr1.pdf.

n Swallow-Worts
Pale Swallow-Wort
Also known as: Swallow-wort, swallowwort, European swallow-wort, dog-strangling vine
Scientific name: Cynanchum rossicum
Synonyms: Vincetoxicum rossicum, Cynanchum medium, Vincetoxicum medium,
Antitoxicum rossicum
Family: Milkweed (Asclepiadaceae)
Black Swallow-Wort
Also known as: Black dog-strangle vine, Louis’ swallow-wort, climbing milkweed
Scientific name: Cynanchum louiseae
Synonyms: Vincetoxicum nigrum
Family: Milkweed (Asclepiadaceae)
Native Range. Pale swallow-wort is native to the Ukraine and to southwestern
European Russia, perhaps endemic to regions north of the Black Sea. Black swallow-
wort is from Mediterranean southwestern Europe, including Italy, France, Spain, and
Portugal.
Distribution in the United States. Midwestern and New England states. Pale swallow-
wort is more frequently found in the Great Lakes region and New England. Black swallow-
wort is more widespread, from Nebraska and Kansas east through New England, and has
also been found in California.
Description. Swallow-worts
are long-lived, herbaceous per-
ennial scramblers or vines that
climb by twining slender stems
around themselves or around
other plants. Stems do not
branch, and depending on the
height of the supporting struc-
ture, they can grow 2–8 ft.
(0.6–2.4 m) long. Typical of
members of the milkweed fam-
ily, stems and leaves exude a
milky latex when cut. Pale
swallow-wort stems have dis-
tinct lines of dense pubescence,
which are absent or sparse in
black swallow-wort. Black
Swallow-worts were brought to the United States as ornamental plants. swallow-wort stems may turn
(Native range adapted from USDA GRIN and selected references.) red in sunny locations. Dark-
 SWALLOW-WORTS n 637

green leaves are oblong to oval or lance-shaped, 2–5 in. (5–13 cm) long and 0.5–3 in.
(1.5–8 cm) wide. Leaf arrangement is opposite, and petioles are 0.2–0.8 in. (0.5–2 cm) long.
Leaf tips are pointed, bases are rounded, and the margins are entire (not serrated). Upper
leaf surfaces are glabrous and shiny, but lower surfaces, especially the veins, and leaf mar-
gins are pubescent. The pubescence on black swallow-wort leaves may be red. The largest
and widest leaves grow in the middle of the vine, while the leaves at both the base and the
growing tip are smaller. Both species have a fleshy, fibrous root system that is firmly anch-
ored in the soil.
Star-shaped flowers, appearing May through June, are small, approximately 0.25 in.
(6 mm). They grow in small clusters of 4–20 in leaf axils. Open for 6–8 days, flowers emit
a slight odor variously described as sweet or of rotting fruit. Petals of pale swallow-wort
flowers are pale yellow-purple, creamy pink, or reddish brown. The flowers have a distinctly
five-lobed pink, orange, or yellow ring, called a corona, surrounding pale green or yellow-
green stamens. Black swallow-wort flowers have purplish, almost black petals, with a
maroon corona encircling bright yellow-green stamens. Petals of pale swallow-wort are gla-
brous, while the upper surface of black swallow-wort petals are covered with short, white
hairs, giving the flower a downy appearance.
Fruit are slender, tapered seed pods, 1.5–3 in. (4–8 cm) long and 0.25 in. (6 mm) wide,
often occurring in pairs on the stem. When mature, they turn light brown and split along a
longitudinal seam to release flat, brown seeds. Each seed has a thin membrane along its mar-
gins and a tuft of silky hair. Opened pods appear very silky inside. Although plants turn yel-
low in late summer and die back to the ground in winter, the dried stems may remain, often
with open seed pods with remaining seeds.
Related or Similar Species. White swallow-wort, native to most of Europe, grows
sparsely in the northeastern United States. It does not seem to be invasive in most areas,
but may be threatening green comet milkweed, an endangered species in Connecticut.
Although the United States has several native Cynanchum species, honeyvine is the only
one growing in the midwestern and northeastern United States where swallow-worts are
found. Honeyvine has white flowers, and the bases of its leaves are distinctly heart shaped.
Honeysuckles have distinctly pea-shaped flowers and semi-woody stems. Climbing
nightshade, also an invasive species, has similar-looking star-shaped flowers but is distin-
guished by its berries and leaves, which are alternate and either serrated or lobed. Neither
honeysuckles nor nightshade have latex.
Introduction History. Although they are no longer desirable as ornamental plants,
swallow-worts were probably brought to the United States for horticultural reasons. Pale
swallow-wort was first collected in nature in New York State in 1897. Black swallow-wort
was first collected from a natural area in Massachusetts in 1854, probably escaped from a
botanic garden where it was a weed. Scattered occurrences of both species indicate that they
were probably garden escapees, but swallow-worts are easily dispersed to new areas in con-
taminated hay.
Habitat. Both swallow-wort species are found in similar habitats, primarily upland areas.
They have a wide tolerance to light and moisture conditions, growing both in dry sunny
locations and moist shaded locations, but avoid sites with long-standing water. Plants in
shady sites, however, are less vigorous, with thinner stems, larger and darker leaves, and
fewer roots. Swallow-worts grow in oak and mixed-hardwood forests, prairies, and along
fencerows. Commonly found in disturbed sites, especially human-caused, they grow along
transportation and utility right-of-ways, in quarries, and in abandoned pastures and
agricultural fields. They also invade natural areas that have been disturbed, such as ice- or
638 n VINES

flood-scoured river banks and

talus slopes. Swallow-worts
may dominate a shady under-
story for years, and when a dis-
turbance opens the forest
canopy, the plant will quickly
grow, flower, and set seed.
Swallow-worts are especially
competitive and form large
populations on shallow soils
over limestone, areas prone to
summer drought because the
limestone fails to hold water
after the surface soil dries.
Plants are also found on deep,
well-drained loamy soils. Black
swallow-wort has a less distinct
association with limestone and
is also found on granitic sub-
strate. This species also grows
on rocky coasts, above the
high-tide line, an indication
that it is somewhat salt tolerant.
Reproduction and Dispersal.
Swallow-worts spread primarily
by seed and are prolific produc-
ers. A thick stand in full sun
can produce as many as 200
seeds per sq. ft. (2,000 per m2).
The number of flowers and seed
pods produced is directly
Both swallow-wort species are found primarily in the upper midwestern related to the amount of light.
and northeastern states, in sites disturbed by human activity. Plants in deep shade fail to
(Introduced range adapted from USDA PLANTS Database, Invasive
flower. Flowers are both self-
Plant Atlas of the United States, and selected references.)
pollinating and insect-polli-
nated, by bees, wasps, flies, ants,
and beetles. Seeds are also polyembryonic, meaning that 1–4 plants can emerge from one seed.
Long-distance dispersal is accomplished by wind-disseminated seeds. Many seeds are not via-
ble. Greenhouse tests indicate that under normal conditions, the germination rate is less than
25 percent, and still less than 50 percent under ideal conditions. Some seeds germinate in fall,
others in spring. In gardens, seedlings will flower and produce seed in their second year of
growth. Both species sprout vigorously from dormant buds on the root crown when the top
portions of the plant are damaged, and black swallow-wort spreads locally by rhizomes.
Impacts. Swallow-worts grow in multi-stemmed clumps that outcompete native plants
for soil moisture, nutrients, and light. Both species overtop plants in brushy areas and domi-
nate the understory, but not the trees, of forests. They are problem weeds that require con-
trol in home gardens, perennial pastures, nurseries, and Christmas tree and other tree
plantations, but they are usually not weedy in annual crops.
 SWALLOW-WORTS n 639

A. Vines can cover and smother all other vegetation. B. Vines climb by twining around objects. C. Flowers of pale
swallow-wort are light colored. D. Flowers of black swallow-wort are very dark purple. E. Seed pods are filled
with silky filaments. (Leslie J. Mehrhoff, Ecology and Evolutionary Biology, University of Connecticut,
Bugwood.org.)

Patches of swallow-wort displace or suppress native plant species and impact wildlife.
Swallow-worts replace goldenrods and grasses in old fields and pastures, disrupting the pro-
cess of natural succession. They threaten rare and endangered species. In at least one region
in New York State, pale swallow-wort is displacing Hart’s tongue fern. In Vermont, black
swallow-wort threatens Jesup’s milkvetch, an endangered endemic species. Also in New
York State, swallow-worts threaten the rare and unique plant communities of limestone
pavement barrens, called alvars. Plants modify the microbial community in the soil, thereby
displacing the native species especially adapted to that environment.
Neither swallow-wort species is readily used by native animals or insects. A study in New
York State indicated that an increase in pale swallow-wort was followed by a decrease in
native grassland birds. A Canadian study showed a decrease in diversity of ground and stem
insects, and a total lack of gall-makers and miners. The effect of black swallow-wort on birds
and insects is not known. Because they are in the milkweed family, swallow-worts may det-
rimentally affect monarch butterflies, which are obligated to lay their eggs on milkweeds.
Eggs laid on swallow-worts do not survive. As swallow-worts become more dominant in
the natural landscape, where they have displaced many Asclepia species necessary to the but-
terfly, populations of monarch butterflies could become endangered.
Management. Early detection is important. Because swallow-wort produces so many
seeds, if one plant is found, it is likely that more are nearby. Any management method
should be carried out before seed sets. Because human activity may promote dispersal, areas
that are actively releasing seed should be avoided, unless all the seeds can be collected. Any
machinery or vehicles should be thoroughly cleaned to ensure that no seeds are carried
away. If appropriate to the site, revegetation with grasses and subsequent spraying with a
broadleaf herbicide to kill swallow-wort seedlings will eventually deplete the seed bank.
Physical control is limited. Hand-pulling is less effective than digging out plants because
the stem base is brittle, breaking off just above the root crown. Root crowns should be com-
pletely dug out because even a fragment may sprout a new plant. Any pod-bearing plants
should be bagged and destroyed, taken to a landfill or burned. Mowing several times a year,
when seed pods are small, will prevent seed formation, but timing is critical. If mowed
during the flowering stage, plants will recover and sprout new stems.
Although chemical treatments are best for large populations, the choice of herbicide is
dependent on location and other plants. Herbicides should be applied after flowering begins
but before seed pods form. At that time, plants are large enough to carry lethal doses of
640 n VINES

herbicide to the roots, and no seeds will be produced. Because of the seed bank, several
years may be needed to eradicate the population. Both glyphosate and triclopyr ester are
effective, but glyphosate is better for cut stems. If seed pods are present, the stems with pods
should be collected and bagged or seeds will continue to develop. Because foliar sprays may
not penetrate the plant canopy, allowing seedlings and small plants to survive, repeated
applications may be necessary. An alternative is to cut or mow plants that have seed pods
and spray the regrowth in early fall.
No biological controls are known for swallow-worts. Except for the tarnished plant bug
(Lygus lineolaris), no insects or pathogens in the United States are prominent in swallow-
wort stands, and the plant bug does little damage. It is possible that swallow-worts may be
toxic to North American insects. Little is known of natural enemies in their native range.

Selected References
Lawlor, Fran. “Element Stewardship Abstract, Vincetoxicum nigrum and Vincetoxicum rossicum,
Swallow-wort.” Global Invasive Species Team, Nature Conservancy, 2001; updated 2002. http://
www.invasive.org/gist/esadocs/documnts/vinc_sp.pdf.
Sonday, ReBecca, and R. Burnham. “Plant Diversity Website: Cynanchum rossicum (Kleopow) Borhidi”
and “Plant Diversity Website: Cynanchum louiseae Kartesz and Ghandi.” CLIMBERS Website, 2010.
http://www-personal.umich.edu/~rburnham/climbers.html.
Tewksbury, L., R. Casagrande, and A. Gassmann. 2002. “Swallow-Worts.” In Biological Control of Invasive
Plants in the Eastern United States, by R. Van Driesche et al. USDA Forest Service Publication FHTET
-2002-04, 2002. http://www.invasive.org/eastern/biocontrol/16SwallowWorts.html.

n Winter Creeper
Also known as: Euonymus, dwarf euonymus, wintercreeper, climbing euonymus
Scientific name: Euonymus fortunei
Synonyms: Euonymus hederaceus, E. japonicus, E. japonicus var. acutus, E. radicans var. acutus,
Elaeodendron fortunei,
Family: Staff-vine (Celastraceae)
Native Range. East-central China, Korea, Japan, and the Ryukyu Islands.
Distribution in the United States. Scattered in the eastern and midwestern states, gener-
ally east of the Mississippi River, but also from Nebraska south to Texas and Louisiana north
to Arkansas. From Wisconsin south to Alabama, east to Georgia, throughout the Atlantic
coast north to Maine, and west to Michigan.
Description. Winter creeper is an evergreen perennial with two growth habits. It can be a
dense ground cover or shrub as much as 3 ft. (0.9 m) tall, or it can be a climbing vine. Aerial
roots, used for climbing trees, rocks, and vertical structures, grow from stem nodes, enabling
vines to climb as high as 40–70 ft. (12–21 m). Trailing plants also have many rootlets, which
give the narrow stems a warty appearance. Initially lime green, the hairless stems become
streaked with light-gray reddish bark as they age. Older stems have gray corky bark, at first
fissured, then checkered with fissures. The broadly oval leaves, which are densely arranged
and opposite on the stem, are 1–3 in. (2.5–8 cm) long and 1–1.8 in. (2.5–4.5 cm) wide. The
thick, leathery leaves are smooth and glossy, with prominent silvery-white veins. Leaf blades
are either dark green or green and white variegated on the upper surface. They are light
green below. Leaf edges are finely serrated and slightly turned under or wavy. The leaf base
tapers to the 0.1–0.4 in. (2–10 mm) petiole, and leaves are sometimes sessile.
 WINTER CREEPER n 641

Long cymes of inconspicu-

ous flowers, ranging in color
from white to green, grow from
leaf axils in June and July.
Flowers are 0.1 in. (2–3 mm)
in diameter, with five petals.
Insect-pollination produces
0.25–0.5 in. (6–12 mm) pink-
ish to red, smooth capsules,
hanging either singly or in
groups of two. When ripe,
September through November,
the fruit splits to expose a single
seed. Seeds are covered with a
fleshy orange seed coat, called
an aril.
Related or Similar Species.
Three Euonymus species are
native to the eastern United
States. Running strawberry
bush, also called running euo-
nymus, is a deciduous trailing
shrub or prostrate vine with
thin leaves. Its greenish-purple
flowers are 0.2–0.3 in. (6–
8 mm) in diameter. Fruits are a
three-lobed red capsule, and
the seeds have red arils.
Bursting heart is an understory
shrub in eastern forests, with
distinctive four-angled twigs
and bright red fruit capsules A common ornamental, winter creeper is often found near old home sites
and seeds. Burning bush and is especially widespread in midwestern and eastern states. (Native
has pink capsules with red range adapted from USDA GRIN and selected references. Introduced
seeds. Species of blueberry that range adapted from USDA PLANTS Database, Invasive Plant Atlas of the
have large leaves can be distin- United States, and selected references.)
guished by their alternate arran-
gement. The leaves of rusty blackhaw, a deciduous shrub or small tree native to the
Southeast, are opposite and as long as 2–4 in. (5–10 cm). They are thick and shiny. The
plant has a rusty-red pubescence on young stems and buds, clusters of creamy-white flow-
ers, and purple or dark-blue fruit about 0.5 in. (1.3 cm) long. Coral honeysuckle is easily
distinguished by its clusters of bright pink 2 in. (5 cm) tubular flowers and red berries.
Several introduced species have similarities to winter creeper. Although leaves near the
inflorescences of English ivy (see Vines, English Ivy) may resemble winter creeper leaves,
all other leaves on English ivy are five-lobed with palmate veins. Oriental bittersweet
(see Vines, Oriental Bittersweet), a deciduous shrub or trailing vine, has alternate 2–5 in.
(5–12.5 cm) leaves. Its stems may be 2–4 in. (5–10 cm) in diameter. Although the arils on
its seeds are red-orange, the mature fruit is green to yellow. Japanese honeysuckle (see
642 n VINES

A. This vine can create a dense ground cover. (James H. Miller, USDA Forest Service, Bugwood.org.) B. It can also
climb by using aerial roots. (Chris Evans, River to River CWMA, Bugwood.org.) C. Inconspicuous flowers form
in clusters from leaf axils. (James H. Miller, USDA Forest Service, Bugwood.org.) D. Each fruit contains one seed.
(James H. Miller, USDA Forest Service, Bugwood.org.) E. Aerial roots grow from nodes on the stems. (James H.
Miller, USDA Forest Service, Bugwood.org.)

Vines, Japanese Honeysuckle) lacks aerial roots and climbs by coiling its stems around the
host plant. Its fragrant tubular flowers are white, tinged with pink.
Introduction History. Winter creeper was brought to the United States in 1907 as a ground
cover and ornamental. Two varieties are common in North America, Euonymus fortunei var.
fortunei and E. fortunei var. radicans, and more than 50 cultivars, such as ‘Emerald’n Gold’
and ‘Gaity,’ are available in the horticultural trade. Cultivars vary by leaf size and color.
Habitat. Winter creeper is most frequently found in old or abandoned homesites and
near urban areas, where it escapes from gardens into adjoining forests. It may be in open
grasslands, gravel pit debris, rocky bluffs, and floodplains, and along roads. Shade tolerant,
it is quick to invade natural forest openings created by wind-throw, fire, or insect damage. It
also invades human-disturbed sites. This vine commonly occurs with other invasives, such
as multiflora rose (see Shrubs, Multiflora Rose) and oriental bittersweet. Winter creeper
grows in a variety of environments, including full sun to dense shade and on fine- to
coarse-textured soil. It prefers mesic sites, however, and is not found in heavy, wet soils or
in wetlands. It grows in several types of forest communities, including those dominated by
maples, oaks, elms, hickories, and pines. Although it can survive climates where the mean
minimum temperature is as low as −30ºF (−34ºC), it requires protection from winter snow
cover. It grows much better where minimum temperatures drop no lower than −20ºF
(−29ºC). In China, winter creeper grows from near sea level to 11,200 ft. (3,400 m) elevation.
Reproduction and Dispersal. Winter creeper reproduces both vegetatively and sexually.
As a ground cover, independent plants grow from rootlets along the stems that rest on the
ground. Lateral shoots grow from the main branches, and damage to the plant stimulates
more lateral stems and more leaves to emerge from existing nodes. When plants are cut,
sprouts develop from roots, root crowns, and cut stems.
Conflicting accounts of whether or not winter creeper flowers is probably because only
climbing plants with access to sufficient light produce flowers. Birds and wildlife that feed
on the arils are responsible for long-distance dispersal of seeds. Both seeds and plant pieces
can be transported by water to new sites. Little information is known concerning seed pro-
duction, seed bank, viability, germination, or seedling establishment. Seedlings, however,
 WINTER CREEPER n 643

are known to grow on disturbed or inhospitable sites, even through cracks in pavement.
Growth rates vary with environmental conditions, such as sunlight, insect damage, water,
and foliage damage by rabbits. Under the best conditions, plants need only one year to cover
75 percent of the ground with a dense mat of foliage.
Impacts. Because winter creeper is tolerant of many environmental conditions and grows
rapidly, it is a serious invader. Although some leaves do drop in spring, the plant is ever-
green and begins growing sooner than native ephemerals in spring, thereby replacing them.
It outcompetes native plants by depleting the soil of moisture and nutrients. As a ground
cover, it forms a dense mat that eliminates understory plants by impeding seedling growth.
By eliminating native ground-layer plants, it threatens native fauna, such as butterflies. As a
climbing vine over taller shrubs and trees, especially those less than 20 ft. (6 m) tall, it blocks
light and interferes with photosynthesis, weakening or killing the plants.
Management. The best management is to prevent its establishment. Minimizing its use as
a landscape plant and preventing fruit set will limit its ability to increase its range. Once
established, the entire plant must be removed. Because winter creeper predominantly
invades disturbed sites, maintenance of healthy communities and monitoring areas for signs
of regrowth several times a year can keep the plant in check.
Physical methods of control have limitations. Although labor intensive, small plants or
small populations can be hand-pulled or dug out, especially in sensitive environments. All
plant material, including roots and runners, must be removed, because any root pieces will
sprout. Debris should be bagged and removed from the site. Cutting plants without using
herbicides on the remaining rooted portion is ineffective because damage causes latent buds
to sprout. Mowing, which is not practical in natural areas, is also ineffective without follow-
up of treatment with a herbicide. Severing the climbing stems from the rest of the plant will
prevent fruit development because only climbing stems flower. At a site in Kentucky, cover-
ing a monoculture stand of winter creeper with black plastic for a growing season sup-
pressed its growth. The effectiveness of that method might be increased if the site were
kept covered for two seasons. Burning selected plants with a blow torch may provide control
when used in combination with other methods.
Physical control must be combined with chemical applications. Foliar sprays of triclopyr
ester or glyphosate can be applied from June through October, but must be repeated for sev-
eral years. Use of a string trimmer on trailing plants to reduce growth and injure leaves will
enhance the absorption of herbicide. Because glyphosate is nonselective, it should be used
only where no native plants will be harmed. Triclopyr is selective only to broadleaf plants
and can be used where grasses are intermixed with winter creeper. The best time for foliar
sprays is late fall or early spring, when native vegetation is dormant. Where plants are well
established or have climbed into tree canopies, application of herbicide to cut stems is effec-
tive, although it may have to be followed up with a foliar spray on resprouts.
No effective biological control is yet known, but winter creeper is one of the major Asian
invasive plants currently being studied in China for host-specific pests. It is susceptible to
Asian Euonymus scale (Unaspis euonymi), which occurs with the plant in most of its U.S.
range. By feeding on the leaves and causing them to drop, the scale damages tissue and
reduces the plant’s photosynthesis capability. The scale can be lethal, especially to some cul-
tivars, but it is also attacking native Euonymus species, such as burning bush and bursting
heart, and also kills the rare subshrub Canby’s mountain-lover. Release in southern New
England of five species of organisms from Asia in 1991–1995 provided no firm results.
Plants are susceptible to attack by a fungal parasite, Texas root rot (Phymatotrichum omnivo-
rum), which is endemic to the southwestern United States and adjacent Mexico.
644 n VINES

The parasite’s presence may prevent the spread of winter creeper into that geographic area.
Winter creeper is browsed by white-tailed deer in winter, and browsing by domestic goats
or sheep is a possibility to keep plants small and prevent them from climbing and flowering.

Selected References
Miller, James H. “Winter Creeper.” In Nonnative Invasive Plants of Southern Forests: A Field Guide for
Identificaiton and Control. Gen. Tech. Rep. SRS-62. U.S. Department of Agriculture, Forest Service,
Southern Research Station, 2003. http://www.invasive.org/eastern/srs/WC.html.
Remaley, Tom. “Climbing Euonymus.” Weeds Gone Wild; Alien Plant Invaders of Natural Areas. Plant
Conservation Alliance’s Alien Plant Working Group, 2009. http://www.nps.gov/plants/alien/fact/
pdf/eufo1.pdf.
Zouhar, Kris. “Euonymus fortunei.” Fire Effects Information System. U.S. Department of Agriculture,
Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, 2009. http://
www.fs.fed.us/database/feis/plants/vine/euofor/all.html.

n Wisteria
Chinese Wisteria
Also known as: Chinese-glysine
Scientific name: Wisteria sinensis
Synonyms: Kraunhia floribunda var. sinensis, K. sinensis, Rehsonia sinensis,
Wisteria sinensis var. albiflora, Glysine sinensis

Japanese Wisteria
Also known as: No other names
Scientific name: Wisteria floribunda
Synonyms: Kraunhia floribunda, K. floribunda var alba, K. japonica, Millettia japonica,
Rehsonia floribunda.
Family: Pea (Fabaceae)
Native Range. Chinese wisteria is from east- and south-central China. Japanese wisteria is
native to Japan.
Distribution in the United States. Both species are primarily in the southeastern and
eastern states, from Texas east to Florida, north to Maine, and west to Michigan, Illinois,
and Arkansas. Chinese wisteria is considered more invasive and is also in Hawai’i.
Description. These two exotic wisterias are deciduous, woody perennial vines that climb
trees and other structures. Because naturalized plants may be hybrids of the two, their character-
istics are variable. The height of vines, which can reach 65 ft. (20 m), is limited only by the
height of the object on which they climb. The vines have no tendrils or root hairs, but climb
by twining their stems around host plants or structures. Stems may be slender to stout, with
brown or white bark that may or may not be hairy. In old plants, stems can be 15 in. (38 cm)
in diameter. Leaves are alternate and pinnately compound, varying in length, 4–16 in. (10–
40 cm) and in number of leaflets, 7–19. Chinese wisteria usually has fewer leaflets, 7–13.
Leaflets, each 1.5–3 in. (4–8 cm) long, are oval to lance-shaped and have slightly wavy margins.
Long hanging clusters of fragrant violet to purple flowers appear in spring, before the
leaves fully develop. Individual flowers, supported by 0.6–0.8 in. (1.5–2 cm) pedicels, are
typically pea-shaped and 0.5–1 in. (1.3–2.5 cm) long. Fruit is a velvety, fuzzy seed pod,
either green or brown. The flattened pods are 4–7.5 in. (10–19 cm) long, narrowed at the
 WISTERIA n 645

base and with slight constric-

tions between the seeds. Pods,
each containing 2–6 seeds, are
retained on the vine after they
mature. The flat, round seeds
are 0.5 in. (1.3 cm) in diameter.
Although most naturalized
wisteria plants, at least in the
southeastern United States,
may be considered hybrids, the
original two species are distin-
guishable by several character-
istics. Chinese wisteria vines
twist clockwise, from lower left
to upper right, while Japanese
wisteria twists counterclock-
wise, from lower right to upper Japanese wisteria and Chinese wisteria are distinct in their native ranges
left. Chinese wisteria has dark- but hybridize in the United States. (Native range adapted from USDA
gray bark and hairy stems. GRIN and selected references.)
Japanese wisteria usually has
white or chalky-gray bark, and only the young stems are hairy. The flower clusters, called
racemes, on Chinese wisteria are shorter, 4–20 in. (10–50 cm) long, and the flowers in the
cluster all open together. Japanese wisteria racemes can be very long, 1–3 ft. (0.3–0.9 m),
and the flowers open sequentially from the base to the tip of the raceme.
Related or Similar Species. American wisteria, also called Atlantic wisteria, is native to
the southeastern U.S. coastal plain from Virginia south to Florida and west to eastern
Texas. It can be found climbing over shrubs and fences in moist thickets adjacent to water
bodies. Like the exotic wisterias, it can climb 50 ft. (15 m). The brown to reddish-brown
stems are hairless, and the vines twine clockwise. Leaves are usually 12 in. (30 cm) long,
with 9–15 leaflets, each 1–3 in. (2–8 cm). The lower leaf surface is a milky green color.
Purplish-blue flowers occur in May, after the plant has leafed out. Flower clusters are shorter
than those of either Chinese wisteria or Japanese wisteria, 1.5–6 in. (4–15 cm) long,
and pedicels are 0.2–0.4 in. (0.5–1 cm) long. The most distinct feature is that the 2–4 in.
(5–10 cm) brown seedpods are not hairy, but glabrous. Several cultivars, such as ‘Nivea’
with white flowers, ‘Magnifica’ with purple-blue flowers, and a yellow blotch, have been
developed from American wisteria. American wisteria is host for many butterflies and bees.
Trumpet creeper, native to the central and eastern United States, is distinguished from
wisterias by its opposite leaves with toothed margins and its orange-red flowers, which
bloom from late spring into autumn.
Introduction History. Both exotic wisterias were imported for ornamental use, Chinese
wisteria from China in 1816, and Japanese wisteria from Japan around 1830. Both species
are used extensively in the southeastern states to decorate porches, gazebos, and garden
walls. Because of its vigorous growth, wisteria escaped landscape plantings. Although
Chinese wisteria and Japanese wisteria were imported as two distinct species, recent evi-
dence indicates that naturalized and invasive plants are hybrids between the two. It is
believed that the hybrids were propagated and spread through the nursery trade. Both spe-
cies and several cultivars, with white, purple, pink, or lavender flowers, continue to be sold
as popular ornamentals.
646 n VINES

Habitat. Exotic wisterias are

common in right-of-ways, road-
sides, forest edges, ditches, and
in riparian zones. Plants also
invade open or disturbed areas
after burns or clearing. Al-
though they can be found in
part shade, plants grow best in
full sun. They tolerate a variety
of soils and moisture, but prefer
deep loamy soils that are well
drained.
Reproduction and Dispersal.
Reproduction is primarily veg-
etative. Nodes on stolons,
which creep along the ground
surface, sprout new plants.
Plants spread locally from
homesites into surrounding
woodlands, from neglected gar-
dens or due to improper dis-
posal of garden waste. Seeds
are produced under favorable
conditions, and can be carried
long distances downstream by
water to new riparian sites.
Long-distance dispersal is accom-
plished through Internet and
nursery sales. The reproductive
biology of the hybrids, how seeds
form and disperse to new loca-
Both alien wisteria species, as well as their hybrids, are invasive, especially tions, is not yet known.
in the southern and eastern states. (Introduced range adapted from USDA Impacts. Aggressive wisteria
PLANTS Database, Invasive Plant Atlas of the United States, and selected plants are hardy and can live
references.) for 50 or more years. New
vines, either from seeds or from
root sprouts, that trail on the ground can form thickets that exclude all other plants. Stems
climb onto and over native shrubs and trees, either shading them out or strangling them.
The twining vines circle the host tree tightly, cutting through the bark and girdling and kill-
ing them. When trees die, the forest canopy is altered, creating sunny openings that may ini-
tially encourage growth of native species, but ultimately favor the expansion of wisteria.
Seeds and pods of Chinese wisteria are toxic, causing nausea, vomiting, stomach pain, or
diarrhea if eaten.
Management. Because wisteria will invade disturbed areas, it is important to maintain
healthy ecosystems. Physical control is practical on small plants or small infestations.
Small plants can be dug up in sensitive sites where herbicides are prohibited. Vines with
seed pods should be bagged and disposed of properly, but piles of pulled material can
remain on site as long as the area is monitored frequently for regrowth. Vines that grow into
 WISTERIA n 647

A. Vines climb by twining around other plants, girdling and killing them. (Chris Evans, River to River CWMA,
Bugwood.org.) B. Long compound leaves have many leaflets. (James H. Miller, USDA Forest Service,
Bugwood.org.) C. Pendulant clusters of showy flowers may cover the plant. (Chris Evans, River to River CWMA,
Bugwood.org). D. Fuzzy seed pods each contain 2–6 seeds. (Ted Bodner, Southern Weed Science Society,
Bugwood.org.)

trees can be severed at the base, close to the root crown. Cut resprouts as they occur, every
two weeks during the growing season. Cutting vines repeatedly to ground level slows
growth, prevents seed production, and decreases nutrient reserves.
Chemical control is effective, either sprayed on foliage or applied to basal bark or cut
stems. Foliar sprays work best when applied to resprouts after vines have been cut.
Spraying is efficient for large stands that cover all other vegetation. Foliar application is also
best where cutting the stumps or physical removal is too disruptive. Applications of herbi-
cide to cut stumps are appropriate for large stands where thick vines are well established.
If applied with care, systemics such as glyphosate or triclopyr can be used without harming
nontarget plants. In order to ensure protection of native species, spraying should be done
before or after the spring bloom of wildflowers, either early in spring or in the fall.
Because triclopyr is selective to broadleaf plants, it is safe to apply around native grasses.
Chlopyralid is a herbicide specifically targeted to the sunflower, buckwheat, and pea fami-
lies, but it seeps into groundwater. Picloram can be used where no desirable vegetation
remains.
No biological control for wisteria is known.

Selected References
“Exotic Wisterias.” Weed of the Week. USDA Forest Service, Invasive Plants, 2006. http://na.fs.fed.us/
fhp/invasive_plants.
Swearingen, J., and Tom Remaley. “Chinese Wisteria.” Weeds Gone Wild: Alien Plant Invaders of
Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2010. http://www.nps
.gov/plants/alien/fact/pdf/wisi1.pdf.
Swearingen, J., and Tom Remaley. “Japanese Wisteria.” Weeds Gone Wild: Alien Plant Invaders of
Natural Areas. Plant Conservation Alliance’s Alien Plant Working Group, 2010. http://www.nps
.gov/plants/alien/fact/pdf/wifl1.pdf.
Trusty, J. L., B. G. Lockaby, W. C. Zipperer, and L. R. Goertzen. “Identity of Naturalised Exotic Wisteria
(Fabaceae) in the South-Eastern United States.” Weed Research 47: 479–87, 2007. http://
www.srs.fs.usda.gov/pubs/ja/ja_trusty003.pdf.
This page intentionally left blank
Tables and Lists about
Invasive Plants

n Common Names and Scientific Names

All invasive plant species profiled in the 80 entries of Volume 2 show their common and
scientific names in the entry headings for these plants. Below are common and scientific
names of other species discussed within the entries of Volume 2.

Common Name Scientific Name

Aquatic
Anchored waterhyacinth Eichhornia azurea
Brazilian waterhyacinth Eichhornia paniculata
Brazilian waterweed, Brazilian elodea Egeria densa
Canadian waterweed Elodea canadensis
Common salvinia, water spangles Salvinia minima
Coon’s tail, common hortwort Ceratophyllum demersum
Devil pod, bat nut, horn nut Trapa bicornis
Duckweed Lemna spp.
Northern watermilfoil, shortspike Myriophyllum sibiricum
watermilfoil
Mexican waterfern Azolla mexicana
Mosquito fern Azolla caroliniana
Pacific mosquito fern Azolla filiculoides
Parrotfeather Myriophyllum aquaticum
Pickerelweed Pontederia cordata
Pondweed Potamogeton spp.
Tapegrass Vallisneria americana
Variable leaf waterhyacinth Eichhornia diversifolia
Variable watermilfoil Myriophyllum heterophyllum
(Continued )
650 n COMMON NAMES AND SCIENTIFIC NAMES

Common Name Scientific Name

Water clover Marsilea vestilia

Forbs
Arrowhead Sagittaria spp.
Bagpod, bladderpod Sesbania vesicaria
Barbwire Russian thistle Salsola paulsenii
Beach layia Layia carnosa
Blazing star Liatris spp.
Bloodroot Sanguinaria canadensis
Bohemian knotweed Polygonum × bohemicum (P. cuspidatum ×
P. sachalinense)
Branched tearthumb Polygonum meisneranum
Broomleaf toadflax Linaria genistifolia
Buffalobur nightshade Solanum rostratum
Bull thistle Cirsium vulgare
Burning bush, kochia, summer-cypress Bassia scoparia
Canada germander Teucrium canadense
Canada toadflax, blue toadflax Nuttallanthus canadensis
Caper spurge, gopher plant Euphorbia lathyris
Carolina horsenettle Solanum carolinense
Celandine Chelidonium majus
Celandine poppy Stylophorum diphyllum
Clasping pepperweed Lepidium perfoliatum
Coastal sand spurge Euphorbia exserta
Common parsnip, cow parsnip Heracleum maximum
Crinkleroot, twoleaf toothwort Cardamine diphylla
Cutleaf toothwort Cardamine laciniata
Cypress spurge Euphorbia cyparissias
Diffuse knapweed Centaurea diffusa
Dutchman’s breeches Dicentra cucullaria
Dwarf St. Johnswort Hypericum mutilum
Early saxifrage Saxifraga virginica
Eggleaf spurge Euphorbia oblongata
 COMMON NAMES AND SCIENTIFIC NAMES n 651

Common Name Scientific Name

Fireweed Epilobium angustifolium

Fringecup Tellima grandiflora
Geraldton carnation weed Euphorbia terracina
Giant knotweed Polygonum sachalinense
Giant wild pine, giant air plant Tillandsia utriculata
Globe artichoke Cynara scolymus
Green comet milkweed Asclepias viridiflora
Ground ivy Glechoma hederaceae
Hairy vetch Vicia villosa
Hairy whitetop Cardaria pubescens
Harbinger-of-spring Erigenia bulbosa
Hart’s tongue fern Asplenium scolopendrium
Heartleaf horsenettle Solanum cardiophyllum
Hepatica Hepatica nobilis
Horsetail Equisetum spp.
Humboldt Bay owl’s clover Castilleja ambigua ssp. humboldtiensis
Iberian knapweed, Iberian starthistle Centaurea iberica
Italian thistle Carduus pycnocephalus
Jesup’s milkvetch Astralagus robbinsii var. Jesupi
Large St. Johnswort Hypericum majus
Lens-pod hoary cress Cardaria chalepensis
Madwoman’s milk, sun spurge Euphorbia helioscopia
Malta starthistle Centaurea melitensis
Marsh jaumea Jaumea carnosa
Marsh marigold Caltha palustris
Milk thistle Silybum marianum
Moth mullein Verbascum blattaria
Oppositeleaf Russian thistle Salsola soda
Partridge pea Chamaecrista fasciculata
Pickleweed Salicornia virginica
Piggyback plant Talmiea menziesii
Pink sand verbena Abronia umbellata ssp. breviflora
(Continued )
652 n COMMON NAMES AND SCIENTIFIC NAMES

Common Name Scientific Name

Plumeless thistle Carduus acanthoides

Pt. Reyes bird’s beak Cordylanthus maritimus var. palustris
Poison hemlock Conium maculatum
Prostrate ticktrefoil, dollar leaf plant Desmodium rotundifolium
Purple starthistle, red starthistle Centaurea calcitrapa
Purplestem angelica Angelica atropurpura
Queen Anne’s lace Daucus carota
Red clover Trifolium pretense
Rio Grande ragweed, South Texas Ambrosia cheiranthifolia
ambrosia
Russian knapweed Acroptilon repens
Sacramento Mountain thistle Cirsium vinaceum
Safflower Carthamus tinctorius
Sand dune thistle Cirsium pitcheri
Scotch thistle Onopordum acanthium
Seabeach amaranth Amaranthus pumilus
Seabeach evening primrose Oenothera humifusa
Seacoast marsh elder Iva imbricata
Sea fig Carpobrotus chilensis
Searocket Cakile edentula
Sea sandwort Honckenya peploides
Seaside goldenrod Solidago sempervirens
Seaside knotweed Polygonum glaucum
Sicilian starthistle Centaurea sulphurea
Slender-flowered thistle Carduus tenuflorus
Slender lespedeza Lespedeza virginica
Slender purslane Sesuvium maritimum
Slender Russian thistle Salsola collina
Spring beauty Claytonia virginica
Squarrose knapweed Centaurea virgata ssp. squarrosa
Squirrel-corn Dicentra canadensis
Swamp verbena Verbena hastata
 COMMON NAMES AND SCIENTIFIC NAMES n 653

Common Name Scientific Name

Sweet cicily Osmorhiza claytonii

Tinker’s penny Hypericum anagalloides
Toothed spurge Euphorbia serrata
Toothworts Cardamine spp.
Tree ferns Cibotium spp.
Trillium Trillium spp.
Tropical curlygrass fern, ray fern Actinostachys pennula
Trout lily Erythronium spp.
Twinleaf Jeffersonia diphylla
Violet Viola spp.
Virginia bluebells Mertensia virginica
Wavyleaf thistle Cirsium undulatum
Western horsenettle, Torrey’s nightshade Solanum dimidiatum
White avens Geum canadense
White ginger Hedychium coronarium
Wild ginger Aserium canadense
Wild parsnip Pastinaca sativa
Wolf’s primrose Oenothera wolfii
Yellow ginger Hedychium flavescens
Yellowspine thistle Cirsium ochrocentrum
Zapata bladderpod Lesquerella thamnophila
Graminoids
African feathergrass Pennisetum macrourum
Alpine hairgrass Deschampsia nubigena
American beach grass Ammophila breviligulata
American cupscale Sacciolepis striata
Arizona cottontop Digitaria californica
Arizona wheatgrass Elymus arizonicus
Bahia grass Paspalum notatum
Bald brome Bromus racemosus
Barley Hordeum spp.
Beach panic grass Panicum amarum
(Continued )
654 n COMMON NAMES AND SCIENTIFIC NAMES
Common Name
Beachstar
Beardless wheatgrass
Bermuda grass
Blue bunchgrass
Bluebunch wheatgrass
Brazilian satintail
Broadleaf cattail
Bulb panicgrass
Bulrush
Bush muhly
California cordgrass
California satintail
Cattail
Coast cockspur
Columbia needlegrass
Cosmopolitan bulrush
Crested wheatgrass
Dune panic grass
Elephant grass
Feathertop
Field brome, Japanese brome
Foxtail
Galleta grass
Hairy wheatgrass
Indian ricegrass
Intermediate wheatgrass
Japanese blood grass
Japanese brome
Jointed grass
Kyasuma grass
Large-headed sedge
Maidencane
Scientific Name
Remirea maritima
Pseudoroegneria spicata
Cynodon dactylon
Pseudoroegneria spicata
Pseudoroegneria spicata
Imperata brasiliensis
Typha latifolia
Panicum bulbosum
Scirpus spp.
Muhlenbergia porteri
Spartina foliosa
Imperata brevifolia
Typha spp.
Echinochloa walteri
Achnatherum nelsonii
Schoenoplectus maritimus
Agropyron cristatum
Panicum amarulum
Pennisetum purpureum
Pennisetum villosum
Bromus arvensus
Alopecuris spp.
Pleuraphis spp.
Thinopyrum intermedium
Achnatherum hymenoides
Thinopyrum intermedium
Imperata cylindrica ‘Rubra’
Bromus japonicus
Arthraxon hispidus
Pennisetum pedicellatum
Carex macrocephala
Panicum hemitomon
 COMMON NAMES AND SCIENTIFIC NAMES n 655

Common Name Scientific Name

Missiongrass, feathery pennisetum Pennisetum polystachyon

Needle and thread grass Hesperostipa comata
Needlegrass Stipa spp.
Northern reedgrass Calamagrostis stricta
Northern wheatgrass Elymus lanceolatus
Pearl millet Pennisetum glaucum
Pili grass, tanglehead Heteropogon contortus
Plains bristlegrass Setaria macrostachya
Reedgrass Calamagrostis stricta
Rush wheatgrass Thinopyrum ponticum
Russian wheatgrass Thinopyrum junceiforme
Ryebrome Bromus secalinus
Ryegrass Lolium multiflorum
St. Augustine grass Stenotaphrum secundatum
Saltgrass Distichlis spicata
Sandberg bluegrass Poa secunda
Sawgrass Cladium jamaicense
Sea oats Uniola paniculata
Seaside arrowgrass Triglochin maritima
Sedge Carex spp., Cyperus spp.
Silverberry Elaeagnus commutata
Sheep fescue Festuca ovina
Small cordgrass Spartina maritima
Smooth brome Bromus inermis
Soft brome Bromus hordeaceus
Sorghum, shattercane Sorghum bicolor
Squirreltail Elymus elymoides
Sudangrass Sorghum bicolor spp. drummondii
Tall wheatgrass Thinopyrum elongatum
Thurber’s needlegrass Achnatherum thurberianum
Tick quackgrass Thinopyrum pungens
Toe toe, Richard’s pampas grass Cortaderia richardii
(Continued )
656 n COMMON NAMES AND SCIENTIFIC NAMES

Common Name Scientific Name

Virginia cutgrass Leersia virginica

Western wheatgrass Agropyron smithii
Wheatgrass Agropyron spp.
White cattail Typha glauca
Wild rye Elymus spp.
Shrubs
Akalakala Rubus macraei
Allegheny blackberry Rubus allegheniensis
American barberry Berberis canadensis
Antelope bitterbrush Purshia tridentata
Bearberry honeysuckle, twinberry Lonicera involucrata
honeysuckle
Big sagebrush Artemisia tridentata
Blueberry Vaccinium spp.
Bridal broom Retama monosperma
Bristletips Oxyspora paniculata
Buffaloberry Shepherdia argentia
Burning bush Euonymus atropurpurea
Bursage Ambrosia spp.
Bursting heart Euonymus americanus
Canadian honeysuckle Lonicera canadensis
Canary Island St. Johnswort Hypericum canariense
Canby’s mountain-lover Paxistima canbyi
Cane ti, cane tibouchina Tibouchina herbacea
Cherokee rose Rosa laevigata
Climbing prairie rose Rosa setigera
Cockroach berry Solanum capsicoides
Coffee colubrina Colubrina arborescens
Common barberry Berberis vulgaris
Common elderberry Sambucus canadensis
Coralberry Symphoricarpos orbiculatus
Creosote bush Larrea tridentata
 COMMON NAMES AND SCIENTIFIC NAMES n 657

Common Name Scientific Name

Crimson bottlebrush Callistemon citrinus

Cuban nakedwood Colubrina cubensis
Dog rose Rosa canina
Drummond rattlebox Sesbania drummondii
Dwarf gorse Ulex minor
European fly honeysuckle, dwarf Lonicera xylosteum
honeysuckle
European wand loosestrife Lythrum virgatum
Georgia bully Sideoxylon thornei
Glorybush, princess flower Tibouchina urvilleana
Grape honeysuckle Lonicera reticulata
Greasewood Sarcobatus vermiculatus
Hawaiian blackberry, akala Rubus hawaiiensis
Himalayan blackberry Rubus armeniacus
Maccartney rose Rosa bracteata
Pasture rose Rosa carolina
Pineland verbena, depressed Lantana depressa
shrubverbena
Prickly rose Rosa acicularis
Privet Ligustrum spp.
Purple African nightshade Solanum marginatum
Purpleflowering raspberry Rubus odoratus
Red honeysuckle Lonicera dioica
Rugosa rose Rosa rugosa
Running strawberry bush Euonymus obovatus
Rusty blackhaw Viburnum rufidulum
Saltbush Atriplex spp.
Sawtooth blackberry Rubus argutus
Shadscale saltbush Atriplex confertifolia
Shrubby nightshade Solanum robustum
Shrubby Russian thistle Salsola vermiculata
Silverleaf nightshade Solanum elaeagnifolium
Silversword Argyroxiphium sandwicense ssp. sandwicense
(Continued )
658 n COMMON NAMES AND SCIENTIFIC NAMES

Common Name Scientific Name

Silverthorn Elaeagnus pungens

Smooth rose Rosa blanda
Snowpeaks raspberry Rubus niveus
Soldierwood Colubrina elliptica
Southern bush honeysuckle Diervilla sessilifolia
Staghorn sumac Rhus typhina
Swamp fly honeysuckle Lonicera oblongifolia
Swamp rose Rosa palustris
Sweetberry honeysuckle Lonicera caerulea
Sweetbriar rose Rosa eglanteria
Turkey berry Solanum torvum
Virginia rose Rosa virginiana
West Indian raspberry Rubus rosifolius
Winter honeysuckle, fragrant Lonicera fragrantissima
honeysuckle
Woods’ rose Rosa woodsii
Wormwood Artemisia spp.
Yellow honeysuckle Lonicera flava
Trees
Alder Alnus spp.
Ash Fraxinus spp.
Aspen Populus spp.
Athel tamarisk Tamarisk aphylla
Australian river oak Casuarina cunninghamiana
Autumn olive Elaeagnus umbellata
Bald cypress Taxodium distichum
Bay cedar Suriana maritima
Beech Fagus spp.
Birch Betula spp.
Brazilian oak Casuarina glauca
Buttonwood Conocarpus erectus
Columnar cactus Cereus spp.
 COMMON NAMES AND SCIENTIFIC NAMES n 659

Common Name Scientific Name

California buckeye Aesculus californica

California live oak Quercus agrifolia
Catalpa Catalpa speciosa
Common guava Psidium guajava
Cottonwood Populus spp.
Crape myrtle Lagerstroemia indica
Elm Ulmus spp.
Elongata princess tree Paulownia elongata
Florida thatch palm Thrinax radiata
Honeylocust Gleditsia triacanthos
Koa Acacia koa
Linden Tilia spp.
Live oak Quercus virginiana
Machineel Hippomane mancinella
Mamane Sophora chrysophylla
Maple Acer spp.
Mesquite Prosopis spp.
Oak Quercus spp.
Ohia Metrosideros polymorpha
Peruvian peppertree Schinus molle
Purple osier willow Salix purpurea
Red mulberry Morus rubra
Sassafras Sassafras albidum
Spruce Picea spp.
Surinam cherry Eugenia uniflora
Texas umbrella chinaberry Melia azedarach cv ‘Umbraculiformis’
Wax myrtle Myrica cerifera
West Indian mahogany Swietenia mahagoni
White basswood Tilia heterophylla
White-flowered paulownia Paulownia fortunei
White mulberry Morus alba
Wild cinnamon Canella winterana
(Continued )
660 n COMMON NAMES AND SCIENTIFIC NAMES

Common Name Scientific Name

Willow Salix spp.

Woman’s tongue Albizia lebbeck
Vines
Alfalfa dodder Cuscuta approximata
Amberique bean, American trailing wild Strophostyles helvula
bean
American bittersweet Celastrus scandens
American climbing fern Lygodium palmatum
American hogpeanut Amphicarpaea bracteata
American wisteria, Atlantic wisteria Wisteria frutescens
Atlantic Ivy, Irish ivy Hedera hibernica
Beach clustervine Jacquemontia reclinata
Black pepper Piper nigrum
Boston ivy Parthenocissus tricuspidata
Bur cucumber Sicyos angulatus
Chinese honeysuckle Lonicera japonica var. chinensis
Chinese yam, air potato Dioscorea bulbifera
Cinammon vine, Chinese yam Dioscorea oppositifolia
Climbing nightshade Solanum dulcamara
Colchis ivy, Persian ivy Hedera colchica
Common dodder, scaldweed, swamp Cuscuta gronovii
dodder
Common hops Humulus lupulus
Coral honeysuckle Lonicera sempervirens
Dutchman’s pipe Aristolochia macrophylla
Grape Vitis spp.
Hall’s honeysuckle Lonicera japonica var. halliana
Heartleaf peppervine Ampelopsis cordata
Hedge false bindweed Calystegia sepium
Honeyvine Cyanchum laeve
Hollyhock bindweed Convolvulvus althaeoides
Hybrid akebia Akebia x pentaphylla
Littleleaf sensitive-briar Mimosa microphylla
 COMMON NAMES AND SCIENTIFIC NAMES n 661

Common Name Scientific Name

Mallow bindweed Convolvulus althaeoides

Morning glories Ipomoea spp.
Orangeberry nightshade Solanum lanceolatum
Peppervine Ampelopsis arborea
Pipevine Aristolochia macrophylla
Poison ivy Toxicodendron radicans
Three-leaf akebia Akebia trifoliata
Trumpet creeper Campsis radicans
Virginia creeper Parthenocissus quinquefolia
Western morning glory Calystegia occidentalis
Wetlands nightshade Solanum tampicense
White swallow-wort Cynanchum vincetoxicum
Wild cucumber Echinocystis lobata
Wild grape Vitis spp.
Woolly Dutchman’s pipe Aristolochia tomentosa
Epiphytes
Bromeliads Tillandsia spp.
Orchids Encyclia bootheriana, Oncidum luridum
Birds
American Robin Turdus migratorius
Black-capped Chickadee Poecile atricapillus
Black Skimmer Rynchops niger
Blue Jay Cyanocitta cristata
Bobwhite Quail Colinus virginianus
California Clapper rail Rallus longirostris obsoletus
Cedar Waxwing Bombycilla cedrorum
Chinese Turtledove Streptopelia chinensis
Chukar Partridge Alectoris chukar
Common Myna Acridotheros tristis
Everglades Snail Kite Rostrhamus sociabilis
Fish Crow Corvus ossifragus
Gambel’s Quail Lophortyx gambelii
(Continued )
662 n COMMON NAMES AND SCIENTIFIC NAMES

Common Name Scientific Name

Golden Eagle Aquila chrysaetos

Grasshopper Sparrow Ammodramus savannarum
Greater Prairie Chicken Tympanuchus cupido
Hawaiian Coot Fulica alai
Hawaiian Gallinule Gallinule chloropus
Hawaiian Stilt Himantopus mexicanus knudseni
Japanese White-eye Zosterops japonica
Least Bell’s Vireo Vireo bellii pusillus
Least Tern Sterna antillarum
Meadowlark Stumella spp.
Mockingbird Mimus polyglottos
Palila Loxioides bailleui
Pheasant Phasianus colchicus
Piping Plover Charadrius melodus
Red-billed Leiothrix Leiothrix lutea
Red-winged Blackbird Agelaius phoeniceus
Ruffed Grouse Bonasa umbellus
Scaled Quail Callipepla squamata
Southwestern Willow Flycatcher Empidonax traillii extimus
Starling Sturnus vulgaris
Wild Turkey Meleagris gallopavo
Wood Stork Mycteria americana
Mammals
Cotton mouse Peromyscus gossypinus
Cotton rats Sigmodon spp.
Coyote Canis latrans
Deer mice Peromyscus maniculatus
Elk Cervus canadensis
Ground squirrels Citellus spp.
Feral pig Sus scrofa
Hispid cotton rat Sigmodon hispidus
Marsh rice rat Oryzomys palustris
 COMMON NAMES AND SCIENTIFIC NAMES n 663

Common Name Scientific Name

Mongoose Herpestes auropunctatus

Mule deer Odocoileus hemionus
Salt marsh harvest mouse Reithrodontomys raviventris
White-tailed deer Odocoileus virginianus
Woodrats Neotoma spp.
Fish
Chain pickerel Esox niger
Chum salmon Oncorhynchus keta
English sole Pleuronectes vetulus
Grass carp Ctenopharyngodon idella
Reptiles and Amphibians
American alligator Alligator mississippiensis
American crocodile Crocodylus acutus
Gopher snake Pituophis spp.
Gopher tortoise Gopherus polyphemus
Green sea turtle Chelonia mydas
Green tree frog Hyla cinerea
Indigo snake Drymarchon corais couperi
Leopard frog Rana spp.
Loggerhead turtle Caretta caretta ssp. caretta
Pig frog Rana grylio
Invertebraes
Caribbean fruit fly Anastrepha suspensa
Colorado potato beetle Leptinotarsa decemlineata
Garden white butterflies Pieris spp.
Green peach aphid Myzus persicae
Gypsy moth Lymantria dispar
Honey bee Apis mellifera
Monarch butterfly Danaus plexippus
Mullein leaf bug Campylomna verbasci
Northeastern sea beach tiger beetle Cicindela dorsalis dorsalis
Pearly eye butterfly Enodia anthedon
(Continued )
664 n COMMON NAMES AND SCIENTIFIC NAMES

Common Name Scientific Name

Silverleaf whitefly Bemisia argentifolii

Soybean loper Pseudoplusia includens
Sugar beet leafhopper Circulifer tenellus
Tobacco budworm Helicoverpa virescens
Tobacco hornworm Manduca sexta
Tomato hornworm Manduca quinquemaculata
Tomato pinworm Keiferia lycopersicella
West Virginia white butterfly Pieris virginiensis
Crustaceans
Dungeness crab Cancer magister
Oyster Crassotrea gigas
n State-by-State Designations of Invasive
or Noxious Weeds

The following states have officially declared certain plants (discussed or mentioned in this
encyclopedia) to be noxious, invasive, or undesirable on some level. Lack of an entry for a
state, however, should not be interpreted as lack of a problem. Some plants may be noxious,
invasive, or undesirable even without official recognition. States have differing levels of
organization for identifying noxious or unwanted plants.

Alabama

Aquatic: Anchored waterhyacinth, Eurasian watermilfoil, giant salvinia, hydrilla, parrotfeather,

water chestnut, waterhyacinth
Forbs: Garlic mustard, giant hogweed, Japanese knotweed, purple loosestrife
Graminoids: African feathergrass, Brazilian satintail, cogongrass, common reed, Japanese stilt
grass, kikuyugrass, kyasuma grass, mission grass
Shrubs: Multiflora rose, shrubby Russian thistle, tropical soda apple, turkey berry.
Trees: Melaleuca
Vines: Dodders, Japanese climbing fern, Old World climbing fern, mile-a-minute, wetlands
nightshade

Alaska

Forbs: Canada thistle, Carolina horsenettle, hairy whitetop, leafy spurge, perennial pepperweed,
Russian knapweed
Graminoids: Quackgrass
Vines: Field bindweed

Arizona

Aquatic: Anchored waterhyacinth, giant salvinia, hydrilla, water chestnut, waterhyacinth

Forbs: Canada thistle, Carolina horsenettle, Dalmatian toadflax, diffuse knapweed, dyer’s woad,
hairy whitetop, halogeton, Iberian knapweed, leafy spurge, lens-pod cress, plumeless thistle,
purple loosestrife, purple starthistle, Russian knapweed, Scotch thistle, Sicilian starthistle,
spotted knapweed, squarrose knapweed, yellow starthistle
Graminoids: Buffelgrass, quackgrass
Shrubs: Tropical soda apple
Vines: Dodders, field bindweed

Arkansas

Forbs: Barbwire Russian thistle, bull thistle, Carduus spp. thistles, Carolina horsenettle, Cirsium
spp. thistles, hemp sesbania, Italian thistle, oppositeleaf Russian thistle, plumeless thistle,
prickly Russian thistle, purple loosestrife, Scotch thistle, slender Russian thistle
Graminoids: Bald brome, ryebrome
Shrubs: Shrubby Russian thistle, silverleaf nightshade
Vines: Dodders, field bindweed

(Continued )
666 n STATE-BY-STATE DESIGNATIONS OF INVASIVE OR NOXIOUS WEEDS

California

Aquatic: Anchored waterhyacinth, giant salvinia, hydrilla, water chestnut, waterhyacinth

Forbs: Barbwire Russian thistle, Canada thistle, Carolina horsenettle, common St. Johnswort,
Dalmatian toadflax, diffuse knapweed, dyer’s woad, giant hogweed, giant knotweed, hairy
whitetop, halogeton, Iberian knapweed, Italian thistle, Japanese knotweed, leafy spurge,
lens-pod cress, musk thistle, perennial pepperweed, plumeless thistle, prickly Russian thistle,
purple loosestrife, purple starthistle, Russian knapweed, Scotch thistle, Sicilian starthistle, slen-
der Russian thistle, spotted knapweed, squarrose knapweed, western horsenettle, yellow
starthistle
Graminoids: African feathergrass, Brazilian satintail, California satintail, cogongrass,
kikuyugrass, kyasuma grass, medusahead, mission grass, quackgrass
Shrubs: French broom, gorse, Scotch broom, shrubby Russian thistle, silverleaf nightshade,
tropical soda apple, turkey berry
Trees: Melaleuca
Vines: Dodders, field bindweed, wetlands nightshade

Colorado

Aquatic: Eurasian watermilfoil, giant salvinia, hydrilla

Forbs: Bull thistle, Canada thistle, Chinese lespedeza, common mullein, common St. Johnswort,
Dalmatian toadflax, diffuse knapweed, dyer’s woad, halogeton, leafy spurge, moth mullein,
musk thistle, perennial pepperweed, plumeless thistle, purple loosestrife, Russian knapweed,
Scotch thistle, spotted knapweed, squarrose knapweed, yellow starthistle
Graminoids: Cheatgrass, medusahead, quackgrass
Trees: Russian olive, tamarisk
Vines: Field bindweed

Connecticut

Aquatic: Eurasian watermilfoil, giant salvinia, hydrilla, parrotfeather, variable watermilfoil, water
chestnut, waterhyacinth
Forbs: Burning bush, Canada thistle, fig buttercup, garlic mustard, giant hogweed, giant
knotweed, goutweed, Japanese knotweed, leafy spurge, perennial pepperweed, purple
loosestrife, Scotch thistle, spotted knapweed
Graminoids: Asiatic sand sedge, cheatgrass, common reed, Japanese stilt grass
Shrubs: Amur honeysuckle, Bell’s honeysuckle, common barberry, Japanese barberry, Morrow’s
honeysuckle, multiflora rose, Tatarian honeysuckle
Trees: Autumn olive, princess tree, Russian olive, tree of heaven
Vines: Black swallow-wort, Japanese honeysuckle, Japanese hops, kudzu, mile-a-minute,
Oriental bittersweet, pale swallow-wort, porcelainberry

Delaware

Forbs: Canada thistle

Florida

Aquatic: Anchored waterhyacinth, common salvinia, Eurasian watermilfoil, hydrilla, water

chestnut, waterhyacinth
Forbs: Giant hogweed, purple loosestrife
Graminoids: African feathergrass, Brazilian satintail, cogongrass, kikuyugrass, kyasuma grass,
mission grass
Shrubs: Asiatic colubrina, shrubby Russian thistle, tropical soda apple, turkey berry
Trees: Australian pine, Brazilian peppertree, carrotwood, fire tree, melaleuca
 STATE-BY-STATE DESIGNATIONS OF INVASIVE OR NOXIOUS WEEDS n 667

Vines: Dodders, field bindweed, Japanese climbing fern, kudzu, Old World climbing fern,
wetlands nightshade

Hawai’i

Forbs: Canada thistle, Carolina horsenettle, common mullein, hairy whitetop, halogeton, leafy
spurge, perennial pepperweed, prickly Russian thistle, Russian knapweed
Graminoids: Cogongrass, crimson fountain grass, jubata grass, quackgrass
Shrubs: French broom, Gorse, Koster’s curse, sawtooth blackberry, Scotch broom, silverleaf
nightshade, snowpeaks raspberry, Spanish broom, turkey berry, yellow Himalayan raspberry
Trees: Fire tree, velvet tree
Vines: Field bindweed

Idaho

Aquatic: Eurasian watermilfoil

Forbs: Buffalobur nightshade, Canada thistle, Dalmatian toadflax, diffuse knapweed, dyer’s
woad, leafy spurge, musk thistle, perennial pepperweed, plumeless thistle, purple loosestrife,
Russian knapweed, Scotch thistle, spotted knapweed, yellow starthistle, yellow toadflax
Shrubs: Scotch broom, silverleaf nightshade
Vines: Field bindweed

Illinois

Forbs: Canada thistle, musk thistle

Vines: Kudzu

Indiana

Forbs: Canada thistle, purple loosestrife

Graminoids: Sorghum, Sudangrass
Shrubs: Multiflora rose

Iowa

Forbs: Bull thistle, Canada thistle, Carduus spp., Carolina horsenettle, Italian thistle, leafy
spurge, purple loosestrife, Russian knapweed
Graminoids: Quackgrass, sorghum, Sudangrass
Shrubs: Multiflora rose
Vines: Field bindweed

Kansas

Forbs: Canada thistle, Chinese lespedeza, leafy spurge, musk thistle, Russian knapweed
Graminoids: Quackgrass
Vines: Field bindweed, kudzu

Kentucky

Forbs: Canada thistle, musk thistle

Shrubs: Multiflora rose
Vines: Kudzu

Maine

Aquatics: Eurasian watermilfoil, hydrilla, parrot feather, variable watermilfoil, water chestnut

(Continued )
668 n STATE-BY-STATE DESIGNATIONS OF INVASIVE OR NOXIOUS WEEDS

Maryland

Forbs: Bull thistle, Canada thistle, musk thistle, plumeless thistle

Graminoids: Sorghum, Sudangrass

Massachusetts

Aquatic: Anchored waterhyacinth, Eurasian watermilfoil, giant salvinia, hydrilla, parrotfeather,

variable watermilfoil, water chestnut
Forbs: garlic mustard, giant hogweed, goutweed, Japanese knotweed, leafy spurge, perennial
pepperweed, purple loosestrife, spotted knapweed
Graminoids: African feathergrass, Asiatic sand sedge, Brazilian satintail, common reed, Japanese
stilt grass, kikuyugrass, kyasuma grass, mission grass
Shrubs: Amur honeysuckle, Bell’s honeysuckle, common barberry, Japanese barberry, Morrow’s
honeysuckle, multiflora rose, shrubby Russian thistle, Tatarian honeysuckle, tropical soda
apple, turkey berry
Trees: Autumn olive, melaleuca, tree of heaven
Vines: Black swallow-wort, dodders, Japanese honeysuckle, Japanese hops, kudzu, mile-a-
minute, Oriental bittersweet, pale swallow-wort, porcelainberry, wetlands nightshade

Michigan

Forbs: Canada thistle, purple loosestrife

Vines: Dodders, field bindweed

Minnesota

Forbs: Bull thistle, Canada thistle, garlic mustard, giant hogweed, leafy spurge, musk thistle,
plumeless thistle, purple loosestrife
Graminoids: African feathergrass, Brazilian satintail, cogongrass, kikuyugrass, kyasuma grass,
mission grass
Shrubs: Shrubby Russian thistle, tropical soda apple, turkey berry
Vines: Dodders, field bindweed

Mississippi

Aquatic: Giant salvinia, hydrilla

Forbs: Fig buttercup
Graminoids: Brazilian satintail, cogongrass
Shrubs: Tropical soda apple
Vines: Kudzu

Missouri

Forbs: Canada thistle, musk thistle, purple loosestrife, Scotch thistle

Shrubs: Multiflora rose
Vines: Field bindweed, kudzu

Montana

Aquatic: Eurasian watermilfoil

Forbs: Canada thistle, common St. Johnswort, Dalmatian toadflax, diffuse knapweed, dyer’s
woad, leafy spurge, perennial pepperweed, purple loosestrife, Russian knapweed, spotted
knapweed, yellow starthistle, yellow toadflax
Trees: Tamarisk
Vines: Field bindweed
 STATE-BY-STATE DESIGNATIONS OF INVASIVE OR NOXIOUS WEEDS n 669

Nebraska

Forbs: Canada thistle, diffuse knapweed, leafy spurge, musk thistle, plumeless thistle, purple
loosestrife, spotted knapweed
Trees: Tamarisk

Nevada

Aquatic: Eurasian watermilfoil, giant salvinia, hydrilla

Forbs: Canada thistle, Carolina horsenettle, common St. Johnswort, Dalmatian toadflax, diffuse
knapweed, dyer’s woad, Iberian knapweed, leafy spurge, Malta starthistle, musk thistle,
perennial pepperweed, purple loosestrife, purple starthistle, Russian knapweed, Scotch
thistle, spotted knapweed, Sudangrass
Shrubs: Silverleaf nightshade
Trees: Tamarisk
Vines: Black swallow-wort, field bindweed, pale swallow-wort

New Hampshire

Forbs: Garlic mustard, giant hogweed, Japanese knotweed spotted knapweed

Shrubs: Bell’s honeysuckle, common barberry, Morrow’s honeysuckle, multiflora rose, Tatarian
honeysuckle
Trees: Autumn olive, tree of heaven
Vines: Black swallow-wort, Japanese honeysuckle, Oriental bittersweet, pale swallow-wort

New Mexico

Aquatic: Eurasian watermilfoil, hydrilla

Forbs: Bull thistle, Canada thistle, Dalmatian toadflax, diffuse knapweed, dyer’s woad,
halogeton, leafy spurge, Malta starthistle, musk thistle, perennial pepperweed, purple
loosestrife, purple starthistle, Russian knapweed, Scotch thistle, spotted knapweed, yellow
starthistle, yellow toadflax
Trees: Russian olive, tamarisk
Vines: Field bindweed

North Carolina

Aquatic: Anchored waterhyacinth, common salvinia, Eurasian watermilfoil, hydrilla, water

chestnut
Forbs: Canada thistle, giant hogweed, musk thistle, plumeless thistle, purple loosestrife
Graminoids: African feathergrass, Brazilian satintail, cogongrass, kikuyugrass, kyasuma grass,
mission grass
Shrubs: Shrubby Russian thistle, tropical soda apple, turkey berry
Trees: Melaleuca
Vines: Dodders, mile-a-minute, Oriental bittersweet, wetlands nightshade

North Dakota

Forbs: Canada thistle, Dalmatian toadflax, diffuse knapweed, leafy spurge, musk thistle, purple
loosestrife, Russian knapweed, spotted knapweed, yellow starthistle
Trees: Tamarisk
Vines: Field bindweed

Ohio

Forbs: Canada thistle, giant hogweed, musk thistle, prickly Russian thistle, purple loosestrife

(Continued )
670 n STATE-BY-STATE DESIGNATIONS OF INVASIVE OR NOXIOUS WEEDS

Graminoids: Sorghum, Sudangrass

Vines: Mile-a-minute

Oklahoma

Forbs: Canada thistle, musk thistle, Scotch thistle

Oregon

Aquatic: Anchored waterhyacinth, Eurasian watermilfoil, giant salvinia, hydrilla, water chestnut
Forbs: Buffalobur nightshade, bull thistle, burning bush, Canada thistle, common St. Johnswort,
Dalmatian toadflax, diffuse knapweed, dyer’s woad, garlic mustard, giant hogweed, giant
knotweed, hairy whitetop, halogeton, Iberian knapweed, Italian thistle, Japanese knotweed,
leafy spurge, lens-pod cress, musk thistle, perennial pepperweed, plumeless thistle, purple
loosestrife, purple starthistle, Russian knapweed, Scotch thistle, spotted knapweed, squarrose
knapweed, yellow starthistle, yellow toadflax
Graminoids: African feathergrass, Brazilian satintail, cogongrass, common cordgrass, dense-
flowered cordgrass, kikuyugrass, kyasuma grass, medusahead, mission grass, quackgrass, salt
meadow cordgrass, smooth cordgrass
Shrubs: French broom, Gorse, Himalayan blackberry, Portuguese broom, Scotch broom, shrubby
Russian thistle, silverleaf nightshade, Spanish broom, tropical soda apple, turkey berry
Trees: Melaleuca, tamarisk
Vines: Dodders, English ivy, field bindweed, kudzu, wetlands nightshade

Pennsylvania

Forbs: Bull thistle, Canada thistle, giant hogweed, musk thistle, purple loosestrife
Graminoids: Sorghum, Sudangrass
Shrubs: Multiflora rose
Vines: Kudzu, mile-a-minute

South Carolina

Aquatic: Anchored waterhyacinth, Eurasian watermilfoil, giant salvinia, hydrilla, water chestnut,
waterhyacinth
Forbs: Giant hogweed, purple loosestrife, Russian knapweed
Graminoids: African feathergrass, Brazilian satintail, cogongrass, common reed, kikuyugrass,
kyasuma grass, mission grass
Shrubs: Shrubby Russian thistle, tropical soda apple, turkey berry
Trees: Melaleuca
Vines: Dodders, mile-a-minute, wetlands nightshade

South Dakota

Aquatic: Eurasian watermilfoil

Forbs: Canada thistle, common St. Johnswort, Dalmatian toadflax, diffuse knapweed, leafy
spurge, musk thistle, perennial pepperweed, plumeless thistle, purple loosestrife, Russian
knapweed, spotted knapweed, yellow starthistle, yellow toadflax
Shrubs: Multiflora rose
Trees: Tamarisk
Vines: Dodders, field bindweed

Tennessee

Forbs: Purple loosestrife

Shrubs: Tropical soda apple
 STATE-BY-STATE DESIGNATIONS OF INVASIVE OR NOXIOUS WEEDS n 671

Texas

Aquatic: Anchored waterhyacinth, common salvinia, Eurasian watermilfoil, giant salvinia,

hydrilla, waterhyacinth
Forbs: Purple loosestrife
Graminoids: Giant reed
Shrubs: Tropical soda apple
Trees: Brazilian peppertree, melaleuca, tamarisk
Vines: Dodders, field bindweed, kudzu

Utah

Forbs: Canada thistle, diffuse knapweed, dyer’s woad, leafy spurge, musk thistle, perennial
pepperweed, purple loosestrife, Russian knapweed, Scotch thistle, spotted knapweed,
squarrose knapweed, yellow starthistle
Graminoids: Medusahead, quackgrass
Vines: Field bindweed

Vermont

Aquatic: Anchored waterhyacinth, Eurasian watermilfoil, giant salvinia, hydrilla, parrotfeather,

variable watermilfoil, water chestnut, waterhyacinth
Forbs: Garlic mustard, giant hogweed, goutweed, Japanese knotweed, purple loosestrife,
spotted knapweed
Graminoids: African feathergrass, Brazilian satintail, cogongrass, common reed, kikuyugrass,
kyasuma grass, mission grass
Shrubs: Amur honeysuckle, Bell’s honeysuckle, Morrow’s honeysuckle, shrubby Russian thistle,
Tatarian honeysuckle, tropical soda apple, turkey berry
Trees: Melaleuca, tree of heaven
Vines: Black swallow-wort, dodders, Japanese honeysuckle, Oriental bittersweet, pale swallow-
wort, wetlands nightshade, white swallow-wort

Virginia

Forbs: Purple loosestrife

Washington

Aquatic: Eurasian watermilfoil, hydrilla, parrotfeather, water chestnut

Forbs: Buffalobur nightshade, bull thistle, burning bush, Canada thistle, common St. Johnswort,
Dalmatian toadflax, diffuse knapweed, dyer’s woad, garlic mustard, giant hogweed, giant
knotweed, hairy whitetop, Italian thistle, Japanese knotweed, leafy spurge, musk thistle,
perennial pepperweed, plumeless thistle, purple loosestrife, purple starthistle, Russian
knapweed, Scotch thistle, spotted knapweed, yellow starthistle, yellow toadflax
Graminoids: Common cordgrass, common reed, dense-flowered cordgrass, salt meadow cord-
grass, smooth cordgrass
Shrubs: Gorse, Scotch broom, silverleaf nightshade, Spanish broom
Trees: Tamarisk
Vines: Alfalfa dodder, English ivy, field bindweed, kudzu

West Virginia

Forbs: Musk thistle, plumeless thistle

Shrubs: Multiflora rose
Vines: Kudzu

(Continued )
672 n STATE-BY-STATE DESIGNATIONS OF INVASIVE OR NOXIOUS WEEDS

Trees: Autumn olive

Wisconsin

Forbs: Canada thistle, leafy spurge, purple loosestrife

Shrubs: Multiflora rose
Vines: Field bindweed

Wyoming

Forbs: Canada thistle, common St. Johnswort, Dalmatian toadflax, diffuse knapweed, dyer’s
woad, hairy whitetop, leafy spurge, musk thistle, perennial pepperweed, plumeless thistle,
purple loosestrife, Russian knapweed, Scotch thistle, spotted knapweed, yellow toadflax
Graminoids: Quackgrass
Trees: Tamarisk
Vines: Field bindweed
n Pathways of Introduction for Plants
Intentional

Aesthetics, Brazilian peppertree (Schinus terebinthifolius)

Ornamentals, and
Garden Escapees
Brooms (Cytisus spp., Genista spp.)
Burning bush (Bassia scoparia, Euonymus atropurpurea)
Carrotwood (Cupaniopsis anacardioides)
Cherokee rose (Rosa laevigata)
Chinaberry (Melia azedarach)
Chocolate vine (Akebia quinata)
Crimson fountain grass (Pennisetum setaceum)
English ivy (Hedera helix)
Exotic bush honeysuckles (Lonicera spp.)
Field bindweed (Convolvulus arvensis)
Fig buttercup (Ficaria verna)
Fire tree (Morella faya)
Giant hogweed (Heracleum mantegazzianum)
Giant reed (Arundo donax)
Gorse (Ulex europaeus)
Goutweed (Aegopodium podagraria)
Japanese barberry (Berberis thunbergii)
Japanese climbing fern (Lygodium japonicum)
Japanese honeysuckle (Lonicera japonica)
Japanese hops (Humulus japonicus)
Japanese knotweed (Fallopia japonica)
Jubata grass (Cortaderia jubata)
Kahili ginger (Hedychium gardnerianum)
Koster’s curse (Clidemia hirta)
(Continued )
674 n PATHWAYS OF INTRODUCTION FOR PLANTS

Kudzu (Pueraria montana)

Lantana (Lantana camara)
Mccartney rose (Rosa bracteata)
Melaleuca (Melaleuca quinquenervia)
Multiflora rose (Rosa multiflora)
Old World climbing fern (Lygodium microphyllum)
Oriental bittersweet (Celastrus orbiculatus)
Pampas grass (Cortaderia selloana)
Paper mulberry (Broussonetia papyrifera)
Porcelainberry (Ampelopsis glandulosa, var. brevipedunculata
Princess tree (Paulownia tomentosa)
Purple loosestrife (Lythrum salicaria)
Russian olive (Elaeagnus angustifolia)
Silk tree (Albizia julibrissin)
Strawberry guava (Psidium cattleianum)
Swallow-worts (Cynanchum spp.)
Tamarisk (Tamarix chinensis, T. ramosissima)
Toadflax (Linaria dalmatica ssp. dalmatica, L. vulgaris)
Tree of heaven (Ailanthus altissima)
Velvet tree (Miconia calvescens)
Waterhyacinth (Eichhornia crassipes)
Winter creeper (Euonymus fortunei)
Wisteria (Wisteria sinensis, W. floribunda)
Aquarium Trade Common salvinia (Salvinia minima)
Giant salvinia (Salvinia molesta)
Hydrilla (Hydrilla verticillata)
Waterhyacinth (Eichhornia crassipes)
Botanical Gardens Exotic bush honeysuckles (Lonicera spp.)
Japanese barberry (Berberis thunbergii)
Koster’s curse (Clidemia hirta)
Velvet tree (Miconia calvescens)
Water chestnut (Trapa natans)
Domestic Use (Dye, Common mullein (Verbascum thapsus)
Fish Poison)
 PATHWAYS OF INTRODUCTION FOR PLANTS n 675

Dyer’s woad (Isatis tinctoria)

Gorse (Ulex europaeus)
Yellow toadflax (Linaria vulgaris)
Erosion Control/Bank Asiatic sand sedge (Carex kobomugi)
Stabilization
Australian pine (Casuarina equisetifolia)
Brooms (Cytisus spp., Genista spp., Sarothamnus spp.)
Buffelgrass (Pennisetum ciliare)
Cogongrass (Imperata cylindrica)
Exotic bush honeysuckles (Lonicera spp.)
Giant reed (Arundo donax)
Ice plant (Carpobrotus edulis)
Japanese knotweed (Fallopia japonica)
Kikuyugrass (Pennisetum clandestinum)
Kudzu (Pueraria montana)
Melaleuca (Melaleuca quinquenervia)
Multiflora rose (Rosa multiflora)
Russian olive (Elaeagnus angustifolia)
Smooth cordgrass (Spartina alterniflora)
Livestock Forage or Buffelgrass (Pennisetum ciliare)
Fodder
Cogon grass (Imperata cylindrica)
Common cordgrass (Spartina anglica)
Gorse (Ulex europaeus)
Johnsongrass (Sorghum halepense)
Kikuyugrass (Pennisetum clandestinum)
West Indian marsh grass (Hymenachne amplexicaulis)
Medicinal Asiatic colubrina (Colubrina asiatica)
Common mullein (Verbascum thapsus)
Common St. Johnswort (Hypericum perforatum)
Field bindweed (Convolvulus arvensis)
Garlic mustard (Alliaria petiolata)
Japanese hops (Humulus japonicus)
Purple loosestrife (Lythrum salicaria)
(Continued )
676 n PATHWAYS OF INTRODUCTION FOR PLANTS

Food and Drink Fire tree (Morella faya)

Garlic mustard (Alliaria petiolata)
Giant hogweed (Heracleum mantegazzianum)
Himalayan blackberry (Rubus armeniacus)
Strawberry guava (Psidium cattleianum)
Yellow Himalayan raspberry (Rubus ellipticus)
Packing Material Cogon grass (Imperata cylindrica)
Japanese stilt grass (Microstegium vimineum)
Princess tree (Paulownia tomentosa)
Smooth cordgrass (Spartina alterniflora)
Reclamation Brooms (Cytisus spp., Genista spp., Sarothamnus spp.)
Dense-flowered cordgrass (Spartina densiflora)
Exotic bush honeysuckles (Lonicera spp.)
Silk tree (Albizia julibrissin)
Melaleuca (Melaleuca quinquenervia)
Timber/Reforestation/ Carrotwood (Cupaniopsis anacardioides)
Firewood
Fire tree (Morella faya)
Melaleuca (Melaleuca quinquenervia)
Wildlife Habitat or Chinese lespedeza (Lespedeza cuneata)
Food
Exotic bush honeysuckles (Lonicera spp.)
Multiflora rose (Rosa multiflora)
Russian olive (Elaeagnus angustifolia)
Windbreaks/ Melaleuca (Melaleuca quinquenervia)
Fencerows
Russian olive (Elaeagnus angustifolia)
Tamarisk (Tamarisk chinensis, T. ramosissima)

Unintentional

Ballast Water Asiatic sand sedge (Carex kobomugi)a

Discharge
Cheatgrass (Bromus tectorum)
 PATHWAYS OF INTRODUCTION FOR PLANTS n 677

Common reed (Phragmites australis)

Crystalline ice plant (Mesembryanthemum crystallinum)
Dense-flowered cordgrass (Spartina densiflora)
Eurasian watermilfoil (Myriophyllum spicatum)
Hoary cress (Cardaria draba)
Leafy spurge (Euphorbia esula)
Musk thistle (Carduus nutans)
Purple loosestrife (Lythrum salicaria)
Smooth cordgrass (Spartina alterniflora)
Spotted knapweed (Centaurea stoebe)
Contaminant in Seeds Canadian thistle (Cirsium arvense)a
or Plants
Cheatgrass (Bromus tectorum)
Dyer’s woad (Isatis tinctoria)
Halogeton (Halogeton glomeratus)
Koster’s curse (Clidemia hirta)
Leafy spurge (Euphorbia esula)
Medusahead (Taeniatherum caput-medusae)a
Perennial pepperweed (Lepidium latifolium)
Prickly Russian thistle (Salsola tragus)
Quackgrass (Elymus repens)
Spotted knapweed (Centaurea stoebe)
Tropical soda apple (Solanum viarum)
Yellow starthistle (Centaurea solstitialis)a
Escapes from Common salvinia (Salvinia minima)
Aquaculture
Giant salvinia (Salvinia molesta)
Hydrilla (Hydrilla verticillata)
Water chestnut (Trapa natans)
Waterhyacinth (Eichhornia crassipes)
Ocean Currents Asiatic colubrina (Colubrina asiatica)
West Indian marsh grass (Hymenachne amplexicaulis)
a
Probable means of introduction
n Impacts of Invasive Plants
Agriculture: Crops and Orchards Forbs
Canada thistle
Dyer’s woad
Leafy spurge
Prickly Russian thistle
Toadflax
Graminoids
Cheatgrass
Johnsongrass
Kikuyugrass
Quackgrass
Shrubs
Lantana
Multiflora rose
Tropical soda apple
Trees
Strawberry guava
Vines
Field bindweed
Japanese dodder
Mile-a-minute
Swallow-worts
Animal Health Forbs
Canada thistle
Common St. Johnswort
Giant hogweed
Halogeton
Leafy spurge
 IMPACTS OF INVASIVE PLANTS n 679

Toadflax
Yellow starthistle
Graminoids
Buffelgrass
Cheatgrass
Johnsongrass
Kikuyugrass
Medusahead
Shrubs
Brooms
Lantana
Rattlebox
Vines
Field bindweed
Fire Hazard Forbs
Canada thistle
Common St. Johnswort
Prickly Russian thistle
Spotted knapweed
Graminoids
Buffelgrass
Cheatgrass
Cogongrass
Fountain grass
Giant reed
Johnsongrass
Jubata grass
Kikuyugrass
Medusahead
Pampas grass
Shrubs
Brooms
Gorse
(Continued )
680 n IMPACTS OF INVASIVE PLANTS

Lantana
Trees
Melaleuca
Russian olive
Vines
Climbing ferns
Forage: Rangeland, Pasture, Hay Forbs
Chinese lespedeza
Common St. Johnswort
Dyer’s woad
Halogeton
Leafy spurge
Musk thistle
Perennial pepperweed
Spotted knapweed
Toadflax
Yellow starthistle
Graminoids
Cheatgrass
Johnsongrass
Kikuyugrass
Medusahead
Quackgrass
Shrubs
Gorse
Lantana
Multiflora rose
Tropical soda apple
Vines
Swallow-worts
Human Health: Disease, Illness, Allergies, Aquatic
Poisonous
Eurasian watermilfoil
Waterhyacinth
 IMPACTS OF INVASIVE PLANTS n 681

Forbs
Giant hogweed
Leafy spurge
Graminoids
Johnsongrass
Shrubs
Brooms
Exotic bush honeysuckles
Lantana
Rattlebox
Tropical soda apple
Trees
Australian pine
Chinaberry
Melaleuca
Paper mulberry
Vines
English ivy
Japanese hops
Oriental bittersweet
Wisteria
Hydrology: Drainage, Water Aquatic
Transportation, Water Quality
Eurasian watermilfoil
Common salvinia
Giant salvinia
Hydrilla
Waterhyacinth
Forbs
Japanese knotweed
Kahili ginger
Purple loosestrife
Graminoids
Giant reed
(Continued )
682 n IMPACTS OF INVASIVE PLANTS

Cordgrasses
West Indian marsh grass
Shrubs
Rattlebox
Trees
Australian pine
Brazilian peppertree
Russian olive
Strawberry guava
Tamarisk
Velvet tree
Native Ecosystems: Displacing Plants Aquatic
and Reducing Biodiversity
Eurasian watermilfoil
Common salvinia
Giant salvinia
Hydrilla
Water chestnut
Waterhyacinth
Forbs
Canada thistle
Chinese lespedeza
Common mullein
Common St. Johnswort
Dyer’s woad
Fig buttercup
Garlic mustard
Giant hogweed
Goutweed
Halogeton
Ice plant
Japanese knotweed
Kahili ginger
Leafy spurge
 IMPACTS OF INVASIVE PLANTS n 683

Musk thistle
Perennial pepperweed
Prickly Russian thistle
Purple loosestrife
Spotted knapweed
Toadflax
Yellow starthistle
Graminoids
Asiatic sand sedge
Buffelgrass
Cheatgrass
Cogongrass
Common reed
Cordgrasses
Fountain grass
Giant reed
Japanese stiltgrass
Johnsongrass
Jubata grass
Pampas grass
Medusahead
Quackgrass
West Indian marsh grass
Shrubs
Asiatic colubrina
Brooms
Exotic bush honeysuckles
Gorse
Japanese barberry
Koster’s curse
Lantana
Multiflora rose
Rattlebox
(Continued )
684 n IMPACTS OF INVASIVE PLANTS

Tropical soda apple

Yellow Himalayan raspberry
Trees
Australian pine
Brazilian peppertree
Carrotwood
Chinaberry
Firetree
Melaleuca
Paper mulberry
Princess tree
Russian olive
Silk tree
Strawberry guava
Tamarisk
Tree of heaven
Velvet tree
Vines
Chocolate vine
Climbing ferns
English ivy
Field bindweed
Japanese dodder
Japanese honeysuckle
Japanese hops
Kudzu
Mile-a-minute
Oriental bittersweet
Porcelainberry
Swallow-worts
Winter creeper
Wisteria
Plant Health: Hosts for Pests or Parasitic Forbs
 IMPACTS OF INVASIVE PLANTS n 685

Canada thistle
Common mullein
Prickly Russian thistle
Graminoids
Johnsongrass
Shrubs
Tropical soda apple
Vines
English ivy
Field bindweed
Japanese dodder
Japanese honeysuckle
Recreational Activities Aquatic
Eurasian watermilfoil
Common salvinia
Giant salvinia
Hydrilla
Water chestnut
Waterhyacinth
Forbs
Japanese knotweed
Yellow starthistle
Graminoids
Jubata grass
Pampas grass
Shrubs
Lantana
Multiflora rose
Rattlebox
Trees
Brazilian peppertree
Vines
Mile-a-minute
(Continued )
686 n IMPACTS OF INVASIVE PLANTS

Water Use (Excess) Forbs

Toadflax
Yellow starthistle
Graminoids
Giant reed
Shrubs
Strawberry guava
Trees
Russian olive
Tamarisk
n Major Organizations and Publications
Concerned about Invasive Plants

From USDA Invasive Plants website (http://plants.usda.gov/java/noxious), with the excep-

tion of World’s Worst 100.
Abbreviations at front of listings are used in the table following this one, “Plant Species
Listed as Invasive or Noxious by Organizations and State and Federal Governments.”

Global Scope
World’s Worst 100: Lowe, S., M. Browne, S. Boudjelas, and M. De Poorter. 100 of the World’s Worst
Invasive Alien Species. A selection from the Global Invasive Species Database. Invasive Species
Specialist Group (ISSG), Species Survival Commission (SSC) of the World Conservation Union
(IUCN), 2000. http://www.issg.org/database/species/search.asp?st=100ss.

National Scope
Federal: U.S. Department of Agriculture. Animal and Plant Health Inspection Service. Plant Protection
and Quarantine. Federal Noxious Weed List (24 May 2006). USDA Animal and Plant Health
Inspection Service, Washington, DC, May 24, 2006. http://plants.usda.gov/java/noxious
?rptType=Federal. Also, USDA APHIS Plant Protection and Quarantine. Federal Domestic
Quarantines. USDA APHIS Plant Protection and Quarantine, Washington, DC, May 24, 2006.
http://www.aphis.usda.gov/plant_health/plant_pest_info.

State and Regional Scope

Cal-IPC: California Invasive Plant Council. California Invasive Plant Inventory. Cal-IPC Publication
2006-02. California Invasive Plant Council, Berkeley, February 1, 2007. http://www.cal-ipc.org/
ip/inventory/index.php.
FLEPPC: Florida Exotic Pest Plant Council. Invasive Plant List. Florida Exotic Pest Plant Council,
Florida, October 19, 1999. http://www.fleppc.org.
HEAR: Hawaiian Ecosystems at Risk Project. Information Index for Selected Alien Plants in Hawaii. USDI,
Geological Survey. Biological Resources Division, Haleakala Field Station, Makawao, HI,
October 20, 2003. http://www.hear.org/plants/.
KY: Haragan, P. D. Weeds of Kentucky and Adjacent States: A Field Guide. Lexington: University Press of
Kentucky, 1991.
N’EAST: Uva, R. H., J. C. Neal, and J. M. DiTomaso. Weeds of the Northeast. Ithaca, NY: Cornell
University Press, 1997.
NE&GP: Nebraska Department of Agriculture, Bureau of Plant Industry, Lincoln, NE. Publications:
Stubbendieck, James, Mitchell L. Coffin, and L. M. Landholt, Weeds of the Great Plains, 2005.
(Previous edition: Stubbendieck, J., G. Y. Friisoe, and M. R. Bolick. Weeds of Nebraska and the
Great Plains. 1994.) http://www.agr.state.ne.us/division/bpi/nwp/nwp1.htm.
688 n MAJOR ORGANIZATIONS AND PUBLICATIONS CONCERNED ABOUT INVASIVE PLANTS

SEEPPC: Southeast Exotic Pest Plant Council. Invasive Exotic Pest Plants in Tennessee. Research
Committee of the Tennessee Exotic Pest Plant Council, Tennessee, October 19, 1999. http://
www.se-eppc.org/weeds.cfm.
SWSS: Southern Weed Science Society. Weeds of the United States and Canada. CD-ROM. Southern
Weed Science Society, Champaign, IL. http://www.swss.ws.
WI: Wisconsin Department of Natural Resources. Hoffmann, R., and K. Kearns, eds. Wisconsin Manual
of Control Recommendations for Ecologically Invasive Plants. Madison: Wisconsin Department of
Natural Resources, 1997. http://dnr.wi.gov/invasives/publications/manual/manual_toc.htm.
WSWS: Western Society of Weed Science. Whitson, T. D. (ed.) et al. Weeds of the West. Western
Society of Weed Science in cooperation with Cooperative Extension Services, University of
Wyoming, Laramie, WY, 2006. http://www.wsweedscience.org/.
n Plant Species Listed as Invasive
or Noxious by Organizations
and State and Federal Governments
Following are species discussed in the text that have been declared invasive by organizations
or by federal or state governments.

Key to Organization Abbreviations

Cal-IPC California Invasive Plant Council.

Federal Federal Noxious Weed List.
FLEPPC Florida Exotic Pest Plant Council.
HEAR USDI, Geological Survey. Hawaiian Ecosystems at Risk Project.
KY Weeds of Kentucky and Adjacent States: A Field Guide.
N’EAST Weeds of the Northeast.
NE&GP Weeds of Nebraska and the Great Plains.
SEEPPC Southeast Exotic Pest Plant Council.
SWSS Southern Weed Science Society.
WI Wisconsin Manual of Control Recommendations for Ecologically
Invasive Plants.
World’s Worst 100 100 of the World’s Worst Invasive Alien Species.
WSWS Western Society of Weed Science.

More information about these organizations is found in the previous list: “Major
Organizations and Publications Concerned about Invasive Plants.”

Aquatics Organizations and Federal or State Governments

Anchored waterhyacinth Federal, AL, AZ, CA, FL, MA, NC, OR, SC, TX, VT
Common salvinia TX, FL, NC
Eurasian watermilfoil Cal-IPC, FLEPPC, SEEPPC, WI, AL, CO, CT, FL, ID, MA, MT, NV, NM,
NC, OR, SC, SD, TX, VT, WA
Giant salvinia Cal-IPC, Federal, AL, AZ, CA, CO, CT, FL, MA, MS, NV, OR, SC,
TX, VT
Hydrilla Cal-IPC, FLEPPC, SEEPPC, Federal, AL, AZ, CA, CO, CT, FL, ME,
MA, MS, NV, NM, NC, OR, SC, TX, VT, WA
Parrotfeather Cal-IPC, SEEPPC, AL, CT, ME, MA, VT, WA
Variable watermilfoil CT, ME, MA, VT
Water chestnut AL, AZ, CT, FL, ME, MA, NC, OR, SC, VT, WA
Waterhyacinth World’s Worst 100, Cal-IPC, FLEPPC, SWSS, AL, AZ, CA, CT, FL, SC,
TX, VT
690 n PLANT SPECIES LISTED AS INVASIVE

Forbs Organizations and Federal or State Governments

Barbwire Russian thistle AR, CA

Buffalobur nightshade NE&GP, SWSS, WSWS, ID, OR, WA
Bull thistle Cal-IPC, N’EAST, NE&GP, SEEPPC, SWSS, WSWS, AR, CO, IA, MD,
MN, NM, OR, PA, WA
Burning bush N’EAST, NE&GP, SWSS, WSWS, Federal, CT, OR, WA
Canada thistle Cal-IPC, N’EAST, NE&GP, SEEPPC, SWSS, WI, WSWS, Federal, AK,
AR, AZ, CA, CO, CT, DE, HI, ID, IL, IN, IA, KS, KY, MD, MI, MN,
MO, MT, NB, NV, NM, NC, ND, OH, OK, OR, PA, SD, UT, WA, WI,
WY
Canary Island St. Johnswort Cal-IPC, HEAR
Carolina horsenettle KY, N’EAST, NE&GP, SWSS, AK, AZ, AR, CA, HI, IA, NV
Chinese lespedeza SEEPPC, Federal, CO, KS
Common mullein Cal-IPC, HEAR, KY, N’EAST, NE&GP, SEEPPC, SWSS, WI, WSWS,
Federal, CO, HI
Common St. Johnswort Cal-IPC, NE&GP, SEEPPC, SWSS, WI, WSWS, Federal, CA, CO, MT,
NV, OR, SD, WA, WY
Crystalline ice plant Cal-IPC
Dalmatian toadflax WSWS, Federal, AZ, CA, CO, ID, MT, NV, NM, ND, OR, SD, WA,
WY
Diffuse knapweed WSWS, Federal, AZ, CA, CO, ID, MT, NB, NV, NM, ND, OR, SD, UT,
WA, WY
Dyer’s woad WSWS, Federal, AZ, CA, CO, ID, MT, NV, NM, OR, UT, WA, WY
Fig buttercup CT, MS
Garlic mustard SEEPPC, WI, Federal, AL, CT, MA, MN, NH, OR, VT, WA
Giant hogweed Federal, AL, CA, CT, FL, MA, MN, NH, NC, OH, OR, PA,
SC, VT, WA
Giant knotweed SEEPC, Federal, CA, CT, OR, WA
Goutweed Federal, CT, MA, VT
Hairy whitetop Federal, AK, AZ, CA, HI, OR, WA, WY
Halogeton Cal-IPC, WSWS, Federal, AZ, CA, CO, HI, NM, OR
Hemp sesbania KY, SWSS, AR
Iberian knapweed WSWS, AZ, CA, NV, OR
Ice plant Cal-IPC
Italian thistle Cal-IPC, WSWS, AR, CA, IA, OR, WA
Japanese knotweed World’s Worst 100, KY, N’EAST, SEEPPC, WSWS, Federal, AL, CA,
CT, MA, NH, OR, VT, WA
Kahili ginger World’s Worst 100, HEAR
Leafy spurge World’s Worst 100, Cal-IPC, N’EAST, NE&GP, SWSS, WI, WSWS,
Federal, AK, AZ, CA, CO, CT, HI, ID, IA, KS, MA, MN, MT, NB, NV,
NM, ND, OR, SD, UT, WA, WI, WY
Lens-pod hoary cress Cal-IPC, Federal, AZ, CA, OR
Malta starthistle Cal-IPC, NV, NM
Moth mullein SWSS, WSWS, CO
Musk thistle KY, N’EAST, NE&GP, SEEPPC, SWSS, WI, WSWS, Federal, CA, CO,
ID, IL, IA, KS, KY, MD, MN, MO, NB, NV, NM, NC, ND, OH, OK, OR,
PA, SD, UT, WA, WV, WY
Oppositeleaf Russian thistle AR
Perennial Pepperweed Cal-IPC, WSWS, Federal, AK, CA, CO, CT, HI, ID, MA, MT, NV, NM,
OR, SD, UT, WA, WY
 PLANT SPECIES LISTED AS INVASIVE n 691

Forbs Organizations and Federal or State Governments

Plumeless thistle NE&GP, WI, WSWS, AZ, AR, CA, CO, ID, MD, MN, NB, NC, OR, SD,
WA, WV, WY
Prickly Russian thistle NE&GP, SWSS, WSWS, AR, CA, HI, OH
Purple loosestrife World’s Worst 100, Cal-IPC, N’EAST, NE&GP, SEEPPC, SWSS, WI,
WSWS, Federal, AL, AZ, AR, CA, CO, CT, FL, ID, IN, IA, MA, MI,
MN, MO, MT, NB, NV, NM, NC, ND, OH, OR, PA, SC, SD, TN, TX,
UT, VT, VA, WA, WI, WY
Purple starthistle AZ, CA, NV, NM, OR, WA
Russian knapweed NE&GP, SWSS, WSWS, Federal, AK, AZ, CA, CO, HI, ID, IA, KS, MT,
NV, NM, ND, OR, SC, SD, UT, WA, WY
Scotch thistle NE&GP, SWSS, WSWS, AZ, AR, CA, CO, CT, ID, MO, NV, NM, OK,
OR, UT, WA, WY
Sicilian starthistle AZ, CA
Slender Russian thistle AR, CA
Spotted knapweed Cal-IPC, KY, N’EAST, NE&GP, SEEPPC, SWSS, WI, WSWS, Federal,
AZ, CA, CO, CT, ID, MA, MT, NB, NV, NM, ND, OR, SD, UT, WA,
WY
Squarrose knapweed WSWS, AZ, CA, CO, NV, OR, UT
Western horsenettle CA
Yellow starthistle Cal-IPC, SWSS, WSWS, Federal, AZ, CA, CO, ID, MT, NV, NM, ND,
OR, SD, UT, WA
Yellow toadflax N’EAST, NE&GP, WSWS, Federal, ID, MT, NV, NM, OR, SD, WA, WY

Graminoids Organizations and Federal or State Governments

African feathergrass Federal, AL, CA, FL, MA, MN, NC, OR, SC, VT
Asiatic sand sedge CT, MA
Bald brome SEEPPC, SWSS, AR
Brazilian satintail Federal, AL, CA, FL, MA, MN, MS, NC, OR, SC, VT
Buffelgrass HEAR, Federal, AZ
California satintail CA
Cheatgrass Cal-IPC, FLEPPC, HEAR, N’EAST, NE&GP, SEEPPC, SWSS, WSWS,
Federal, CO, CT
Cogongrass World’s Worst 100, FLEPPC, SWSS, Federal, AL, CA, FL, HI, MN,
MS, NC, OR, SC, VT
Common cordgrass Cal-IPC, CA, WA
Common reed N’EAST, SEEPPC, Federal, AL, CT, MA, SC, VT, WA
Crimson fountain grass Cal-IPC, HEAR, HI
Dense-flowered cordgrass Cal-IPC, OR, WA
Elephant grass FLEPPC
Giant reed Cal-IPC, HEAR, SEEPPC, Federal, TX
Japanese brome KY, NE&GP, SEEPPC, SWSS, WSWS, Federal
Japanese stilt grass SEEPPC, Federal, AL, CT, MA
Jubata grass Cal-IPC, HEAR, HI
Kikuyugrass HEAR, WSWS, Federal, AL, CA, FL, MA, MN, NC, OR, SC, VT
Kyasuma grass Federal, AL, CA, FL, MA, MN, NC, OR, SC, VT
Medusahead Cal-IPC, SWSS, WSWS, Federal, CA, CO, NV, OR, UT
Mission grass Federal, AL, CA, FL, MA, MN, NC, OR, SC, VT
Pampas grass Cal-IPC, HEAR
Quackgrass KY, N’EST, NE&GP, SWSS, WI, WSWS, Federal, AK, AZ, CA, CO, HI,
IA, KS, OR, UT, WY

(Continued )
692 n PLANT SPECIES LISTED AS INVASIVE

Graminoids Organizations and Federal or State Governments

Ryebrome SEEPPC, SWSS, WSWS, AR

Salt meadow cordgrass Cal-IPC, OR, WA
Smooth brome NE&GP, SEEPPC, WI, Federal
Smooth cordgrass Cal-IPC, OR, WA
Soft brome SEEPPC, SWSS, WSWS, Federal
Sorghum N’EAST, NE&GP, SWSS, IN, IA, MD, NV, OH, PA
Sudangrass N’EAST, NE&GP, SWSS, IN, IA, MD, NV, OH, PA
West Indian marsh grass FLEPPC

Shrubs Organizations and Federal or State Governments

Amur honeysuckle SEEPPC, WI, Federal, CT, MA, VT

Asiatic colubrina FLEPPC
Bell’s honeysuckle CT, MA, NH, VT
Canary Island St. JohnswortHEAR
Common barberry WI, Federal, CT, MA, NH
French broom Cal-IPC, Federal, CA, HI, OR
Gorse Cal-IPC, HEAR, WSWS, Federal, CA, HI, OR, WA
Himalayan blackberry Cal-IPC, HEAR, N’EAST, Federal, OR
Japanese barberry SEEPPC, WI, Federal, CT, MA
Koster’s curse World’s Worst 100, HEAR, HI
Lantana World’s Worst 100, FLEPPC, HEAR
Morrow’s honeysuckle SEEPPC, WI, Federal, CT, MA, NH, VT
Multiflora rose N’EAST, SEEPPC, SWSS, WI, Federal, AL, CT, IN, IA, KY, MA, MO,
NH, PA, SD, WV, WI
Portuguese broom Cal-IPC, Federal, OR
Rattlebox Cal-IPC, FLEPPC
Sawtooth blackberry HEAR, N’EAST, HI
Scotch broom Cal-IPC, WSWS, Federal, CA, HI, ID, OR, WA
Shrubby Russian thistle Federal, AL, AR, CA, FL, MA, MN, NC, OR, SC, VT
Silverleaf nightshade SWSS, WSWS, AR, CA, HI, ID, NV, OR, WA
Snowpeaks raspberry HEAR, N’EAST, HI
Spanish broom Cal-IPC, Federal, HI, OR, WA
Tatarian honeysuckle N’EAST, WI, Federal, CT, MA, NH, VT
Tropical soda apple FLEPPC, SEEPPC, SWSS, Federal, AL, AZ, CA, FL, MA, MN, MS, NC,
OR, SC, TN, TX, VT
Turkey berry FLEPPC, WSWS, Federal, AL, CA, FL, HI, MA, MN, NC,
OR, SC, VT
West Indian raspberry HEAR, N’EAST
Yellow Himalayan raspberry World’s Worst 100, HEAR, N’EAST, HI

Trees Organizations and Federal or State Governments

Australian pine FLEPPC, HEAR, Federal, FL

Autumn olive SEEPPC, WI, CT, MA, NH, WV
Brazilian peppertree World’s Worst 100, Cal-IPC, FLEPPC, HEAR, FL, TX
Carrotwood FLEPPC, FL
Chinaberry FLEPPC, SEEPPC, Federal
Fire tree World’s Worst 100, HEAR, HI
Melaleuca World’s Worst 100, Federal, AL, CA, FL, MA, NC, OR, SC, TX, VT
 PLANT SPECIES LISTED AS INVASIVE n 693

Trees Organizations and Federal or State Governments

Paper mulberry SEEPPC

Princess tree SEEPPC, Federal, CT
Russian olive Cal-IPC, NE&GP, WI, WSWS, Federal, CO, CT, NM
Silk tree FLEPPC, SEEPPC, Federal
Strawberry guava World’s Worst 100, FLEPPC, HEAR
Tamarisk species World’s Worst 100, Cal-IPC, WSWS, Federal, CO, MT, NB, NV, NM,
ND, OR, SD, TX, WA, WY
Tree of heaven Cal-IPC, SEEPC, WI, Federal, CT, MA, NH, VT
Velvet tree World’s Worst 100, HEAR, HI
Woman’s tongue FLEPPC, HEAR

Vines Organizations and Federal or State Governments

Black swallow-wort N’EAST, federal, CT, MA, NH, VT

Chinese wisteria SEEPC, PLEPPC, Federal
Dodder species N’EAST, Federal, AL, AZ, AR, CA, FL, MA, MI, MN, NC, OR, SC, SD,
TX, VT, WA
English ivy Cal-IPC, SEEPPC, WI, Federal, OR, WA
Field bindweed KY, N’EAST, NE&GP, SWSS, WI, WSWS, Federal, AK, AZ, AR, CA,
CO, HI, ID, IA, KS, MI, MN, MO, MT, NM, ND, OR, SD, TX, UT, WA,
WI, WY
Japanese climbing fern FLEPPC, Federal, AL, FL
Japanese honeysuckle FLEPPC, HEAR, N’EAST, SEEPPC, SWSS, WI, Federal, CT, MA,
NH, VT
Japanese hops CT, MA
Japanese wisteria SEEPPC, Federal
Kudzu World’s Worst 100, FLEPPC, HEAR, KY, N’EAST, SEEPPC, SWSS,
Federal, CT, FL, IL, KS, KY, MA, MS, MO, OR, PA, TX, WA, WV
Mile-a-minute N’EAST, Federal, AL, CT, MA, NC, OH, PA, SC
Old World climbing fern FLEPPC, Federal, AL, FL
Oriental bittersweet N’EAST, SEEPPC, WI, Federal, CT, MA, NH, NC, VT
Pale swallow-wort Federal, CT, MA, NH, VT
Peppervine SWSS
Porcelainberry WI, Federal, CT, MA
Wetlands nightshade FLEPPC, Federal, AL, CA, FL, MA, NC, OR, SC, VT
White swallow-wort Federal, VT
Winter creeper SEEPC, WI, Federal
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Set Appendices

n Appendix A: American Species That Are

Invasive Abroad

Many species native to the United States are invasive in other parts of the world. The follow-
ing presents a sample of those that are having or are expected to have a significant impact on
ecosystems and/or economies in the regions to which they have been introduced. The list
reflects those regions best studied for exotic species and does not necessarily represent the
full nonnative range of a given taxon.

I. Plants

Common Name Scientific Name Areas Where It Is Invasive

Trees
Black cherry Prunus serotina Northwest Europe
Black locust Robinia pseudoacacia Europe, Australia, southern Africa
Douglas fir Pseudotsuga menzeiseii Argentina
Honey locust Gleditsia triacanthus Europe
Honey mesquite* Prosopis glandulosa South America, southern Africa, Australia
Loblolly pine Pinus taeda Argentina, Brazil
Lodgepole pine Pinus contorta Argentina, Australia, New Zealand
Monterey pine Pinus radiata Argentina, southern Africa, Australia, New
Zealand
Parkinsonia Parkinsonia aculeata Australia
Slash pine Pinus elliotti Brazil, southern Africa, Australia
Velvet mesquite Prosopis velutina South America, Africa
Shrubs
Desert false indigo Amorpha fruticosa Southern Europe, temperate Asia
Oregon grape Mahonia aquifolium Europe
Sand blackberry Rubus cuneifolius Chile, Australia, southern Africa
Forbs
Annual ragweed Ambrosia artemisiifolia Europe
Canada goldenrod Solidago canadensis Europe, temperate Asia
Giant goldenrod Solidago gigantea Europe
Jerusalem artichoke Helianthus tuberosa Europe

(Continued )
696 n APPENDIX A

Common Name Scientific Name Areas Where It Is Invasive

Large-leaved lupine Lupinus polyphyllus Europe, New Zealand

Rough cocklebur Xanthium strumarium Southern Africa, Australia
Graminoids
Broomsedge bluestem Andropogon virginicus Tropical Africa
Finestem needlegrass Nassella tenuissima Southern Africa
Jointed rush Juncus articularis Australia, New Zealand
Rice cutgrass Leersia oryzoides Australia
Smooth cordgrass Spartina alterniflora New Zealand
Soft rush Juncus effusus Australia
Tall flatsedge Cyperus eragrostis Southern Europe, Australia
Succulents
Century plant Agave americana Southern Europe, southern Africa, Canary
Islands, Azores
Erect prickly pear* Opuntia stricta Australia, Madagascar southern Europe,
southern Africa, temperate Asia,
Tree cholla Cylindropuntia imbricata Southern Africa
Aquatic Plants
American water fern Azolla filiculoides Europe
American waterweed Elodea canadensis Southern Europe, Australia

II. Invertebrates

Common Name Scientific Name Areas Where It Is Invasive

Trematode
American liverfluke Fuscioloides magna Europe
Ctenophore
Warty comb jelly* Mnemiopsis leidyi Black Sea, Mediterranean Sea,
North Sea, Baltic Sea
Mollusks
American oyster drill Urosalpinx cinerea Northern Europe
Atlantic jackknife clam Ensis directus Northwest Europe
Bay barnacle Balanus improvisus Europe
Common slipper shell Crepidula fornicata Europe
Rosy wolfsnail * Euglandia rosea Islands of Pacific and Indian
oceans
Crustaceans
American lobster Homarus americanus Europe
Louisiana crayfish Procambarus clarkii Africa, China, Europe
Signal crayfish Pacifastacus lenisculus Europe, Japan
Spiny-cheeked crayfish Oronectes limosus Europe
Recluse spiders Loxosceles spp. Europe
Insects
Ambrosia beetle Gnathotrichus materarius Europe
American rice water weevil Lissorhoptrus oryzophilus China
Cypress tip moth Argyresthia cupressella Europe
Eastern subterranean termite Reticulitermes flavipes Europe
 APPENDIX A n 697

Common Name Scientific Name Areas Where It Is Invasive

Fall webworm Hyphantria cunea China

Loblolly pine mealybug Oracella acuta China
Oak lacebug Corythucha arcuata Europe
Orange spruce needleminer Coleotechnites piceaella Europe
Sycamore lacebug Corythucha ciliata Europe
Western conifer seed bug Leptoglossus occidentalis Europe

III. Vertebrates

Common Name Scientific Name Areas Where It Is Invasive

Fish
Black bullhead Ameiurus melas Europe
Brook trout Salvelinus fontinalis Europe
Brown bullhead Ameiurus nebulosa Europe, Chile, New Zealand
Channel catfish Ictalurus punctatus Europe
Fathead minnow Pimephales promelas Europe
Lake trout Salvelinus namaycush Europe
Largemouth bass * Micropterus salmoides Southern Africa
Western mosquitofish* Gambusia affinis Argentina, China, Europe
Pumpkinseed sunfish Lepomis gibbosus Europe
Rainbow trout* Oncorhynchus mykiss Argentina
Smallmouth bass Micropterus dolomieu Europe
Amphibian
American bullfrog* Lithobates catesbeianus South America, Asia
Reptile
Red-eared slider* Trachemys scripta Europe
Birds
California Quail Callipepla californica Chile
Canada Goose Branta canadenis Europe
Ruddy Duck Oxyura jamaicensis Europe
Mammals
American beaver Castor canadensis Tierra del Fuego (Argentina and
Chile)
American mink Mustela vison Argentina, Chile; Europe
Eastern cottontail Sylvilagus floridanus Europe
Grey Squirrel * Sciurus carolinensis Europe
Muskrat Ondatra zibethicus China, Europe, South America,
Japan
Raccoon Procyon lotor Europe
698 n APPENDIX A

IV. Fungi

Common Name Scientific Name Areas Where It Is Invasive

Crayfish plague* Aphanomyces astaci Europe

Cypress canker Seiridium cardinale Southern Europe

*Nominated as one of 100 of the world’s worst invading alien species by the Invasive Species Specialist Group.

Sources
Global Invasive Species Programme. http://www.gisp.org/index.asp.
Hulme, Philip E. Handbook of Alien Species in Europe. Dordrecht, Netherlands: Springer, 2009. http://
dx.doi.org/10.1007/978-1-4020-8280-1.
Lowe, S., M. Browne, S. Boudjelas, and M. Poorter. “100 of the World’s Worst Invasive Alien Species. A
Selection from the Global Invasive Species Database.” Invasive Species Specialist Group (ISSG) of
the Species Survival Commission (SSC) of the World Conservation Union (IUCN), 2004.
Electronic version available at http://www.gisp.org/publications/reports/100worst.pdf.
U.S. Department of Agriculture, National Invasive Species Information Center, International. http://
www.invasivespeciesinfo.gov/international/main.shtml.
Weber, Ewald. Invasive Plant Species of the World: A Reference Guide to Environmental Weeds. Cambridge,
MA: CABI Publishing, 2003.
n Appendix B: Major Federal Legislation
and Agreements Pertaining to Invasive
Species
1900 Lacey Act (amended 1998)
One of the oldest laws pertaining to wildlife in the United States. With its sub-
sequent amendments, it prohibits the importation into the United States of wild
mammals, wild birds, reptiles, amphibians, fish, mollusks, and crustaceans that
appear on a list of injurious wildlife without a permit issued by the U.S. Fish
and Wildlife Service. It also prohibits the transport of such species between states,
the District of Columbia, the Commonwealth of Puerto Rico, or any territory or
possession of the United States.
1912 Plant Quarantine Act (repealed Jun 20, 2000, with passage of the Plant Protection
Act of 2000)
Gave the Animal and Plant Health Inspection Service (APHIS) the authority to
regulate importation and interstate movement of nursery stock and other plants
that may carry harmful pests and diseases. Preempted state quarantines in inter-
state commerce.
1916 National Park Service Organic Act
Promotes the eradication and control of nonindigenous species and prohibits most
introduction in national parks.
1931 Animal Damage Control Act
Gives APHIS authority to control wildlife damage, including that of nonindigenous
species, on federal, state, or private land. Protects field crops, vegetables, fruits, nuts,
horticultural crops, commercial forests; freshwater aquaculture ponds and marine
species cultivation areas; livestock on public and private range and in feedlots; public
and private buildings and facilities; civilian and military aircraft; public health.
1940 Federal Seed Act (amended 1998)
Requires accurate labeling and purity standards for seeds in commerce. Prohibits
importation and interstate movement of adulterated or misbranded seeds.
1944 Organic Act
Gives APHIS the authority to conduct plant pest eradication programs.
1944 Public Health Services Act
Regulates entry of living organisms into the U.S. that may carry or cause human
diseases.
700 n APPENDIX B

1947 Federal Insecticide, Fungicide, and Rodenticide Act. 7 U.S.C. §136

Gives EPA authority to regulate importation and distribution of substances,
including microorganisms that are intended to function as pesticides.
1951 Importation of Certain Mollusks
Provides for the inspection and treatment of goods entering the U.S. from areas
infested with any terrestrial or freshwater mollusks to control entry of such organisms.
1952 International Plant Protection Convention (IPPC)
Creates an international regime to prevent the spread and introduction of plant
and plant product pests premised on the exchange of Phytosanitary certificates
between importing and exporting countries’ national plant protection offices.
Parties have national plant protection organizations established according to the
Convention with authority in relation to quarantine control, risk analysis and
other measures required to prevent the establishment and spread of all invasive
alien species that, directly or indirectly, are pests of plants. Parties agree to co-
operate on information exchange and on the development of International
Standards for Phytosanitary Measures.
Defines pests of plants or plant products as “any form of plant or animal life, or
any pathogenic agent, injurious or potentially injurious to plants or plant
products.”
Provides for quarantine of pests involved with international trade, specifically any
“pest of potential national economic importance to the country endangered
thereby and not yet present there, or present but not widely distributed and being
actively controlled.”
1955 Convention on Great Lakes Fisheries Between the United States and Canada
Establishes the Great Lakes Fisheries Commission, whose purpose is to control
and eradicate the nonnative, highly invasive Atlantic sea lamprey from the Great
Lakes.
1970 National Environmental Policy Act (NEPA). Public Law 91-190
Requires federal government agencies to consider the environmental effects of
their actions through preparation of environmental impact statements (EIS).
Effects of nonnative species, if harmful to the environment, must be included in
the EIS, but APHIS may approve and issue permits for importing nonindigenous
species following preparation of an environmental assessment rather than an envi-
ronmental impact statement. Permits for importing nonindigenous species into
containment facilities or interstate movement between containment facilities are
excluded from NEPA requirements.
1973 Endangered Species Act
When nonnative invasive species threaten endangered species, this act could be
used as basis for their eradication.
 APPENDIX B n 701

1974 Federal Noxious Weed Act (Secs. 2801 to 2813—repealed; superseded by the
Plant Protection Act, except for Sec. 2814)
Defines noxious weeds as “any living stage (including, but not limited to, seeds
and reproductive parts) of any parasitic or other plant of a kind, or subdivision
of a kind, which is of foreign origin, is new to or not widely prevalent in the
United States, and can directly or indirectly injure crops, other useful plants, live-
stock, or poultry or other interests of agriculture, including . . . the fish and wild-
life resources of the United States or the public health.”
Authorizes APHIS to restrict the introduction and spread of nonnative noxious
weeds through port-of-entry and follow-up activities.
Authorizes permanent restrictions and emergency regulations.
1975 Convention on International Trade in Endangered Species (CITES)
Regulates intentional introductions through trade (export, re-export, import and
introduction from the sea) of plants and animals which are threatened or endan-
gered in the exporting countries. Represents an alternate model for regulating
invasive species not already covered by the IPPC or other agreements. Although
the Convention is intended to prevent harm in exporting country; it can be
applied when a species is endangered in the exporting country and considered
an invasive in the importing country.
1975 Convention on the prohibition of the development, production, and stockpiling of
bacteriological (biological) and toxin weapons and on their destruction (Biological
Weapons Convention)
While the Convention prohibits parties from developing, producing, stockpiling,
acquiring or retaining microbial or other biological agents for hostile purposes,
it allows for the “international exchange of bacterial agents and toxins and
equipment for the processing, use or production of bacterial agents and
toxins for peaceful purposes.” Could lead to the unintentional release of such
agents.
1977 Executive Order 11987, Exotic Organisms
Restricts the introduction of exotic species into natural ecosystems under federal
agency authority.
1978 Cooperative Forestry Assistance Act
Provides for detection, identification, surveys, and controls of forest pests.
1980 Act to Prevent Pollution from Ships (amended by the Marine Plastics Pollution
Research and Control Act of 1987)
Requires ships in U.S. waters to comply with the International Convention for the
Prevention of Pollution from Ships.
702 n APPENDIX B

1987 Clean Water Act (as amended in 1987)

Allows releases of ballast water to be permitted or otherwise controlled under sec-
tions 402 (National Pollution Discharge Elimination System) and 303(d) (Total
Maximum Daily Load program).
1990 Agricultural Quarantine Enforcement Act
Prohibits shipping of plants, fruits, and vegetables via first-class mail.
1990 Food, Agriculture, Conservation, and Trade Act
Establishes Genetic Resources Program to collect, classify, preserve, and dissemi-
nate genetic material important to agriculture.
1990 Toxic Substances Control Act
Enables EPA to regulate nonindigenous microbes.
1990 Non-indigenous Aquatic Nuisance Prevention and Control Act (NANPCA). Public
Law 101-646
Establishes Aquatic Nuisance Species Task Force to identify areas where ballast
water does not pose an environmental threat, assess whether aquatic nuisance spe-
cies threaten the ecological characteristics and economic uses of U.S. waters (other
than the Great Lakes), determine the need for controls on vessels entering U.S.
waters (other than Great Lakes), identify and evaluate approaches for reducing risk
of adverse consequences associated with intentional introduction of aquatic species.
Directs Coast Guard to issue regulations to prevent the introduction and spread of
aquatic nuisance species into the Great Lakes through ballast water
Directs Corps of Engineers to develop a program of research and technology to
control zebra mussels in and around public facilities and make available informa-
tion on control methods
1992 Wild Bird Conservation Act. Public Law 102-440
Regulates importation of foreign wild birds.
1992 Hawaii Tropical Forest Recovery Act. Public Law 102-574
Authorizes Sec. of Agriculture and USFS to establish biological control agents for
non-native species.
Creates task force to develop action plan to “promote public awareness of the
harm caused by introduced species” develop recommendations on “the benefits
of fencing or other management activities for the protection of Hawaii’s native
plants and animals from non-native species, including the identification and prior-
ities for the areas where these activities are appropriate.”
1992 Alien Species Prevention and Enforcement Act. Public Law 102-393
Makes illegal the shipment through U.S. mail of plants or plant matter whose ship-
ment is prohibited under the Federal Plant Pest Act or Plant Quarantine Act and of
animals whose shipment is prohibited under 18 U.S.C. 42; 43, or the Lacey Act.
 APPENDIX B n 703

1994 North American Agreement on Environmental Cooperation

Article 10 (2)(h) states that the Council of the Commission on Environmental Co-
operation may develop recommendations regarding exotic species that may be
harmful.
1995 Agreement on the Application of Sanitary and Phytosanitary Measures (SPS
Agreement)
A supplementary agreement to the World Trade Organization Agreement.
Provides a uniform interpretation of the measures governing safety and plant
and animal health regulations. Applicable to all sanitary and phytosanitary mea-
sures directly or indirectly affecting international trade. Sanitary and phytosanitary
measures are defined as any measure applied to protect animal or plant life or
health within (a Members’ Territory) from entry, establishment or spread of pests,
diseases, disease carrying organisms; and to prevent or limit other damage within
the (Members Territory) from the entry, establishment or spread of pests.
1996 National Invasive Species Act. Public Law 104-332
Reauthorizes and amends the Non-indigenous Aquatic Nuisance Prevention and
Control Act of 1990. Mandates regulations to prevent introduction and spread of
aquatic nuisance species into Great Lakes through ballast water.
Authorizes funding for research on aquatic nuisance species prevention and con-
trol (Chesapeake Bay, Gulf of Mexico, Pacific Coast, Atlantic Coast, San
Francisco Bay–Delta Estuary).
Requires ballast water management program to demonstrate technologies and
practices to prevent nonindigenous species from being introduced.
Modifies composition of Aquatic Nuisance Species Task Force.
Requires Task Force to develop and implement comprehensive program to control
the brown tree snake in Guam.
1998 Lacey Act (1900; amended in 1998)
Prohibits import of a number of designated species and other vertebrates, mollusks,
and crustaceans that are “injurious to human beings, to the interests of agriculture,
horticulture, forestry, or to wildlife or the wildlife resources of the United States.”
Declares importation or transportation of any live wildlife as injurious and prohib-
ited, except as provided for under the Act; but allows import of almost all species
for scientific, medical, education, exhibition, or propagation purposes.
1999 Executive Order 13112 on Invasive Species
Defines invasive species as “an alien species whose introduction does or is likely to
cause economic or environmental harm or harm to human health.”
Prevents the introduction of invasive species, provides for their control, and
reduces impacts through improved coordination of federal agencies under a
National Invasive Species Management Plan.
704 n APPENDIX B

1999 Water Resources Development Act. Public Law Public Law 106-53
Authorizes the Secretary of the Interior, in conjunction with the Great Lakes
Fishery Commission, to undertake a program for the control of sea lampreys in
and around waters of the Great Lakes.
2000 Plant Protection Act. Public Law 106-224 (Replaces the Federal Noxious Weed
Act and many other APHIS Plant Protection Authorities)
Consolidates and modernizes all major statutes pertaining to plant protection and
quarantine (Federal Noxious Weed Act, Plant Quarantine Act). Permits APHIS to
address all types of weed issues.
Increases maximum civil penalty for violation.
Authorizes APHIS to take both emergency and extraordinary emergency actions to
address incursions of noxious weeds.
2002 Farm Security and Rural Investment Act. Public Law 107-171 (Animal Health
Protection Act)
Directs the USDA to take measures ensuring the prevention, detection, control,
and eradication of diseases and pests of animals when essential to protect (a) ani-
mal health; (b) the health and welfare of the people of the U.S.; (c) the economic
interests of the livestock and related industries of the U.S.; (d) the environment
of the U.S.; and (e) interstate commerce and foreign commerce of the U.S. in ani-
mals and other articles.
2002 Public Health Security and Bioterrorism Preparedness and Response Act. Public
Law 107-188
Improves the ability of the United States to prevent, prepare for, and respond to
bioterrorism and other public health emergencies.
Ensures coordination and minimizing duplication of Federal, State, and local
planning, preparedness, and response activities, including during the investiga-
tion of a suspicious disease outbreak or other potential public health emergency.
Particularly directs attention to invasive pests and pathogens affecting livestock.
2003 Nutria Eradication and Control Act. Public Law 108-016
Authorizes the Secretary of the Interior to provide financial assistance to the State
of Maryland and the State of Louisiana for a program to implement measures to
eradicate or control nutria and restore marshland damaged by nutria.
2004 Brown Tree Snake Control and Eradication Act. Public Law 108-38
Subject to the availability of appropriated funds for this purpose, authorizes the
expansion of research and eradication and control programs in Guam to reduce
the impact of the brown tree snake and reduce the risk of its introduction or
spread to other areas.
2004 Noxious Weed Control and Eradication Act. Public Law 108-412
 APPENDIX B n 705

Establishes a program to provide financial and technical assistance to control or

eradicate noxious weeds. Subject to the availability of appropriations under sec-
tion 457(a), the Secretary of Agriculture shall make grants to weed management
entities for the control or eradication of noxious weeds. Subject to the availability
of appropriations under section 457(b), the Secretary of Agriculture shall enter
into agreements with weed management entities to provide financial and technical
assistance for the control or eradication of noxious weeds.
2004 National Plan for Control and Management of Sudden Oak Death. Public Law
108-488 (amends the Plant Protection Act)
Subject to the availability of appropriated funds for this purpose, the Secretary of
Agriculture, acting through APHIS, shall develop a national plan for the control
and management of Sudden Oak Death, a forest disease caused by the fungus-
like pathogen Phytophthora ramorum.
2005 §6006 of the Safe, Accountable, Flexible, Efficient Transportation Equity Act: A
Legacy for Users (SAFETEA-LU). Public Law 109-59 (implements 23 U.S.C. §
329, a new provision of law added to Title 23 by §6006 of SAFETEA-LU)
Includes a provision that makes activities for the control of noxious weeds and the
establishment of native species eligible for Federal-aid funds under the National
Highway System (NHS) and the Surface Transportation System (STP). The control
of terrestrial noxious weeds and aquatic weeds is commonly done by maintenance
districts or contracted crews of each State department of transportation.
Historically, maintenance activities have been the responsibility of the State and
therefore have not been eligible for Federal-aid dollars.
2005 Public Lands Corps Healthy Forests Restoration Act. Public Law 109-154 (amends
the Public Lands Corps Act of 1993)
Addresses the impact of insect or disease infestations or other damaging agents on
forest and rangeland health
2006 Salt Cedar and Russian Olive Control Demonstration Act. Public Law 109-320
Directs the Secretary of the Interior, acting through the Commissioner of
Reclamation, to carry out an assessment and demonstration program to control
salt cedar and Russian olive, and for other purposes.
2006 Great Lakes Fish and Wildlife Restoration Act of 2006. Public Law 109-326
Amends the Great Lakes Fish and Wildlife Restoration Act of 1990 to provide for
implementation of recommendations of the United States Fish and Wildlife
Service contained in the Great Lakes Fishery Resources Restoration Study.
2007 Water Resources Development Act of 2007. Public Law 110-114; Sec. 3061
Provides for the conservation and development of water and related resources, to
authorize the Secretary of the Army to construct various projects for improve-
ments to rivers and harbors of the United States, and for other purposes, including
Asian carp dispersal barrier demonstration project, Upper Mississippi River.
706 n APPENDIX B

2008 National Defense Authorization Act (NDAA) for Fiscal Year (FY) 2008
To prevent the introduction of the brown tree snake into Hawaii, the
Commonwealth of the Northern Mariana Islands, the continental United States,
or any other non-native environment as a result of the movement from Guam of
military aircraft, personnel, and cargo, including the household goods of military
personnel and other military assets.
2008 Food, Conservation, and Energy Act of 2008 (The 2008 Farm Bill) (amends the
Lacey Act)
Amends the Lacey Act to cover a broad range of plants and plant products, includ-
ing timber deriving from illegally harvested plants.
2009 Public Land Management Act of 2009. Public Law 111-11
Allows the Secretary of the Interior to prescribe measures to control nonnative
invasive plants and noxious weeds within the National Wilderness Preservation
System.

Bills Relating to Invasive Species before the U.S. Congress

The National Invasive Species Center tracks pending legislation at http://www.invasive

speciesinfo.gov/laws/federal.shtml.
GovTrack.us also allows tracking of legislation, by subject, at http://www.govtrack.us/
congress/legislation.xpd.

Sources
“Federal Laws and Regulations. Public Laws and Acts.” National Invasive Species Information Center,
U.S. Department of Agriculture, National Agricultural Library, 2010. http://www.invasive
speciesinfo.gov/laws/publiclaws.shtml.
“An Initial Survey of Aquatic Invasive Species in the Gulf of Mexico Region.” Version 4.0. A report to
the Gulf of Mexico Program, U.S. Environmental Protection Agency, EPA/OCPD Contract # 68-C
-00-121, Battelle, 2000.
n Appendix C: Selected International
Agreements and Conventions
Pertaining to Invasive Species
1959 Agreement Concerning Cooperation in the Quarantine of Plants and the
Protection against Pests and Diseases
Signature parties agree to take measures to prevent the introduction from one
country into another of plant pests and diseases and weeds specified in lists drawn
up by the parties.
(Not ratified by the United States.)
1972 International Plant Protection Convention (IPPC)
Creates an international system to prevent the spread and introduction of plant
and plant product pests based on the exchange of phytosanitary certificates
between importing and exporting countries’ national plant protection offices.
Parties have national plant protection organizations established according to the
Convention with authority in relation to quarantine control, risk analysis, and
other measures required to prevent the establishment and spread of all invasive
alien species that, directly or indirectly, are pests of plants.
(Ratified by the United States, August 18, 1972.)
1974 Convention of International Trade in Endangered Species (CITES)
Offers an alternate method for regulating invasive species not already covered in
IPPC or other agreements. Can only be used when the species in question is
endangered in the export country and considered invasive in the importing coun-
try and the pathway is intentional.
(Ratified by the United States, January 14, 1974.)
1979 Convention of Migratory Species of Wild Animals
Article III (4)(c): Parties in the range of endangered migratory species can, to the
extent feasible and appropriate, take measures to prevent, reduce, or control fac-
tors that are endangering or likely to further endanger the species, including
strictly controlling the introduction of, or controlling or eliminating already intro-
duced exotic species detrimental to a migrating species.
(Not ratified by the United States.)
1982 United Nations Convention on the Law of the Sea (UNCLOS)
Signature states will take all measures necessary to prevent, reduce and control the
intentional or accidental introduction of species, alien or new, to a particular part
708 n APPENDIX C

1982 of the marine environment, which may cause significant and harmful changes
(cont.) thereto.
(Not ratified by the United States.)
1992 Framework Convention of Climate Control
Promotes stabilization and eventual reduction of greenhouse gas concentrations in
the atmosphere. Relevance to invasive species is that changes in climate could
stimulate new invasions and increase the rate or impact of existing invasions.
(Ratified by the United States, October 15, 1992.)
1992 Agenda 21, United Nations Conference on Environment and Development (Earth
Summit, Rio de Janeiro)
A blueprint for action on many fronts where humans affect the natural environ-
ment, the program addressed unintentional and intentional introductions of alien
plants and animals and ballast water problems.
1992 Convention of Biological Diversity (CBD)/Invasive Alien Species
Calls on signature parties, to the degree possible and appropriate, to “prevent the
introduction of, control or eradicate those alien species which threaten ecosys-
tems, habitats or species.”
(Not ratified by the United States.)
1994 Agreement on the Application of Sanitary and Phytosanitary Measures (SPS
Agreement)
A supplement to the World Trade Organization Agreement that gives a uniform
interpretation of the measures governing safety and plant and animal health regu-
lations that is applied to all measures directly or indirectly affecting international
trade. Sanitary and phytosanitary measures are defined as any measure designed
to protect animal or plant life or health within a member’s territory from the intro-
duction, establishment or spread of pests, diseases, and diseases carrying organ-
isms. Calls upon members to prevent or limit other damage within their
territories from the introduction, establishment, or spread of pests. (The United
States became a member of the World Trade Organization in 1995.)
1997 Convention on the Law of Non-navigational Uses of International Watercourses
States on international watercourses shall take all necessary steps to prevent the
intentional or unintentional introduction of species, alien or new, into an
international watercourse which may have effects detrimental to the ecosystem
and result in significant harm to other countries on the waterway.
1997 International Maritime Organization
Annex. Nonmandatory “Guidelines for the Control and Management of Ships’ Ballast
Water to Minimize the Transfer of Harmful Aquatic Organisms and Pathogens.”
Among the provisions are (1) recognition that ballast water is “the most promi-
nent” pathway for transferring aquatic organisms that could pose threats to native
human, animal, and plant life and the marine environment; and (2) “Every ship
 APPENDIX C n 709

that carries ballast water should be provided with a ballast water management plan
to assist in the minimization of transfer of harmful aquatic organisms and
pathogens.”

Source
“International Laws and Regulations,” National Invasive Species Information Center, U.S. Department
of Agriculture, National Agricultural Library, 2009. http://www.invasivespeciesinfo.gov/laws/
intlagree.shtml.
n Appendix D: ISSG’s 100 of the World’s
Worst Invasive Alien Species

Those shown in bold type are described in entries in this encyclopedia.

MICRO-ORGANISM
avian malaria (Plasmodium relictum)
banana bunchy top virus (Banana bunchy top virus)
rinderpest virus1 (Rinderpest virus)
MACRO-FUNGI
chestnut blight (Cryphonectria parasitica)
crayfish plague (Aphanomyces astaci)
Dutch elm disease (Ophiostoma ulmi)
frog chytrid fungus (Batrachochytrium dendrobatidis)
phytophthora root rot (Phytophthora cinnamomi)
AQUATIC PLANT
caulerpa seaweed (Caulerpa taxifolia)
common cord-grass (Spartina anglica)
wakame seaweed (Undaria pinnatifida)
water hyacinth (Eichhornia crassipes)
LAND PLANT
African tulip tree (Spathodea campanulata)
black wattle (Acacia mearnsii)
Brazilian pepper tree (Schinus terebinthifolius)
cogongrass (Imperata cylindrica)
cluster pine (Pinus pinaster)
erect pricklypear (Opuntia stricta)
fire tree (Morella faya)
giant reed (Arundo donax)
gorse (Ulex europaeus)
hiptage (Hiptage benghalensis)
Japanese knotweed (Fallopia japonica)
Kahili ginger (Hedychium gardnerianum)
Koster’s curse (Clidemia hirta)
kudzu (Pueraria montana var. lobata)
lantana (Lantana camara)
leafy spurge (Euphorbia esula)
leucaena (Leucaena leucocephala)
melaleuca (Melaleuca quinquenervia)
mesquite (Prosopis glandulosa)
miconia (Miconia calvescens)
mile-a-minute weed (Mikania micrantha)
mimosa (Mimosa pigra)
privet (Ligustrum robustum)
 APPENDIX D n 711

pumpwood (Cecropia peltata)

purple loosestrife (Lythrum salicaria)
quinine tree (Cinchona pubescens)
shoebutton ardisia (Ardisia elliptica)
Siam weed (Chromolaena odorata)
strawberry guava (Psidium cattleianum)
tamarisk (Tamarix ramosissima)
wedelia (Sphagneticola trilobata)
yellow Himalayan raspberry (Rubus ellipticus)
AQUATIC INVERTEBRATE
Chinese mitten crab (Eriocheir sinensis)
comb jelly (Mnemiopsis leidyi)
fish hook flea (Cercopagis pengoi)
golden apple snail (Pomacea canaliculata)
green crab (Carcinus maenas)
marine clam (Potamocorbula amurensis)
Mediterranean mussel (Mytilus galloprovincialis)
Northern Pacific seastar (Asterias amurensis)
zebra mussel (Dreissena polymorpha)
LAND INVERTEBRATE
Argentine ant (Linepithema humile)
Asian longhorned beetle (Anoplophora glabripennis)
Asian tiger mosquito (Aedes albopictus)
big-headed ant (Pheidole megacephala)
common malaria mosquito (Anopheles quadrimaculatus)
common wasp (Vespula vulgaris)
crazy ant (Anoplolepis gracilipes)
cypress aphid (Cinara cupressi)
flatworm (Platydemus manokwari)
Formosan subterranean termite (Coptotermes formosanus shiraki)
giant African snail (Achatina fulica)
gypsy moth (Lymantria dispar)
khapra beetle (Trogoderma granarium)
little fire ant (Wasmannia auropunctata)
red imported fire ant (Solenopsis invicta)
rosy wolf snail (Euglandina rosea)
sweet potato whitefly (Bemisia tabaci)
AMPHIBIAN
bullfrog (Rana catesbeiana)
cane toad (Bufo marinus)
Caribbean tree frog (Eleutherodactylus coqui)
FISH
brown trout (Salmo trutta)
carp (Cyprinus carpio)
large-mouth bass (Micropterus salmoides)
Mozambique tilapia (Oreochromis mossambicus)
Nile perch (Lates niloticus)
rainbow trout (Oncorhynchus mykiss)
walking catfish (Clarias batrachus)
Western mosquito fish (Gambusia affinis)
(Continued )
712 n APPENDIX D

BIRD
Indian myna bird (Acridotheres tristis)
red-vented bulbul (Pycnonotus cafer)
starling (Sturnus vulgaris)
REPTILE
brown tree snake (Boiga irregularis)
red-eared slider (Trachemys scripta)
MAMMALS
brushtail possum (Trichosurus vulpecula)
domestic cat (Felis catus)
goat (Capra hircus)
grey squirrel (Sciurus carolinensis)
macaque monkey (Macaca fascicularis)
mouse (Mus musculus)
nutria (Myocastor coypus)
pig (Sus scrofa)
rabbit (Oryctolagus cuniculus)
red deer (Cervus elaphus)
red fox (Vulpes vulpes)
ship rat (Rattus rattus)
small Indian mongoose (Herpestes javanicus)
stoat (Mustela erminea)
1
Declared eradicated in 2010.

Source
Lowe S., Browne M., Boudjelas S., De Poorter M. 100 of the World’s Worst Invasive Alien Species, A selec-
tion from the Global Invasive Species Database. The Invasive Species Specialist Group (ISSG) a special-
ist group of the Species Survival Commission (SSC) of the World Conservation Union (IUCN),
2002. http://www.issg.org/database/species/reference_files/100English.pdf.
(First published as special lift-out in Aliens 12, December 2000. Updated and reprinted version:
November 2004.) Used with permission.
n Glossary
Achene. A small, dry, hard one-seed fruit.
Adventive. Refers to an introduced species that has arrived in a new habitat or environment
without the aid of humans and that has not established a self-replacing population.
Aeciospore. A fungal spore produced in an aecium. Each spore has two nuclei and is part of
a chain of spores.
Aecium. The cuplike fruiting body of some rust fungi.
Aerenchyma. Pithy respiratory tissue, common in stems of some aquatic plant species.
Agnathan. Member of the class Agnatha, the jawless fish.
Alate. Winged reproductive adult of a social insect, such as ants and termites.
Alien (species). A nonnative species. A species found beyond its normal range limits.
Synonyms: exotic, nonindigenous.
Allee effect. The consequences of low population density when the presence of too few
individuals greatly reduces reproductive success.
Alleleopathy. Condition in which one plant or species exudes chemicals that prevent the
growth of other plants in the immediate vicinity.
Altricial. Refers to recently hatched birds or other newborn animals that have closed eyes
and little or no down or fur, and that are unable to leave the nest and therefore must depend
upon the parents for food.
Anadromous. Refers to fish that spend most of their lives in salt water but ascend freshwater
streams to spawn.
Anecic. Refers to deep-burrowing earthworms that inhabit the lower layers of the soil.
Annelid worm. Any member of the phylum Annelida, the segmented worms.
Annual. A plant that germinates from seed, matures, and dies in one season.
Apical (snail). The tip of a spiraling shell.
Apomictic. Refers to a flower than does not require pollination to produce seed.
Aquatic. Refers to a plant growing primarily or entirely in water, either rooted or free-floating.
Aril. The fleshy coating around a seed.
Arthropod. Member of the phylum Arthropoda, invertebrates with exoskeletons, segmented
bodies, and jointed appendages. The phylum includes arachnids, insects, and crustaceans.
Ascospore. A type of spore bearing a single copy of each chromosome formed by sexual
reproduction in fungi in the Division/Phylum Ascomycetes.
Asexual reproduction. The multiplication of individuals without the fusion of gametes. Can
occur in fungi and animals through cell splitting, budding, cloning, or sporation. In plants,
formation of new plants without the transfer of pollen. In some plants, new individuals can
be generated vegetatively from parts of the parent plant.
Auricle. Earlike appendage at the base of some leaves, which clasps the stem.
714 n GLOSSARY

Awn. A bristle-shaped appendage on a grass.

Axis. The central line of any organ, such as a stem.
Barbel. Whiskerlike tactile organ in catfish and carp that houses taste glands and helps them
to find food in murky water.
Basidiospore. A spore bearing a single copy of each chromosome and found in fungi of the
Division/Phylum Basidiomycetes.
Beak (bivalve). The highest raised part of each valve, which is generally pointed and located
near the hinge.
Benthos. A collective term referring to organisms living on the seabed.
Bergmann’s Rule. An ecogeographic pattern wherein the higher the latitude or colder the
climate, the larger the body size of warmed-blooded animals compared to close relatives
living at lower latitudes and/or in warmer climates.
Biennial. A plant that lives for two years, usually flowering and setting seed in the second year.
Bilabiate. Refers to a corolla, two-lipped.
Biodiversity. The total variation and variability of life found in genes, species, communities,
ecosystems, and landscapes.
Biogeography. The science that studies the distribution patterns of species and the pro-
cesses that determine those patterns.
Biotype. A subset of a species with a particular set of genetic features.
Bivalve. A mollusk, such as a clam or mussel, that has its body covered by two rigid shells
joined by a hinge.
Blade. The portion of the leaf that extends from the leaf sheath, flat, folded, or with rolled
margins.
Bolt: Rapid growth of flower stalk.
Bract. A small scale-like leaf, usually associated with a flower.
Bulbil. Small bulb, usually growing from leaf axils.
Byssal threads. Filaments that some mollusks produce and use to fasten themselves to hard
surfaces.
Calyx. The leaf-like sepals that enclose the petals of a flower.
Canker. A localized area of dead tissue on the trunk or branch of a woody plant.
Cardinal teeth (bivalve). Ridges and grooves on the inner surfaces of both valves of a
bivalve near the front end of the hinge that help hold the shells in alignment.
Carton. Material made of undigested cellulose, mud, and termite saliva.
Catadromous. Refers to fish that spend most of their lives in freshwater but migrate to the
sea to breed.
Chasmogamous. Refers to flowers that open to allow cross-pollination.
Chitin. A strong, semitransparent, horny substance that is the main material composing the
exoskeletons of arthropods and the internal structures of certain other invertebrates.
Chlamydospore. Large, thick-walled resting spore of several kinds of certain fungi. It is the
part of the life cycle that allows survival during unfavorable conditions, such as excessive
drought or heat.
 GLOSSARY n 715

Cilia. Hairlike structures used by some cells to move themselves or to move food particles.
Clambering. Refers to shrubs or vines with stems that climb onto and over other plants.
Cleistogamous. Refers to flowers that do not open and are self-pollinated as buds.
Clitellum. Thickened, saddlelike section of the body of earthworms that secretes a viscous
fluid in which the worm’s eggs are deposited.
Colubrid. Any snake of the large and poorly defined group of nonvenomous snakes placed
in the family Colubridae.
Columella (snail). The central structural spine of coiled snail and whelk shells.
Community (ecological). All species living in the same area or a subset of them, such as the
bird community or the plant community.
Compound. Refers to leaves that are divided into leaflets.
Conidia. Asexual, nonmotile spores of a fungus such as chestnut blight.
Contact: Refers to herbicides that kill only the plant portions contacted.
Coppicing. Refers to trees that sprout many shoots from a cut stump.
Corm. A type of bulb.
Corolla. The petals of a flower.
Corona. A distinct circular growth between the corolla and the stamens, especially in the
milkweed family.
Culm. Stem of a grass, usually hollow.
Cuticle (insect). The exoskeleton, composed mostly of chitin.
Cuticle (plant). A protective waxy coating produced by the outermost cells of a leaf or other
aerial part of a plant.
Cyme. A broad and flat-topped determinate flower cluster, with central flowers opening
first.
DBH (Diameter at breast height). Standard way of expressing the diameter of a living tree.
There is no universal standard, however, as to what breast height is. In the United States,
DBH is usually measured at a height of 1.4 m (about 4.5 ft.) above ground.
Decumbent. Refers to a stem that is reclining or lying on the ground, but with the tip
upright.
Dehiscent. Refers to a seed capsule that opens, sometimes explosively.
Determinate. Refers to when a branch or stem ceases to grow after flowering.
Detritus. Organic debris composed of parts of plants, the remains of animals, and waste
products that accumulates on the ground or moves into water bodies from surrounding
terrestrial areas.
Diapause. A suspension of development in response to adverse environmental conditions.
Dioecious. Refers to male and female flowers being on different plants.
Disjunct. A distribution pattern in which parts of the range are noncontiguous, i.e., separated
geographically.
Drupe. A fleshy, one-seeded fruit. Druplet is one part of a berry fruit.
Ecology. The interrelationships among organisms and the nonliving aspects of the environ-
ment in which they live; the science that studies such interrelationships.
716 n GLOSSARY

Ecosystem. The totality of living and nonliving elements in a given area that function as a
unit to cycle nutrients and maintain a flow of energy.
Ecotype. A population that is adapted to a particular environment and displays characteris-
tics that set it apart from related populations but that has not evolved into a distinct species.
Emergent: Refers to aquatic plants that grow primarily above the water surface.
Entire. Refers to leaf margins that are smooth, not toothed or serrated.
Epiphyte. A plant that physically lives on another but obtains no nutrients from the host.
Bromeliads and tropical orchids are frequently epiphytic.The leaves of some epiphytic bro-
meliads fuse to form a tank in which water collects, creating prime breeding grounds for
mosquitoes and some treefrogs.
Erect: Growing upright, not sprawling or trailing.
Established (species). A species not native to a geographic area and with a self-replacing
population.
Exotic (species). Any nonnative species. Synonyms: alien, nonnative, nonindigenous.
Fasciated: Abnormal growth of a plant part, such as an inflorescence, causing it to be
twisted or incurved. Also called crested.
Floret. The individual flower of a grass, comprised of two bracts, the lemma and the palea,
and the pistil and stamens. Also the individual flower of a composite.
Follicle. A dry, dehiscent fruit or seed pod that splits open on the front.
Forb. A broad-leaved, green-stemmed, nonwoody plant. One type of herb.
Fouling (organism). Any organism that accumulates on solid surfaces in an aquatic envi-
ronment and impedes the normal mechanical functioning of the equipment or host on
which it resides.
Frass. Fine, powdery material that wood-eating insects produce as waste after digesting
plant matter.
Fruiting body. Multicellular structure of fungi that carries spore-forming bodies. When the
sexual stages of the life cycle are aerial, they are usually visible to the naked eye.
Gamete. A mature sexual reproductive cell, either sperm, pollen, or egg, that fuses with
another cell during fertilization to form a new organism.
Genotype. The total complement of genes in an individual or an entire species.
Gill rakes. Bony or cartilaginous, finger-like projections off the gill arch of fish that allow
filter-feeders to retain food particles and keep solids from entering the gill cavity. Also called
gill rakers.
Glabrous: Smooth; not rough, fuzzy, or hairy.
Glaucous. Covered with a bloom, a whitish substance that rubs off.
Gloger’s Rule. An ecogeographic pattern in which warm-blooded animals in humid envi-
ronments tend to have darker pigments in skin, feathers, or hair than close relatives living
in drier environments.
Glume. Bract on a grass that does not have associated flowers.
Gonopodium. An anal fin that on some male live-bearing fish has been modified to allow
passage of sperm and internal fertilization.
Graminoid. Herbaceous plant that includes grasses, reeds, rushes, and sedges.
 GLOSSARY n 717

Granivorous. Feeding on seeds.

Gravid. Refers to a female carrying eggs or developing young; pregnant.
Habitat. The place in which an organism lives and the physical attributes of that place.
Halophyte. Plants adapted to salty conditions.
Hammock. Slightly raised tree islands surrounded by other vegetation, usually sawgrass, in
the Everglades.
Harborage (insects). Shelter or refuge.
Haustoria. The root-like, absorbing organs of a parasitic plant.
Herb. A plant with no persistent woody stem above ground. Herbaceous.
Hibernaculum. The wintering place that shelters hibernating bats.
Holoparasite. A parasitic plant that can obtain nutrients and water in no way other than
from a host plant.
Host specific. Refers to a biological control that affects only the intended plant.
Hyphae. Long, branching threads that are the main vegetative structural feature of many
fungi. See also Mycelium.
Indehiscent. Refers to a fruit that does not split open at maturity to release seeds.
Indeterminate. Refers to when a branch or stem continues to grow after flowering.
Inflorescence. Flower stalk and how the flowers are arranged.
Injurious wildlife. Species of wild mammals, wild birds, fish, mollusks, crustaceans,
amphibians, and reptiles listed under the auspices of the Lacey Act that the secretary of
the interior has determined may be harmful to the health and welfare of humans; the inter-
ests of agriculture, horticulture, or forestry; and the welfare and survival of wildlife resources
of the United States. Such species require a permit in order to be imported or trans-
ported between states.
Instar. A developmental stage in larval insects that begins and ends with a molt until the
individual is sexually mature. Often, but not always, morphological changes occur from
one instar to the next.
Introduced species. A species that has been transported, either deliberately or unintention-
ally, by humans to a location it had not previously occupied.
Introduction. The transport and release into a free-living state of a nonnative species.
Invasive species. (1) A nonnative species that is currently spreading rapidly or that has
done so in the past; (2) “An alien species whose introduction does or is likely to cause eco-
nomic or environmental harm or harm to human health” (Executive Order 13112).
Involucre. Whorl of small leaves or bracts beneath a flower or flower cluster, especially
thistles.
Irruption. A sudden, rapid increase in numbers in an animal population, usually accompa-
nied by the migration of many individuals.
Keratin. A fibrous structural material composed of protein in skin, hair, nails, feathers, and
beaks of vertebrates.
Lateral line. A sense organ in aquatic organisms such as fish and amphibians that is used to
detect movement and vibration. Commonly visible as a faint line running lengthwise down
each side.
718 n GLOSSARY

Lateral teeth (bivalve). Elongated, interlocking projections along the hinge line of a shell
that prevent the two valves from sliding against each other when the shell is closed.
Lemma. The lower of the two bracts that enclose the flower in a grass.
Lenticel. Corky cells in the bark of trees that allow air to penetrate into the interior.
Ligule. Ring on the inside of a grass leaf where the blade meets the sheath.
Lore (bird). The area between the eye and the bill on the side of the head
Macroalgae. Large, multicellular algae. Seaweed.
Macrophyte. Any plant large enough to be visible to the naked eye.
Margins. The edges of leaves or leaflets.
Membranous. Thin, parchment-like texture.
Meristem. The growing point of a plant.
Monocarpic. Refers to a plant in which the growing point of the plant becomes the flower-
ing stem, and the plant dies after flowering.
Monoecious. Refers to both female and male flowers on the same plant.
Monospecific. Consisting of one species.
Monotypic. Consisting of one genotype or ecotype, such as a clone.
Mucilaginous. Soft, moist, sticky, or gel-like.
Mycelium. A mass of branching, filamentous hyphae through which a fungus absorbs
nutrients and decomposes plant material.
Naı̈ve. Previously unexposed to a pathogen and therefore having no natural immunity.
Native. In this encyclopedia, describes species, habitats, or ecosystems known to have
existed in North America prior to European colonization. Considered by many to be the
natural elements of a continent’s biodiversity that would occur even if humans had not set-
tled the region.
Native transplant. A species that is native to the country or region in question but has been
transported beyond its natural range limits.
Naturalized (species). Refers to a nonindigenous species that is able to sustain itself repro-
ductively in the wild outside of cultivation, and has become a functioning member of a
native ecosystem.
Nitrogen fixer. A plant that, with the help of certain soil bacteria that form nodules on its
roots, can utilize atmospheric nitrogen.
Node. Joint in a stem, usually where the leaves grow.
Nonindigenous (species). A species that is not native to the place in which it now occurs.
Nonnative (species). An alien, exotic, or nonindigenous species.
Noxious (weed). A plant specified by law as being especially undesirable, troublesome, and
difficult to control.
Nuisance (species). According to the Nonindigenous Nuisance Aquatic Prevention and
Control Act of 1990, an alien aquatic species that “threatens the diversity or abundance of
native species or the ecological stability of infested waters, or commercial, agricultural or
recreational activities dependent upon such waters.”
 GLOSSARY n 719

Nymph (insect). The immature form of insects that undergo a gradual and incomplete
metamorphosis before reaching the adult stage. A nymph resembles the adult form and
never enters a pupal stage. It becomes an adult after the final molt.
Opercle. A bony plate that supports the gill covers of fishes, especially the most posterior
one.
Organelle. Any of the distinct structures within a cell that performs a specific and vital
function.
Outcompete. When a plant or animal displaces another plant or animal by being a better
competitor for some resource.
Palate (plant). A bulge in the lower lip of a figwort (Scrophulariaceae) flower that closes off
the throat.
Palea. The upper of the two bracts that enclose the flower in a grass.
Palpus. A jointed organ for touching or tasting attached to a mouthpart in arthropods.
Panicle. A loose, irregularly compound inflorescence with flowers on pedicels.
Pantropical. Found throughout the tropics.
Pappus. Appendage to a flower in the sunflower family (Asteraceaceae), such as a thistle,
which may remain attached to the fruit; may be bristled, plume-like, or scaly.
Paradioecious. Refers to male and female flowers occurring on separate plants, but any
individual can develop flowers of either gender.
Parietal callus. In some snails, a thickened deposit on the margin of the aperture and the
wall of the body whorl closest to the central spine (columella). It is often smooth and glossy
and may be adorned with raised ribs or wrinkles.
Parthenogenesis. A form of asexual reproduction in which growth and development of
embryos occurs without fertilization of the ovum by sperm.
Pathway. The means by which a species arrives at a new region.
Pedicel. Stalk that supports an individual flower or fruit.
Peduncle. Stalk that supports a flower cluster.
Perennial. A plant that lives for more than one season, although aerial parts may die back.
Perigynium. The inflated sac, which encloses the ovary in Carex species.
Petiole. A leaf stalk.
pH. The measure of acidity or alkalinity of soil or water. Using a logarithmic scale, it
describes the amount of hydrogen ions in the solution.
Pharyngeal teeth (fish). Teeth in the throat located at the back of a fish’s head.
Pheromone. Chemicals released by an organism into its environment to communicate with
other members of its own species. Some pheromones are alarm signals, while others attract
individuals to food or to a mate.
Phloem. Plant tissues that conduct foods made in the leaves to all other parts of the plant.
Photoperiod. The number of hours of daylight.
Phreatophyte. Refers to plants with roots that extend into the water table.
Phytoplankter. A tiny, usually microscopic plant that is part of the plankton.
720 n GLOSSARY

Pinna. The primary division, or branch, of a pinnate leaf. Leaflets are on the pinna. Plural,
pinnae.
Pinnate. Refers to compound leaves that have pairs of leaflets on either side of a stalk.
Evenly pinnate leaves have an even number of paired leaflets and terminate in a pair.
Oddly pinnate leaves have an uneven number of leaflets and terminate in a single leaflet.
Pinnatifid. Refers to leaves that resemble pinnately compound leaves, but with lobes that do
not reach the midrib of the leaf.
Plankton. A collective term referring to all organisms that drift in open water unable to
move under their own power against tides and currents.
Pollard. Refers to the method of severely pruning tree limbs back to the trunk or to a main
branch.
Polychaete. A member of the Polychaete class of annelid worms characterized by having
bristles on each body segment. Also called bristle worms and lugworms.
Postemergent. Refers to a herbicide that affects growing plants.
Precocial. Referring to hatchlings or newborns that are born with their eyes open, fully
feathered, or furred, and that leave the nest a short time after birth or hatching,
Preemergent. Refers to a herbicide that prevents seeds from germinating.
Pronotum. In insects, the upper surface of the first segment of the thorax.
Propagule. In animals, the minimum number of individuals of a species capable of coloniz-
ing a new area. This may be fertilized eggs, a mated female, a single male and a single female,
or a whole group of organisms, depending upon the biological and behavioral requirements
of the species. In plants, a propagule is whatever structure functions to reproduce the spe-
cies, such as a seed, spore, stem, or root cutting,
Protist. A microorganism that is either single-celled or multicellular, but lacking specialized
tissues, and has the genetic information carried in a cell nucleus.
Pubescence. Describes plant parts covered with soft, fine hairs.
Pupa. In the development of those insects that undergo complete metamorphosis, the life
stage that immediately precedes the adult stage. Some pupae remain inside the exoskeleton
of the final larval instar, but others are encased in a cocoon or chrysalis.
Pustule. A blister-like spot.
Pycnia. A flask-shaped or conical fruiting body of a rust fungus that develops below the
epidermis of the host and bears pycniospores.
Pycniospore. A spore produced in a pycnia of a rust fungus. It fuses with a hypha of the
opposite mating type to produce the sexual generation.
Raceme. Inflorescence with all the individual flowers on a single axis.
Rachilla. A small or secondary axis or rachis, especially the axis that bears florets in sedges
and grasses.
Rachis. An axis bearing flowers or leaflets.
Recurved. Bent backward.
Rhizoid (fungi). A structure that functions like a root to anchor the fungus and absorb
nutrients. Rhizoid also releases enzymes that break down organic matter.
 GLOSSARY n 721

Rhizome. A root structure below the soil surface, distinguished from a root by having nodes;
can grow shoots that produce new plants. Also called a rootstock.
Rootcrown. Top portion of a root, often containing dormant buds that sprout.
Rosette. Arrangement of leaves radiating from a central point.
Ruderal. Waste places.
Samara. An indehiscent winged fruit.
Saprotroph. Any organism that gains energy and nutrients from dead organic material.
Savanna. Grassland with scattered trees.
Scrambling. Sprawling or climbing over other plants.
Sebaceous gland. A gland in the skin that secretes an oily substance to lubricate the skin
and hair.
Semievergreen. Remaining green only in warm climates or sheltered locations.
Senesce. To grow old, turn brown. Senescence.
Sepal. A leaf-like bract that encloses a flower or flower bud. Sepals form the calyx.
Sessile (plant). Refers to flowers or leaves attached directly to stems, without pedicels or
petioles.
Settle (mollusks and crustaceans). The process by which larvae leave the plankton stage of
their life and attach to a substrate.
Sexual reproduction. The formation of new individuals from the union of two gametes, an
ovum and a sperm. In the higher plants it takes places with the transfer of pollen from a male
flower to a female flower.
Sheath. A leaf structure that surrounds and encloses a grass stem.
Shrub. Woody perennial smaller than a tree, usually with several stems.
Silicle. A small seed pod in the Mustard family.
Silique. A seed pod in the Mustard family.
Simple. Refers to leaves that are not divided or compound.
Species. A group of individuals of the same kind that can interbreed and produce viable
offspring.
Spike. A simple inflorescence with sessile flowers on a single axis. Also branch of a grass
inflorescence.
Spikelet. Secondary spike, especially a grass structure that includes glumes and florets, the
cluster of grass flowers.
Spirochete. A bacterium of the phylum Spirochaete, distinguished by its spirally twisted form.
Sporangium. Structure in which spores are formed; spore case. Plural, sporangia.
Sporation. Spore formation.
Stellate. Refers to plant hairs, star-shaped.
Stipule. The basal appendage of a petiole.
Stolon. Root stem on the soil surface that roots at nodes; may produce new plants from
sprouts; also called runners.
Stromata. The connective tissue framework or support of cells or organisms
722 n GLOSSARY

Subdioecious. Male and female flowers usually restricted to separate plants.

Submergent. Refers to aquatic plants, or parts, that grow completely underwater.
Submersed. Refers to aquatic plants that grow primarily underwater. Flowering parts may
be at or slightly above the water surface. May be free-floating or rooted.
Subshrub. A shrub in which the upper branches die back during the unfavorable seasons.
Substrate. Substance in which plants are rooted; can be soil, sand, alluvium, mud, or rock.
Surfactant. Substance added to a herbicide to help the chemicals adhere to the foliage.
Systemic. Refers to herbicides that are absorbed into plant tissues and translocated through-
out the plant.
Talus. Cone- or fan-shaped slope of loose rocks at the base of a cliff.
Telium. The pimplelike cluster of spore cases that is produced by rust fungi.
Ternate. In sets of three.
Thorax (insects). The central of three main segments of an insect’s body: the segment
between the head and the abdomen.
Tree. A woody plant with one main trunk.
Turion. A scaly, young shoot or sucker on a root or tuber.
Two-ranked. Referring to alternate arrangement of leaves; leaves are on opposite sides of the
stem, in the same geometric plane.
Umbel. Often flat-topped inflorescence resembling an umbrella, with individual pedicels
rising from a common point.
Uredinia. A reddish, pimplelike structure on the tissue of a plant infected by a rust fungus.
Vegetative reproduction. Formation of new plants from pieces of the parent plant, such as
stems, leaves, rhizomes, and stolons. Also called asexual reproduction.
Vent (reptile). Cloaca. The common cavity into which the intestinal, genital, and urinary
tracts end.
Vine. Plant whose stem requires support; can be trailing on the ground or climbing by twin-
ing, tendrils, or other means.
Whorl. Arrangement of leaves in a circle around the stem, three or more leaves at one node.
Xylem. Tissue that conducts water and dissolved minerals from the roots to all other parts of
a plant, provides mechanical support, and forms the wood of trees and shrubs.
Zooanthellae. Protozoans that live symbiotically in some jellyfishes as well as corals and
other marine organisms.
Zooid. One of the individual organisms composing a colonial animal, such as a bryozoan.
Zooplankter. Any animal, single-celled or multicelled, that is part of the plankton.
Zoospore. An asexual spore produced by some fungi that can move around by using a
tail-like appendage (flagellum).
Zygomorphic. In plants, irregular corollas that can be equally divided into mirror-image
halves in only one plane, such as pea or orchid flowers.
n General Bibliography: Selected Classic
and Contemporary Works and Major
Internet Data Sources
Alien Plant Working Group, Plant Conservation Alliance. “Weeds Gone Wild, Alien Plant Invaders of
Natural Areas.” http://www.nps.gov/plants/alien.
Animal Diversity Web, University of Michigan Museum of Zoology. http://animaldiversity.ummz
.umich.edu/site/index.html.
Bargeron, Charles T., David J. Moorhead, G. Keith Douce, Richard C. Reardon, and Arthur E. Miller.
Invasive Plants of the Eastern United States: Identification and Control. USDA Forest Service
Publication FHTET-2003-08, 2003. http://www.invasive.org/eastern/.
Baskins, Yvonne. A Plague of Rats and Rubbervines. The Growing Threat of Species Invasions. Washington,
DC: Island Press, 2002.
Bossard, Carla C., John M. Randall, and Marc C. Hoshovsky, eds. Invasive Plants of California’s
Wildlands. Berkeley: University of California Press, 2000. http://www.cal-ipc.org/ip/management/
ipcw.
Burnham, R. J. Plant Diversity Website. CLIMBERS Website, 2010. http://www-personal.umich.edu/
~rburnham/climbers.html.
Burrell, C. Colston. Native Alternatives to Invasive Plants. New York: Brooklyn Botanic Garden, 2006.
Cadotte, Marc William, Sean M. McMahon, and Tadashi Fukami. Conceptual Ecology and Invasion
Biology: Reciprocal Approaches to Nature. Invading Nature, vol. 1. Dordrecht: Springer, 2006.
California Department of Food and Agriculture. Pest Plant and Health Prevention Services (PPHPS).
http://www.cdfa.ca.gov/phpps/.
Center for Aquatic and Invasive Plants. University of Florida. http://plants.ifas.ufl.edu/.
Center for Invasive Species and Ecosystem Health, University of Georgia. http://www.invasive.org/.
Center for Invasive Weed Management (CIPM). Missouri River Watershed Coalition. http://
www.weedcenter.org/index.html.
Coates, Peter. American Perceptions of Immigrant and Invasive Species. Strangers on the Land. Berkeley:
University of California Press, 2006.
Cox, G. W. Alien Species in North America and Hawaii: Impacts on Natural Systems. Washington, DC:
Island Press, 1999.
Elton, Charles S. The Ecology of Invasions by Animals and Plants. London: Chapman and Hall, 1958.
Federal Noxious Weed Act of 1974. 7 U.S.C. §§ 2801–2814, January 3, 1975, as amended 1988 and
1994. http://www.thecre.com/fedlaw/legal2/fedweed.htm.
“Florida Invaders Series.” Entomology and Nematology Department, Cooperative Extension Sevice,
Institute of Food and Agricultural Sciences (IFAS), University of Florida. http://
entnemdept.ifas.ufl.edu/creatures/.
Introduced Species Summary Project, Columbia University. http://www.columbia.edu/itc/cerc/danoff
-burg/invasion_bio/inv_spp_summ/invbio_plan_report_home.html.
Invasispedia. http://wiki.bugwood.org/Invasipedia.
“Invasive and Exotic Species of North America.” Invasive.org: Center for Invasive Species and
Ecosystem Health, University of Georgia, Warnell School of Forestry and Natural Resources and
College of Agricultural and Environmental Sciences, Department of Entomology. http://
www.invasives.org.
Invasive Plant Atlas of New England (IPANE). http://nbii-nin.ciesin.columbia.edu/ipane.
Invasive Plant Atlas of the United States. http://www.invasiveplantatlas.org.
724 n GENERAL BIBLIOGRAPHY

Invasive Plant Information. State and Provinces Weed Lists. Center for Invasive Plant Management
(CIPM). http://www.weedcenter.org/inv_plant_info/state.html.
“Invasive Species.” Upper Midwest Environmental Sciences Center, U.S. Geological Survey. http://
www.umesc.usgs.gov/invasive_species.html.
“Invasive Species Program.” Great Lakes Science Center, U.S. Geological Survey. http://
www.glsc.usgs.gov/main.php?content=research_invasive&title=Invasive%
20Species0&menu=research.
Invasive Species Specialist Group (ISSG). “The Global Invasive Species Database.” International Union
for Conservation of Nature (IUCN) Species Survival Commission, Global Invasive Species
Programme (GISP). http://www.issg.org/database/welcome.
Laycock, George. The Alien Animals: The Story of Imported Wildlife. New York: Ballantine Books, 1966.
Lockwood, Julie L., Martha F. Hoopes, and Michael P. Marchetti. Invasion Ecology. Oxford: Blackwell
Publishing, 2007.
McNeely, Jeffrey A. The Great Reshuffling. Human Dimensions of Invasive Alien Species. Gland,
Switzerland: IUCN–The World Conservation Union, 2001.
Miller, James H., Erwin B. Chambliss, and Nancy J. Lowenstein. A Field Guide for the Identification of
Invasive Plants in Southern Forests. Gen. Tech. Rep. SRS-119. USDA Forest Service, Southern
Research Station. Ashville, NC. 2010. http://www.srs.fs.usda.gov/pubs/35292.
Miller, James H., Steven T. Manning, and Stephen F. Enloe. A Management Guide for Invasive Plants in
Southern Forests. Gen. Tech. Rep. SRS-131. USDA Forest Service, Southern Research Station,
Ashville, NC. 2010. http://www.srs.fs.usda.gov/pubs/35292.
Mooney, H. A., and J. A. Drake, eds. Ecology of Biological Invasions of North America and Hawaii.
Ecological Studies 58. New York: Springer-Verlag, 1986.
Mooney, Harold A., and Richard J. Hobbs. Invasive Species in a Changing World. Washington, DC:
Island Press, 2000.
Motooka, P., L. Castro, D. Nelson, G. Nagai, and L. Ching. 2003. Weeds of Hawaii’s Pastures and Natural
Areas: An Identification and Management Guide. College of Tropical Agriculture and Human
Resources. University of Hawai’i at Manoa. http://www.ctahr.hawaii.edu/invweed/weedsHi.html.
National Association of Exotic Pest Plant Councils (NAEPPC). http://www.naeppc.org/.
“Non-Native and Invasive Species.” Smithsonian Marine Station at Fort Pierce, FL. http://
www.sms.si.edu; http://www.sms.si.edu/irlspec/Nonnatives.htm.
“Non-Native Species” Florida Fish and Wildlife Conservation Commission, Tallahossee. http://
myfwc/wildlifehabitats/nonnatives.
Old, Richard. “1,200 Weeds of the 48 States and Adjacent Canada, An> Interactive Identification
Guide.” XID Services, Inc. 2008.
OTA (Office of Technology Assessment), U.S. Congress. Harmful Nonindigenous Species in the United
States. Washington DC, 1993.
Pimentel, D., L. Lach, R. Zuniga, and D. Morrison. “Environmental and Economic Costs Associated
with Non-Indigenous Species in the United States.” Ithaca, NY: College of Agriculture and Life
Sciences, Cornell University, 1999.
Plant Conservation Alliance. http://www.nps.gov/plants/index.htm.
Plants for a Future, Edible, Medicinal, and Useful Plants for a Healthier World. http://www.pfaf.org/.
PlantWise, Garden Smart. http://www.beplantwise.org.
Randal, John M., and Janet Marinelli, eds. Invasive Plants, Weeds of the Global Garden. New York:
Brooklyn Botanic Garden, 1996.
Richardson, David M., Petr Pyšek, Marcel Rejmánek, Michael G. Barbour, F. Dane Panetta, and Carol J.
West. “Naturalization and Invasion of Alien Plants: Concepts and Definitions.” Diversity and
Distributions 6: 93–107, 2000.
Ruiz, Gregory. M., and James T. Carlton. Invasive Species: Vector and Management Strategies.
Washington, DC: Island Press, 2003.
Sax, Dov F., John J. Stachowicz, and Steven D. Gaines. Species Invasions. Insights into Ecology, Evolution,
and Biogeography. Sunderland, MA: Sinauer Associates, Inc., 2005.
Simberloff, Daniel. “Confronting Introduced Species: A Form of Xenophobia?” Biological Invasions 5:
179–192, 2003.
 GENERAL BIBLIOGRAPHY n 725

Southeast Exotic Pest Plant Council Invasive Plant Manual. SEEPPC. http://www.se-eppc.org/manual/.
“Species of Concern.” United States Federal Aquatic Nuisance Species Task Force. http://
www.anstaskforce.gov/soc.php.
Swearingen, J., K. Reshetiloff, B. Slattery, and S. Zwicker. Plant Invaders of Mid-Atlantic Natural Areas.
National Park Service and U.S. Fish and Wildlife Service, 2002. http://www.invasive.org/eastern/
midatlantic/.
Todd, Kim. Tinkering with Eden: A Natural History of Exotics in America. New York: W. W. Norton,
2001.
U.S. Department of Agriculture, Agriculture Research Service, National Genetic Resources Program.
Germplasm Resources Information Network—(GRIN). National Germplasm Resources Laboratory,
Beltsville, Maryland.http://www.ars-grin.gov/cgi-bin/npgs/html/index.pl.
U.S. Department of Agriculture. Animal and Plant Health and Inspection Service (APHIS). http://
www.aphis.usda.gov/plant_health/plant_pest_info/pest_detection/index.shtml.
U.S. Department of Agriculture. Fire Effects Information System. Rocky Mountain Research Station, Fire
Sciences Laboratory, 2008. http://www.fs.fed.us/database/feis/.
U.S. Department of Agriculture, Forest Service, Invasive Species Program. http://www.fs.fed.us/invasi-
vespecies/index.shtml.
U.S. Department of Agriculture, National Invasive Species Information Center (NISIC). http://
www.invasivespeciesinfo.gov/.
U.S. Department of Agriculture, Natural Resources Conservation Service. Plants Database. http://
plants.usda.gov/.
U.S. Department of Agriculture, Natural Resources Conservation Service. Plants Database. “Invasive
and Noxious Weeds.” http://plants.usda.gov/java/noxComposite.
U.S. Department of Agriculture, Natural Resources Conservation Service. Plants Database. Plants
Profile. http://plants.usda.gov/java/nameSearch.
U.S. Geological Survey. Nonindigenous Aquatic Species Database, Gainesville, FL. http://
nas.er.usgs.gov.
Van Driesche, R., B. Blossey, M. Hoodle, S. Lyon, and R. Reardon. Biological Control of Invasive Plants in
the Eastern United States. USDA Forest Service Publication FHTET-2002-04, 2002. http://
wiki.bugwood.org/Archive:BCIPEUS.
Virginia Cooperative Extension. Virginia Tech Weed Identification Guide. http://www.ppws.vt.edu/
weedindex.htm.
Vitousek, P. M., C. M. D’Antonio, L. L. Loope, and R. Westbrook. “Biological Invasions as Global
Environmental Change.” American Scientist 84: 468–478, 1996.
Weber, Ewald. 2003. Invasive Plant Species of the World: A Reference Guide to Environmental Weeds.
Cambridge, MA: CABI Publishing, 2003.
Zheng, Hao, Yun Wu, Jianquing Ding, Denise Binion, Weidong Fu, and Richard Reardon. 2004.
Invasive Plants of Asian Origin Established in the US and Their Natural Enemies, vol. 1, 2004. http://
www.invasive.org/weeds/asian.
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n Index
Page numbers in boldfaced type refer to a main entry in the encyclopedia and “t” indicates table.
Acarapis woodi. See Honeybee Tracheal Mite
Acclimatization societies, 238, 245
Acentria ephemerella, biological control (plants)
Eurasian watermilfoil, 324–25
Aceria imperata, biological control (plants)
cogongrass, 447
Aceria malherbae, biological control (plants)
field bindweed, 609
Aceria salsolae, biological control (plants)
prickly Russian thistle, 413
Achatina fulica. See Giant African Snail
Acidotheres tristis. See Common Myna
Acremonium zonaatum, biological control (plants)
waterhyacinth, 342–43
Adedarach species. See Chinaberry
Adelges tsugae. See Hemlock Woolly Adelgid
Adoretus sinicus. See Chinese rose beetle
Adventive species, xiv (v. 1)
Aecidium mori var. broussonetia, biological
control (plants)
paper mulberry, 564
Aedes albopictus. See Asian Tiger Mosquito
Aedes eagypti. See Yellow fever mosquito
Aegopodium podagraria. See Goutweed
African Clawed Frog, xxivt (v. 1), xxvt (v. 1), 18,
19, 201–5
state-by-state occurrences, 295, 296
African feathergrass, 460–61
noxious designation, 653, 665, 666, 668,
669, 670, 671, 691
African fountain grass. See Crimson
Fountain Grass
African foxtail grass. See Buffelgrass
Africanized Honey Bee, xix (v. 1), xxvt (v. 1),
106–10
as varroa mite host, 103, 108
African pyle. See Giant Salvinia
Agonopterix nervosa, biological control (plants)
brooms, 501
Agreement on the Application of Sanitary and
Phytosanitary Measures, 703
Agrilus aurichalceus. See rose stemgirdler
Agrilus hyperici, biological control (plants)
common St. Johnswort, 361
Agrilus planipennis. See Emerald Ash Borer
Agropyron repens. See Quackgrass
AHB. See Africanized Honey Bee
Ailanthus. See Tree of Heaven
Ailanthus altissima. See Tree of Heaven
Ailanthus glandulosa. See Tree of Heaven
Ailanthus peregrine. See Tree of Heaven
Air potato, 607, 660
Aizoaceae, stone plant family, 383
Akala, 536, 538, 657
Akalakala, 536, 656
Akebia quinata. See Chocolate Vine
Akepa, as affected by avian malaria, 248
Alabama jumper (earthworm), 49
ALB. See Asian Longhorned Beetle
Albizia julibrissin. See Silk Tree
Albizzia julibrissin. See Silk Tree
Albonia peregrina. See Tree of Heaven
Albugo. See white leaf rust
Alelaila tree. See Chinaberry
Aleppo grass. See Johnsongrass
Alewife, xviii (v. 1), xxiiit (v. 1), 157–60,
190, 193
state-by-state occurrences, 296, 298–310
Alfalfa dodder, 611, 660
Alfalfa dwarf (disease), Glassy-Winged
Sharpshooter as vector, 137
Alien Animals (Laycock), xxxi (v. 1)
Alien, definition, xiii (v. 1)
Alkali bulrush. See cosmopolitan bulrush
Allegheny blackberry, 524, 656
Alleleopathy
Australian pine, 542
chinaberry, 553
Chinese lespedeza, 352
dyer’s woad, 364
exotic bush honeysuckles, 506
Johnsongrass, 471
kikuyugrass, 480
lantana, 521
quackgrass, 488
tree of heaven, 588
yellow starthistle, 429
Alley cat. See Feral Cat
728 n INDEX
Alliaria alliaria. See Garlic Mustard
Alliaria officinalis. See Garlic Mustard
Alliaria petiolata. See Garlic Mustard
Allorhogas species, biological control (plants)
velvet tree, 592
Alosa pseudoharengus. See Alewife
Alternaria solani, tropical soda apple
as host, 533
Altica carduorum, biological control (plants)
Canada thistle, 348
Alvars, 639
Amberique bean, 623, 660
American alligator, 663
as potentially affected by Nile monitor, 228
and West Indian marsh grass, 491
American barberry, 512, 656
American beach grass, 432, 435, 653
American bittersweet, 630–31, 632, 660
American Bullfrog, xiii (v. 1), xviii (v. 1),
xxvt (v. 1), xxvi (v. 1), 205–8
as chytrid frog fungus vector, 19
ISSG 100 worst invaders, 207, 711
state-by-state occurrences, 295, 296, 298,
299, 301, 303, 304, 306, 308, 309
American bumblebee, as varroa mite host, 103
American climbing fern, 598–99, 660
American crocodile, 663
as potentially affected by Nile monitor, 228
and West Indian Marshgrass, 491
American cupscale, 489, 653
American eel, 160, 183, 191
American elm, 23
cultivars, 25
American hogpeanut, 623, 660
American Robin, 661
and exotic bush honeysuckles, 506
and multiflora rose, 525
American wisteria, 645, 660
Aminopyralid, chemical control (plants)
common St. Johnswort, 361
Ampelopsis brevipedunculata.
See Porcelainberry
Ampelopsis glandulosa var. brevipedunculata.
See Porcelainberry
Ampelopsis glandulosa ‘Elegans,’ 633, 634
Ampelopsis heterophylla. See Porcelainberry
Amphibians, 201–14
ISSG 100 worst invaders, 711
Amur Honeysuckle 502–8, 614
noxious designation, 666, 668, 671, 692.
See also Exotic Bush Honeysuckles
Amur peppervine. See Porcelainberry
Amynthas agrestis. See Alabama jumper
Anabasis glomerata. See Halogeton
Anacardaceae. See sumac family
Anchored waterhyacinth, 339, 649
noxious designation, 665, 666, 668, 669,
670, 671, 689
Andean pampas grass. See Jubata Grass
Andes grass. See Jubata Grass
Andropogon vimineum. See Japanese Stilt Grass
Anecic worms, 50
Angle worm, 49. See European Earthworms
Anisantha tectorum. See Cheatgrass
Anitimicrobial chemicals, African Clawed
Frog, 204
Anjan grass. See Buffelgrass
Annelid worms, 48–53, 216
Anolis distichus. See Bark anole
Anolis equestris. See Knight anole
Anolis garmani. See Jamaican giant anole
Anoplophora glabripennis. See Asian Longhorned
Beetle
Antelope bitterbrush, 442, 656
Anthonomus tenebrosus, biological control
(plants), tropical soda apple, 534
Antiblemma acclinalis, biological control (plants)
Koster’s curse, 518
Antitoxicum rossicum. See Pale Swallow-Wort
Aphalara itadori, biological control (plants)
Japanese knotweed, 390
Aphids, biological control (plants), 663
Canada thistle, 347
exotic bush honeysuckle, 507
giant reed, 466
Japanese hops, 621
kikuyugrass, 481
tropical soda apple, 634
Aphtae epizooticae, Wild Pig as host, 280
Apiaceae. See carrot family
Apion fuscirostre, biological control (plants)
brooms, 501
Apion species, biological control (plants)
velvet tree, 592
Apion ulicis, biological control (plants)
gorse, 511
Apis mellifera scutellata. See Africanized
Honey Bee
Aplocera plagiata, biological control (plants)
common St. Johnswort, 361
Aporrectodea caliginosa. See European
Earthworms
Applesnail. See Golden Applesnail
Apthona species, biological control (plants)
leafy spurge, 398
Aquarium watermoss. See Giant Salvinia
Aquatic invertebrates, ISSG 100 worst
invaders, 711

Aquatic plants, 321–43
American species invasive abroad, 696
ISSG 100 worst invaders, 710
noxious designation, 689
Aquatic soda apple. See wetlands nightshade
Aquatic weeds, grass carp control of, 173, 174
Arachnids, 99–106
Araliacaeae. See ginseng family
Archanara geminipuncta, biological
control (plants)
common reed, 451
Archips asiaticus, biological control (plants)
chocolate vine, 596
Argentine Ant, xxii (v. 1), xxvt (v. 1), 110–13
ISSG 100 worst invaders, 711
state-by-state occurrences, 295–310
Aristolochia. See pipevine and wooly
Duchman’s pipe
Arizona cottontop, 436–37, 653
Arizona wheatgrass, 483, 653
Artimpaza argenteonota, biological
control (plants)
Asiatic colubrina, 496
Artipus floridanus, biological control (plants)
Australian pine, 543
Arundo. See Giant Reed
Arundo donax. See Giant Reed
Arundo selloana. See Pampas Grass
Arundo versicolor. See Giant Reed
Asclepiadaceae. See milkweed family
Asian applesnail. See Chinese Mystery Snail
Asian bittersweet. See Oriental Bittersweet
Asian Clam, xxiiit (v. 1), xxix (v. 1),
53–56, 77
state-by-state occurrences, 295–310
Asian euonymus scale, biological
control (plants)
winter creeper, 643
Asian Green Mussel, xxvt (v. 1), 56–58
state-by-state occurrences, 297, 298, 307
Asian gypsy moth, 138
Asian honey bee, as varroa mite host, 102
Asian lady beetle. See Multicolored Asian
Lady Beetle
Asian Longhorned Beetle, xix (v. 1), xxvt (v. 1),
113–16
ISSG 100 worst invaders, 711
state-by-state occurrences, 299, 301, 304
Asian nakedwood. See Asiatic Colubrina
Asian rapa whelk. See Veined Rapa Whelk
Asian snakeroot. See Asiatic Colubrina
Asian Swamp Eel, 160–62
state-by-state occurrences, 297, 298, 304
Asian tapeworm, grass carp as host, 174
INDEX n 729
Asian Tiger Mosquito, xxiv (v. 1), xxvt (v. 1),
116–20
ISSG 100 worst invaders, 711
state-by-state occurrences, 295–310
Asian water monitor, 226
Asiatic bittersweet. See Oriental Bittersweet
Asiatic clam. See Asian Clam
Asiatic Colubrina, 493–96
impacts, 683
noxious designation, 666, 692
pathways of introduction, 675, 677
uses of, 496
Asiatic Sand Sedge, 432–35
impacts, 683
noxious designation, 666, 668, 691
pathways of introduction, 433, 675, 676
Asiatic tear-thumb. See Mile-A-Minute
Asteraceae. See aster family; sunflower family
Athel tamarisk, 581, 583, 658
Atlantic cordgrass. See Cordgrasses and Their
Hybrids
Atlantic ivy, 603–4, 660
Atlantic shipworm. See Naval Shipworm
Atlantic wisteria, 645, 660
Atomacera petroa, biological control (plants)
velvet tree, 592
Atrazine, chemical control (plants)
cheatgrass, 443
medusahead, 485
mile-a-minute, 628
Australian Pine, 540–44
and Asiatic colubrina, 495
impacts, 681, 682, 684
noxious designation, 666, 692
pathways of introduction, 541, 675
uses of, 543
Australian pine borer, biological
control (plants)
Australian pine, 543
Australian river oak, 540, 541, 542, 658
Australian Spotted Jellyfish, 45–48
state-by-state occurrences, 295, 297, 298,
300, 302, 310
Austromusotima camptozonale, biological control
(plants)
climbing ferns, 601
Autumn olive, 568, 570
noxious designation, 570, 668, 669, 672, 692
Avian Malaria, xix (v. 1), xxvii (v. 1),
1–3, 24, 248
bird reservoir, 234
state-by-state occurrences, 298
Awapuhi kahili. See Kahili Ginger
Azolla species, 327
730 n INDEX

Bacterial leaf scorch (Xylalla fastidiosa), 22, 137 kudzu, 625

Bacterial leaf scorch, and Dutch Elm Disease, 22 leafy spurge, 398–99
English Ivy as host, 605 melaleuca, 561
Bagous affinis, biological control (plants) purple loosestrife, 417
hydrilla, 334 rattlebox, 529
Bagpod, 528, 650 tamarisk, 585
Bahamian brown anole. See Brown Anole toadflax, 426
Bald brome, 440, 653 tropical soda apple, 534
noxious designation, 666, 691 velvet tree, 592
Ball nut. See Water Chestnut water chestnut, 338
Banded mystery snail, 59 Begomovirus. See tobacco leaf curl
Bangasternus orientalis, biological Bell’s honeysuckle, 502–8
control (plants) noxious designation, 666, 668, 669, 691,
yellow starthistle, 430 692. See also Exotic Bush Honeysuckles
Banker horse. See Feral Horse Belut eel. See Asian Swamp Eel
Barberry family, 512 Berberidaceae. See barberry family
Barbwire Russian thistle, 411, 650 Berberis japonica. See Japanese Barberry
noxious designation, 665, 666, 690 Berberis sinensis. See Japanese Barberry
Bark anole, 215 Berberis thunbergia var. atropurpurea.
Barley, 483, 653 See Japanese Barberry
Basal bark application, herbicides, xx (v. 2) Berberis thunbergii. See Japanese Barberry
Basiliscus vittatus. See Brown basilisk Berberis x ottawensis, 512
Bat nut. See devil pod Bergmann’s Rule, xxxi (v. 1), 312, 714
Batrachochytrium dendrobatidis. See Chytrid Bermuda grass, 479, 654
Frog Fungus Big Cypress National Preserve
Bat White-Nose Syndrome Fungus, xix (v. 1), West Indian marsh grass, 490
11–14 melaleuca, 559
state-by-state occurrences, 297, 301–2, 304, Big Cypress National Preserve, spotted
306, 308, 309 tilapia, 197
Bay cedar, 495, 658 Burmese python, 218
Beach clustervine, 546, 660 melaleuca, 559
Beach layia, 386, 650 Big eye herring. See Alewife
Beach panic grass, 432, 435, 653 Bighead. See hyperparathyroidism
Beach she-oak. See Australian pine Bighead Carp, xxvt (v. 1), 163–66, 195
Beachstar, 546, 654 state-by-state occurrences, 295–300,
Bead tree. See Chinaberry 302–3, 306–9
Bean aphid, Canada thistle host, 347 Bignonia tomentosa. See Princess Tree
Bean stalk borer, Canada thistle host, 347 Big sage. See Lantana
Bearberry honeysuckle, 505, 656 Big taper. See Common Mullein
Beardless wheatgrass, 487, 654 Biogeography, xiii (v. 1), xxxii (v. 1)
Bedbug. See Common Bed Bug Biological control (plants). See species entries
Bee colonies destruction, by varroa mite, 104–5 Birds, 228–59
Beefwood. See Australian pine Bishop’s goutweed. See Goutweed
Beetles, biological control (plants) Bishop’s weed. See Goutweed
Asiatic colubrina, 496 Bitter panicum, 432
Brazilian peppertree, 547–48 Black acacia. See Rattlebox
Canada thistle, 348 Black carp, 172
climbing ferns, 601 Black dog-strangle vine. See Black Swallow-Wort
common St. Johnswort, 361 Blackfin cisco, 193
garlic mustard, 372 Black-hooded Parakeet. See Nanday Conure
hydrilla, 334 Black imported fire ant, 152
Japanese hops, 621 Black-legged tick
Japanese knotwood, 390 as Lyme disease vector, 4, 5, 6
Koster’s curse, 518 Black mangrove cichlid. See Spotted Tilapia

Black Rat, xviii (v. 1), xxvit (v. 1), 259–62, 287
ISSG 100 worst invaders, 261, 712
state-by-state occurrences, 295–310
Blacksage. See Lantana
Black snail. See Chinese Mystery Snail
Black spinytail iguana, 222–23
Black stem grain rust, 513
Black Swallow-Wort, 636–40
impacts, 678, 680, 684
noxious designation, 666, 668, 669, 671,
693. See also Swallow-Worts
Bladderpod. See bagpod
Blady grass. See Cogongrass
Blue toadflax. See Canada toadflax
BMSB. See Brown Marmorated Stink Bug
Bohemian knotweed. See giant knotweed
Boiga irregularis. See Brown tree snake
Bootanelleus orientalis, biological control (plants)
Australian pine, 543
Boreioglycaspis melaleucae, biological
control (plants)
melaleuca, 561
Borrelia burgdorferi. See Lyme Disease bacterium
Boston ivy, 603, 633, 660
Bothriocephalus opsarichthydis, 174
Botrylloides diagensis, 40
Botrylloides perspicuum, 40
Botrylloides violaceus. See Chain Tunicate
Botryllus schlosseri, 40
Botrytis cinerea, biological control (plants)
fire tree, 557
Botrytis cinerea, fire tree control, 557
Bowfin, 182, 183
Branch herring. See Alewife
Brachypterolus pulicarius, biological
control (plants)
toadflax, 426
Branched tearthumb, 600, 650
Brassicaceae. See mustard family
Brazilian elodea. See Brazilian waterweed
Brazilian oak, 540, 541–42, 543, 658
Brazilian Peppertree, 495, 544–48, 560
impacts, 682, 684, 685
ISSG 100 worst invaders, 710
noxious designation, 666, 671, 692
pathways of introduction, 673
uses of, 547
Brazilian satintail, 445, 654
noxious designation, 665, 666, 668, 669,
670, 671, 691
Brazilian waterhyacinth, 440, 649
Brazilian waterweed, 332, 334, 649
Breea arvensis. See Canada Thistle
Breea incana. See Canada Thistle
INDEX n 731
Bridal broom, 498, 656
Bristletips, 515, 589–90, 656
British Petroleum oil spill (2010), 329
Broadleaf toadflax, 424, 650
Broadleaved pepperweed. See Perennial
Pepperweed
Broad-leaved toadflax. See Dalmatian
Toadflax
Broghammerus reticulates, 220
Bromacil, chemical control (plants)
Brazilian peppertree, 547
Bromeliads, 495, 660, 661
Bromus tectorum. See Cheatgrass
Broncograss. See Cheatgrass
Broomleaf toadflax, 424, 650
Brooms, 496–502, 509
impacts, 679, 681, 683
noxious designation, 666, 667,
670, 671, 692
pathways of introduction, 673, 675, 676
uses of, 502
Brotogeris chiriri. See Yellow-chevroned
Parakeet
Brotogeris versicolurus. See White-winged
Parakeet
Broussonetia papyrifera. See Paper Mulberry
Brown Anole, xviii (v. 1), 214–17
state-by-state occurrences, 297, 298
Brown basilisk, 223
Brown Marmorated Stink Bug, xix (v. 1),
xxvt (v. 1), xxx (v. 1), 120–23
state-by-state occurrences, 296, 297, 304–6,
307, 309
Brown rat. See Norway Rat
Brown tree snake, xxiv (v. 1), xxvii (v. 1), 210
Brown Trout, xxivt (v. 1), 159, 166–68, 185
ISSG 100 worst invaders, 168, 711
state-by-state occurrences, 295–310
Bruchus atronotatus, biological control (plants)
Brazilian peppertree, 548
Bryozoans, 36–38
Bubulcus ibis. See Cattle Egret
Buckthorn family, 493
Buckwheat family, 387, 626
Budgerigars, 250
Buffalobur, 533, 650
noxious designation, 667, 670, 671, 690
Buff-backed heron. See Cattle Egret
Buffelgrass, 435–39
impacts, 679, 683
noxious designation, 665, 691
pathways of introduction, 675
Bulb panicgrass, 470, 654
Bulbous buttercup. See Fig Buttercup
732 n INDEX

Bull thistle, 345–46, 348–49, 401, 403, 650
noxious designation, 666, 667, 668, 669,
670, 671, 690
Burbot, 182, 183
Bur cucumber, 619, 660
Bureau of Land Management (BLM)
feral burros, 262, 264–65
feral horses, 273, 274–75
Burgdorfer, Dr. Willy, 5
Burmese Python, xviii (v. 1), xxivt (v. 1),
217–21
state-by-state occurrences, 297, 310
Burning, physical control (plants)
Australian pine, 543
Canada thistle, 348
cheatgrass, 442–43
Chinese lespedeza, 352
common reed, 450–51
English ivy, 605
exotic bush honeysuckles, 506
garlic mustard, 371
giant reed, 465
gorse, 511
Japanese barberry, 514
Japanese dodder, 613
Japanese stilt grass, 469
medusahead, 484
quackgrass, 488
tamarisk, 584
tree of heaven, 588
yellow starthistle, 430
Burning bush, 380, 641, 643, 650, 656
noxious designation, 666, 660, 671, 690
pathways of introduction, 673
Bursting heart, 641, 643, 656
Bush currant. See Velvet Tree
Bush honeysuckles. See Exotic Bush
Honeysuckles
Bush morning glory. See western
morning glory
Bush muhly, 437, 654
Bushy white solanum. See turkey berry
Butter and eggs. See Yellow Toadflax
Buttercup family, 366
Butterflies
and Canada thistle, 348
and garlic mustard, 371
and Japanese stiltgrass, 468,
and swallow-worts, 639
Butterflies, biological control (plants)
fire tree, 557
garlic mustard, 371
Bythotrephes cederstroemi, 96
Bythotrephes longimanus. See Spiny Water Flea
California broom, 498
California Clapper Rail, 661
and cordgrass, 457
California cordgrass, 453, 454–55, 456, 457,
654. See also Cordgrasses and Their
Hybrids
California peppertree. See Peruvian peppertree
California satintail, 445, 654
noxious designation, 666, 691
Californian thistle. See Canada Thistle
Calophasia lunula, biological control (plants)
toadflax, 426
Caloptilla sp. nr. schinella, biological
control (plants)
fire tree, 557
Camasey. See Koster’s Curse
Canada germander, 414, 650
Canada Thistle, 344–49, 401
impacts, 678, 679, 682, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 677
Canada thistle stem weevil, xxiv (v. 1), 348
Canada toadflax, 424, 650
Canadian honeysuckles, 505, 656
Canadian waterweed, 332, 649
Canary Island St. Johnswort, 359, 656
Canby’s mountain-lover, 643, 656
Candleberry myrtle. See Fire Tree
Cane. See Common Reed
Cane ti, 515, 589–90, 656
Cane tibouchina. See cane ti
Cannabidaceae. See hemp family
Caper spurge, 396, 650
Capra hircus. See Feral Goat
Caprifoliaceae. See Honeysuckle family
Carcinus maenas. See Green Crab
Carderia draba. See Hoary Cress
Cardaria latifolia. See Perennial Pepperweed
Carduus acanthoides. See plumeless thistle
Carduus arvensis. See Canada Thistle
Carduus nutans. See Musk Thistle
Carduus pycnocephalus. See Italian thistle
Carduus species, 344, 345, 399–404, 651, 652
noxious designation, 605, 607
pathways of introduction, 677. See also Musk
Thistle
Carduus tenuflorus. See slender-flowered thistle
Carex kobomugi. See Asiatic Sand Sedge
Caribbean fruit fly, 663
strawberry guava as host, 578, 579
Carolina horsenettle, 533, 650
noxious designation, 665, 666, 667,
669, 690

Carolina Parakeet, xxix (v. 1), 251
Carpobrotus edulis. See Ice Plant
Carpodacus mexicanus. See House Finch
Carposina bullata, biological control (plants)
Koster’s curse, 518
Carrizo. See Giant Reed
Carrot family, 372, 376
Carrot weed. See Carrotwood
Carrotwood, 548–51
impacts, 684
noxious designation, 666, 692
pathways of introduction, 673, 676
Cartwheel-flower. See Giant Hogweed
Cassida rubiginosa, biological control (plants)
Canada thistle, 348
Casuarina. See Australian Pine
Casuarinaceae. See casuarina family
Casuarina equisetifolia. See Australian Pine
Casuarina litorea. See Australian Pine
Casuarina littorea. See Australian Pine
Casuarina family, 540
Catalpa, 566, 659
Caterpillars, biological control (plants)
giant reed, 466
Japanese stiltgrass, 468
lantana, 521
Cat facing, fruit deformation, 122
Cattle
forage for, 437, 441, 445, 470, 479,
480, 489
Johnsongrass, 472
leafy spurge, 398
plants toxic to, common St. Johnswort, 360
toadflax, 425, 521
Cattle Egret, xiv (v. 1), xxi (v. 1), 228–32
state-by-state occurrences, 295–310
Cattley guava, 576
Cayuga Lake, and Eurasian
watermilfoil, 325
Celandine, 367, 650
Celandine poppy, 367, 650
Celastraceae. See staff-tree family;
staff-vine family
Celastrus articulatus. See Oriental Bittersweet
Celastrus orbiculatus. See Oriental Bittersweet
Celastrus sepiarius. See Asiatic Colubrina
Cenchrus ciliaris. See Buffelgrass
Cenchrus glaucus. See Buffelgrass
Cenchrus setaceus. See Crimson Fountain Grass
Centaurea biebersteinii. See Spotted Knapweed
Centaurea maculosa. See Spotted Knapweed
Centaurea solstitialis. See Yellow Starthistle
Centaurea stoebe. See Spotted Knapweed
Ceonothus asiaticus. See Asiatic Colubrina
INDEX n 733
Ceonothus capsularis. See Asiatic Colubrina
Cercopagis pengoi. See Fishhook water flea
Cercospora rodmanii, biological control (plants)
waterhyacinth, 343
Ceutrhynchus litura. See Canada thistle
stem weevil
Ceutrhynchus species, biological control (plants)
garlic mustard, 372
Ceutrhynchus trimaculatus, biological
control (plants)
musk thistle, 403
Chaetococcus phragmitis. See legless red mealybug
Chaetorellia species, biological control (plants)
yellow starthistle, 431
Chain sea squirt. See Chain Tunicate
Chain Tunicate, xxiv (v. 1), xxvt (v. 1),
39–41, 42
state-by-state occurrences, 295–97, 300,
301, 303, 304, 306, 308, 309
Chamaespecia species. See clearwing moths
Channa argus. See Northern Snakehead
Channeled apple snail. See Golden Apple Snail
Chaparral false bindweed. See western
morning glory
Character displacement, feeding adaptation, 243
Charru mussel, 56
Cheatgrass, 439–43
impacts, 678, 679, 685
and medusahead, 484
noxious designation, 666, 691
pathways of introduction, 676, 677
uses of, 443
Cheeseberry. See Yellow Himalayan Raspberry
Cheilosia corydon, biological control (plants)
musk thistle, 403
Chemical control (plants). See species entries
Chenopodiaceae. See goosefoot family
Cherokee rose, 523, 656
pathways of introduction, 673
Cherry guava. See Strawbery Guava
Chestnut bark disease fungus. See Chestnut
Blight Fungus
Chestnut Blight Fungus, xxvit (v. 1), xxviii (v. 1),
14–17, 34
state-by-state occurrences, 295–310
Chewing disease. See nigropalallidal
encephalomalacia
Chinche. See Common Bed Bug
Chilean iceplant. See sea fig
Chilo phragmitella, biological control (plants)
common reed, 451
Chinaberry, 551–54
impacts, 681, 684
noxious designation, 692
734 n INDEX
pathways of introduction, 673
uses of, 554
China tree. See Chinaberry
Chinese bush clover. See Chinese Lespedeza
Chinese-glysine. See Wisteria
Chinese honeysuckle, 614, 660. See also
Japanese Honeysuckle
Chinese Lespedeza, 349–53
impacts, 680, 682
noxious designation, 666, 667, 690
pathways of introduction, 676
Chinese Mitten Crab, xxiiit (v. 1), 86–89
ISSG 100 worst invaders, 711
state-by-state occurrences, 296
Chinese Mystery Snail, xxiiit (v. 1), 58–61
state-by-state occurrences, 295–97, 299–310
Chinese packing grass. See Japanese Stilt Grass
Chinese rose beetle, biological control (plants)
velvet tree, 592
Chinese sumac. See Tree of Heaven
Chinese tamarisk. See Tamarisk
Chinese turtledove, and lantana, 520, 661
Chinese wisteria. See Wisteria. See also Rattlebox
Chinese yam, 603, 607, 660
Chlopyralid, chemical control (plants)
wisteria, 647
Chlorflurenol, chemical control (plants)
ice plant and crystalline ice plant, 386
Chlorsulfuron, chemical control (plants)
Canada thistle, 348
dyer’s woad, 365
pepperweed and hoary cress, 409
Chocolate Vine, 594–97
impacts, 684
pathways of introduction, 673
uses of, 597
Christmasberry. See Brazilian Peppertree
Chukar Partridge, 661
and cheatgrass, 443
and Medusahead, 484
Chrysobothris tranquebarica. See Australian
pine borer
Chrysolina species, biological control (plants)
common St. Johnswort, 361
Chytonix segregate, biological control (plants)
Japanese hops, 621
Chytrid fungus. See Chytrid Frog Fungus
Chytrid Frog Fungus, xv (v. 1), 13, 18–21
African clawed frog as host, 204
American bullfrog as host, 207
state-by-state occurrences, 295–10
Chytridomycosis, 13, 204, 207
Cicadallid cotton pest, and Japanese
honeysuckle, 617
Cichla ocellaris. See Peacock cichlid
Cimex lectularius. See Common Bed Bug
Cimex hemipterus, 124
Cinnamon vine, 603, 660
Cipangopaludina chinensis malleata. See Chinese
Mystery Snail
Cipangopaludina japonica. See Japanese
mystery snail
Cirsium arvense. See Canada Thistle
Cirsium setosum. See Canada Thistle
Cirsium species, 401
Cirsium vulgare. See bull thistle
Cissus brevipedunculata. See Porcelainberry
Clarias batrachus. See Walking Catfish
Clastoptera undula, biological control (plants)
Australian pine, 543
Clearwing moths, biological control (plants)
leafy spurge, 398
Cleonis pigra, Canada thistle control, 349
Clethodim, chemical control (plants)
quackgrass, 488
Cletus schmidti, biological control (plants)
mile-a-minute, 629
Clidemia. See Koster’s Curse
Clidemia crenata. See Koster’s Curse
Clidemia elegans. See Koster’s Curse
Clidemia hirta. See Koster’s Curse
Climbing euonymus. See Winter Creeper
Climbing fern family, 597
Climbing ferns, 597–602
impacts, 680, 684
noxious designation, 665, 667, 693
pathways of introduction, 673
Climbing milkweed. See Black Swallow-Wort
Climbing nightshade, 532, 637, 660
Climbing prairie rose, 524, 656
Climbing spindleberry. See Oriental Bittersweet
Clitocybe tabescens, biological control (plants)
Australian pine, 543
Clopyralid chemical control (plants)
Canada thistle, 348
Chinese lespedeza, 352
common mullein, 352
spotted knapweed, 421
yellow strarthistle, 430
Clusiaceae. See mangosteen family
Cnidarians, 45–48
Coachella Valley Preserve
tamarisk eradication, 584
Coastal sand spurge, 434, 650
Coast she-oak. See Australian Pine
Cobicula fluminea. See Asian Clam
Coccinella septempunctata. See Seven-spotted
lady beetle

Cochlearia draba. See Hoary Cress
Cockroach berry, 532–33, 656
Cocostroma myconae, biological control (plants)
velvet tree, 592
Codium fragile tomentosoides. See Dead man’s
fingers
Coffee colubrina, 493, 656
Cogon grass. See Cogongrass
Cogongrass, 443–47
impacts, 679, 683
pathways of introduction, 675
ISSG 100 worst invaders, 710
noxious designation, 665, 666, 667, 668,
669, 670, 671, 691
Colchis ivy, 604, 660
Coleophora klimeschiella, biological control
(plants)
prickly Russian thistle, 413
Coleophora parthenica, biological control (plants)
halogeton, 383
prickly Russian thistle, 413
Colletotrichum gloesporioides, biological control
(plants)
Koster’s curse, 518
kudzu, 625
Colletotrichum gloesporioides f. sp. miconiae,
biological control (plants)
velvet tree, 592
Colonial ascidian. See Colonial Tunicate
Colonial sea squirt. See Colonial Tunicate
Colonial Tunicate, xxiv (v. 1), xxvt (v. 1),
42–45
state-by-state occurrences, 296, 300, 303,
304, 306, 309
Colorado potato beetle, and tropical soda apple
as host, 533–34, 663
Colubrina asiatica. See Asiatic Colubrina
Columba livia. See Rock Pigeon
Columnar cactus, 495, 658
Common barberry, 512–13, 656
noxious designation, 666, 668, 669, 692
Common Bed Bug, xix (v. 1), xxvt (v. 1),
xxvt (v. 1), 123–27
state-by-state occurrences, 295–310
Common colubrina. See Asiatic Colubrina
Common coqui. See Coqui
Common cordgrass. See Cordgrasses and Their
Hybrids; noxious designation, 670, 671,
691; pathways of introduction, 675
Common dodder, 611, 660
Common elderberry, 374, 551, 656
Common goatweed. See Common St. Johnswort
Common gorse. See Gorse
Common guava, 577, 578, 579, 659
INDEX n 735
Common hops, 619, 621, 660
Common hornwort. See coontail
Common ice plant. See Crystalline Ice Plant
Common iguana. See Green Iguana
Common Mullein, xiii (v. 2), 353–57
impacts, 682, 685
pathways of introduction, 674, 675
noxious designation, 666, 667, 690
uses of, 357
Common Myna, xxiiit (v. 1), 1, 232–34
ISSG 100 worst invaders, 234, 712
lantana, 520
state-by-state occurrences, 297
strawberry guava, 578
and velvet tree, 591
Common parsnip, 373, 650
Common Periwinkle, 61–63
state-by-state occurrences, 297, 300, 301,
304, 306
Common pigeon. See Rock Pigeon
Common platanna. See African Clawed Frog
Common Reed, 447–51
and Giant Reed, 463
impacts, 683
noxious designation, 665, 666, 668, 670,
671, 691
pathways of introduction, 677
uses of, 451
Common saltwort. See Prickly Russian Thistle
Common salvinia, 327, 649, 328, 329
Common St. Johnswort, 358–62
impacts, 678, 679, 680, 682
in medicine, 361
noxious designation, 666, 668, 669, 670,
671, 672
pathways of introduction, 675
Common toadflax. See Yellow Toadflax
Contact herbicides, xx (v. 2)
Convolvulaceae. See morning glory family
Convolvulus ambigens. See Field Bindweed
Convolvulus arvensis. See Field Bindweed
Convolvulus incanus. See Field Bindweed
Coontail, 322, 649
Copal. See Brazilian Peppertree
Copal-tree. See Tree of Heaven
Copper chelate, chemical control (plants)
waterhyacinth, 342
Copper sulfate biological control (plants)
waterhyacinth, 342
Coptotermes formosanus. See Formosan
Subterranean Termite
Coqui, xxvi (v. 1), xxx (v. 1), 208–11
ISSG 100 worst invaders, 210, 711
state-by-state occurrences, 297, 298
736 n INDEX

Coquı́ común. See Coqui Crystalline Ice Plant, 383–87

Coral honeysuckle, 615, 641, 660 impacts, 682
Coralberry, 505, 656 noxious designation, 690
Cordgrass. See Cordgrasses and Their Hybrids pathways of introduction, 677
Cordgrasses and Their Hybrids, 63, 274, 293, Ctenopharyngodon idella. See Grass Carp
452–56 Ctenosaurus similis. See Black spinytail iguana
American species invasive abroad, 696 Cuban brown anole. See Brown Anole
impacts, 682, 683 Cuban nakedwood, 493, 657
noxious designation, 670, 671, 691, 692 Cuban Treefrog, xx (v. 1), xxviii (v. 1), xxxii (v. 1),
pathways of introduction, 675, 676, 677 211–14, 227
Corn bind. See Field Bindweed state-by-state occurrences, 297, 310
Corn thistle. See Canada Thistle Cucullia verbasci. See mullein moth
Cortaderia. See Jubata Grass Cucumber, 619
Cortaderia argentea. See Pampas Grass Culex quinquefasciatus. See Southern house
Cortaderia atacamensis. See Jubata Grass mosquito
Cortaderia dioica. See Pampas Grass Cultivation, physical control (plants). See tilling
Cortaderia jubata. See Jubata Grass Cupania anacardioides. See Carrotwood
Cortaderia selloana. See Pampas Grass Cupania anacardioides var. parvifolia. See
Cosmopolitan bulrush, 455, 654 Carrotwood
Cotton States Exposition (1884), Waterhyacinth Cupaniopsis anacardioides. See Carrotwood
introduction, 340 Currants, White Pine Blister Rust alternate host,
Couch grass. See Quackgrass 30, 32, 34
Council of Europe, and Water Chestnut, 338 Curvulara lunata, biological control (plants)
Cow parsnip. See common parsnip West Indian marsh grass, 492
Coypu. See Nutria Cuscutaceae family. See morning glory family
Crasimorpha infuscate, biological control (plants) Cuscuta japonica. See Japanese Dodder
Brazilian peppertree, 548 Cutleaf toothwart, 371, 650
Crassostrea gigas. See Japanese oyster Cutting or mowing, physical control (plants)
Crater Lake National Park, and white pine brooms, 501
blister rust, 34 cheatgrass, 442
Crawdad. See Rusty Crayfish chocolate vine, 596
Crawfish. See Rusty Crayfish dyer’s woad, 365
Cream lily. See yellow ginger English ivy, 605
Creeper. See Porcelainberry giant reed, 465
Creeping Charlie. See field bindweed; ground ivy gorse, 511
Creeping Jenny. See field bindweed Japanese barberry, 514
Creeping thistle. See Canada Thistle Japanese dodder, 613
Creeping wild rye. See Quackgrass Japanese hops, 621
Crested wheatgrass, 485, 487, 654 Japanese knotweed, 390
Cricotopus myriophylli. See milfoil midge Japanese stilt grass, 468–69
Crimson beauty. See Japanese Knotweed Johnsongrass, 472
Crimson bottlebrush, 558, 657 kudzu, 625
Crimson Fountain Grass, 458–62 mile-a-minute, 628
noxious designation, 667, 691 multiflora rose, 525
pathways of introduction, 673 oriental bittersweet, 632
Crinkleroot, 371, 650 rattlebox, 529
Cristulariella pyramidalis. See zonate leafspot swallow-worts, 639
Cronartium ribicola. See White Pine Blister Rust toadflax, 426
Crustaceans, 86–99 winter creeper, 643
American species invasive abroad, 696 wisteria, 647
Cryphonectria parasitica. See Chestnut Blight yellow starthistle, 430
Fungus Cyanophyllum magnificum. See Velvet Tree
Cryophytum crystallinum. See Crystalline Cygnus olor. See Mute Swan
Ice Plant Cynanchum medium. See Pale Swallow-Wort
 INDEX n 737

Cynanchum rossicum. See Pale Swallow-Wort tamarisk, 585

Cyperaceae. See sedge family
Cyphocleonus achates, biological control (plants)
spotted knapweed, 421
Cypress spurge, 396, 650
Cytisus monspessulana. See French Broom
Cytisus scoparius. See Scotch Broom
Cytisus striatus. See Portuguese Broom
Crytobagous salviniae, biological
control (plants)
giant salvinia, 330
Dalmatian toadflax, 421–26
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 674
Dark-green white-eye, 246
Daubentonia punicea. See Rattlebox
Davis Pond Freshwater Diversion Project
giant salvinia, 329
Dead man’s fingers (green algae), 37
Deer, and leafy spurge, 397
Deer mice, 281, 662
Deer tick, as Lyme disease vector, 4, 5, 6
Deformed wing virus (DWV), bee virus, 105
Dendrobaena octaedra. See European Earthworms
Dendryphiella broussonetiae, biological
control (plants)
paper mulberry, 564
Dengue fever, xix (v. 1), xx (v. 1), 7, 119
Dense-flowered cordgrass, noxious designation,
671, 691
pathways of introduction, 676, 677. See also
Cordgrasses and Their Hybrids
Depressed shrubverbena. See pineland lantana
Devil firefish, 175, 176, 176
Devil pod, 336, 649
Devil’s apple. See cockroach berry
Devil’s fig. See turkey berry
Devils-grass. See Quackgrass
Devil’s guts. See Field Bindweed
Devil’s hair. See Japanese Dodder
Devil’s tail tear-thumb. See Mile-A-Minute
Diatraea succharalis, biological control (plants)
giant reed, 466
Dicamba, chemical control (plants)
Australian pine, 543
gorse, 511
Japanese honeysuckle, 617
multiflora rose, 525
musk thistle, 403
prickly Russian thistle, 413
spotted knapweed, 421
strawberry guava, 578
tropical soda apple, 534
Didemnid. See Colonial Tunicate
Didemnum vexillum. See Colonial Tunicate
Diffuse knapweed, 419, 650
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
Digging out, hand pulling, or bulldozing,
physical control (plants)
Asiatic colubrina, 495
buffelgrass, 438
cordgrasses, 457
dyer’s woad, 364
giant hogweed, 375
ice plant, 386
Japanese barberry, 514
Japanese hops, 621
Japanese stilt grass, 468
Kahili ginger, 394
mile-a-minute, 628
porcelainberry, 635
tamarisk, 584
velvet tree, 592
Diorhabda elongate. See saltcedar leaf beetle
Dioscorea species, 607, 660
Diplodia natalensis, biological control (plants)
Australian pine, 543
Diquat, chemical control (plants)
giant salvinia, 329
Dittander. See Perennial Pepperweed
Dodder, 386. See also Japanese Dodder
Dog grass. See Quackgrass
Dog rose, 523–24, 657
Dog-strangling vine. See Pale Swallow-Wort
Dolichos hirsutus. See Kudzu
Dolichos lobatus. See Kudzu
Dollar leaf plant. See prostrate tickrefoil
Domesticated livestock, xviii (v. 1), xxiv (v. 1)
Dorosoma cepedianum. See Gizzard Shad
Downy brome. See Cheatgrass
Downy chess. See Cheatgrass
Drake, J. A., xxxi (v. 1)
Dreissena polymorpha. See Zebra Mussel
Dreissena rostriformis bugensis. See Quagga mussel
Drooping brome. See Cheatgrass
Drummond rattlebox, 528–29, 657
Dutch Elm Disease Fungi, xix (v. 1), xxvi (v. 1),
21–25, 115, 130
state-by-state occurrences, 295–310
Dutchman’s pipe, 603, 660
Dwarf euonymus. See Winter Creeper
Dwarf gorse, 509, 657
Dwarf honeysuckle. See European fly
honeysuckle
738 n INDEX
Dwarf St. Johnswort, 359, 650
Dyer’s Woad, 362–66
impacts, 678, 680, 682
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 675, 677
uses of, 365
EAB. See Emerald Ash Borer
Early chess. See Cheatgrass
Early saxifrage, 370, 650
Eastern mosquito fish. See Mosquitofish
Ebola, xix (v. 1)
Ecological impacts, xxvi–xxvii (v. 1)
in natural and semi-natural ecosystems,
xxvii–xxix (v. 1). See also species entries
Ecology of Biological Invasions of North American
and Hawaii (Mooney and Drake),
xxxi (v. 1)
“Ecology of Biological Invasions” (ICSU),
xxxi (v. 1)
Ecology of Invasions by Animals and Plants,
The (Elton), xxx (v. 1)
Economic impacts, xxix–xxxi (v. 1).
See also species entries
Edible Periwinkle. See Common
Periwinkle
Eleutherodactylus coqui. See Coqui
Eleutherodactylus planirostris.
See Greenhouse frog
Eggleaf spurge, 396, 650
Eggs and bacon. See Yellow Toadflax
Eichhornia crassipes. See Waterhyacinth
Eichhornia speciosa. See Waterhyacinth
Elaeagnaceae. See oleaster family
Elaeagnus augustifolia. See Russian Olive
Elaeagnus hortensis. See Russian Olive
Elaeagnus iliensis. See Russian Olive
Elaeagnus umbellate. See autumn olive
Elaeodendron fortunei. See Winter Creeper
Eleocharus dulcis, 335
Elephant grass, 459, 460, 654
noxious designation, 691
Elm yellows, 22
Elongate paulownia, 566, 568, 659
Elton, Charles S., xx (v. 1)
Elymus caput-medusae. See Medusahead
Elymus repens. See Quackgrass
Elytrigia repens. See Quackgrass
Elytrigia vaillantiana. See Quackgrass
Emerald Ash Borer, xxvit (v. 1), xxix (v. 1),
127–31
state-by-state occurrences, 299–302, 305,
309, 310
Emerging infectious diseases, xix (v. 1)
Empoasca biguttula, Japanese honeysuckle as
host, 617
Empress tree. See Princess Tree
Enchanted Lake (O’ahu), giant salvinia, 329
Endothall, chemical control (plants)
hydrilla, 334
“English House Sparrow Has Arrived in Death
Valley, The An Experiment in Nature”
(Grinnell), xxxi (v. 1)
English ivy, 602–6
impacts, 684, 685
noxious designation, 670, 671, 693
pathways of introduction, 673, 681
and winter creeper, 641
English Sparrow. See House Sparrow
Epigeic worms, 51
Epirrhoe sepergressa, biological control (plants)
Japanese hops, 621
Episimus utilis, biological control (plants)
Brazilian peppertree, 548
Equus asinus. See Feral Burro
Equus caballus. See Feral Horse
Eriocheir japonicus. See Japanese mitten crab
Eriocheir sinensis. See Chinese Mitten Crab
Erophora cardui, biological control (plants)
Canada thistle control, 348
Erysiphe cichoracearum, biological
control (plants)
common mullein, 357
Erysium alliaria. See Garlic Mustard
Eteobalea species, biological control (plants)
toadflax, 426
Eucerocoris suspectus, biological
control (plants)
melaleuca, 561
Eucalyptus family, 557
Eucerocoris suspectus. See leaf-blotching bug
Euhrychiopsis lecontei, biological
control (plants)
Eurasian watermilfoil, 324–25
Eulalia. See Japanese Stilt Grass
Eulalia viminea. See Japanese Stilt Grass
Eunectes murinus, 220
Eunectes notaeus, 220
Euonymus. See Winter Creeper
Euonymus fortunei. See Winter Creeper
Euonymus species. See Winter Creeper
Euphorbiaceae. See spurge family
Euphorbia esula. See Leafy Spurge
Euphorbia virgata. See Leafy Spurge
Eurasian Collared-Dove, 234–37
state-by-state occurrences, 295, 296,
297–300, 302–8

Eurasian Watermilfoil, 173, 321–26, 334, 336
grass carp, 173, 325
impacts, 680, 681, 682, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671, 689
pathways of introduction, 677
Eurasian wild boar. See Feral Pig
European bindweed. See Field Bindweed
European Earthworms, 48–53
state-by-state occurrences, 296–310
European fly honeysuckle, 504–5, 614, 657
European green crab. See Green Crab
European gypsy moth. See Gypsy Moth
European honey bee, xix (vol 1), 99, 102, 103,
105, 106, 107, 108, 109
European rose chalicid, biological control
(plants)
multiflora rose control, 526
European shore crab. See Green Crab
European Starling, xviii (v. 1), xxiiit (v. 1),
xxx (v. 1), 237–40
ISSG 100 worst invaders, 240, 712
and oriental bittersweet, 631
state-by-state occurrences, 295–310
European swallow-wort. See Pale Swallow-Wort
European Tree Sparrow, 244
European wand loosestrife, 414, 657
European water chestnut. See Water Chestnut
European wild boar. See Feral Pig
Eustenopus villosus, biological control (plants)
yellow starthistle, 431
Evecliptopera decurrens, biological control (plants)
chocolate vine, 596
Everglades
threats from invasive species,
xvi (v. 1), 162, 197, 199, 218, 219,
220, 223, 490
Australian pine, 541, 542–43
Brazilian peppertree, 545, 546
climbing ferns, 599, 600
melaleuca, 558, 559, 560
West Indian marsh grass, 490
Everglade Snail Kite, 661
and waterhyacinth, 342,
and West Indian marshgrass, 491
Everglades National Park, threats from invasive
species, xvi (v. 1), xviii (v. 1)
Asian swamp eel, 162
Burmese python, 218, 219, 220–21
green iguana 223
spotted tilapia, 197
walking catfish, 199
Executive Order 13112 (February 3, 1999),
xiii (v. 1), xv (v. 1), 703
INDEX n 739
Exotic Bush Honeysuckles, 121, 502–8
impacts, 681, 683
and Japanese honeysuckle, 614
noxious designation, 666, 668, 669,
671, 692
pathways of introduction, 673, 674,
675, 676
Fabaceae. See pea family
Fallopia japonica. See Japanese Knotweed
False peacock fly, biological control (plants)
yellow starthistle, 431
False poinciana. See Rattlebox
Faya bush. See Fire Tree
Fayatree. See Fire Tree
Feathertop, 460, 654
Feathery pennisetum. See missiongrass
Federal Noxious Weed Act, xv (v. 1), 701
Felis silvestris catus. See Feral Cat
Feral Burro, 262–65
state-by-state occurrences, 295, 296, 303,
306–8
Feral Cat, xxivt (v. 1), xxxv (v. 1), 265–68
ISSG 100 worst invaders, 268, 712
state-by-state occurrences, 295–310
Feral Goat, xviii (v. 1), xxiiit (v. 1), 268–71
ISSG 100 worst invaders, 712
state-by-state occurrences, 296, 298
Feral hog/swine, 275
and tropical soda apple, 533
Feral Horse, xxiiit (v. 1), 271–75
state-by-state occurrences, 295, 296, 298,
303–10
Feral Pig, xviii (v. 1), xxiiit (v. 1), 275–81
fire tree, 556
ISSG 100 worst invaders, 280, 712
state-by-state occurrences, 295, 296,
297–300, 302, 303, 305, 306, 307, 308,
309, 310
and strawberry guava, 578
Feral pigeon. See Rock Pigeon
Fergusonina species, biological control (plants)
melaleuca, 561
Ficaria ranunculoides. See Fig Buttercup
Ficaria verna. See Fig Buttercup
Field Bindweed, 606–10
impacts, 678, 679, 684, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 693
pathways of introduction, 673, 675
Field morning glory. See Field Bindweed
Field thistle. See Canada thistle
Fig Buttercup, 366–68
impacts, 682
740 n INDEX

noxious designation, 668, 690 Floating fern. See Giant Salvinia

pathways of introduction, 673 Floating fern family, 326
Figwort family, 421, 422, 565, 719 Floating waterhyacinth. See Waterhyacinth
Fire, physical control (plants). See burning Florida apple snail, 68
Firebush. See Fire Tree Florida elodea. See Hydrilla
Fire hazard Florida holly. See Brazilian Peppertree
brooms, 500 Florida Keys National Marine Sanctuary, 178
climbing ferns, 600 Florida thatch palm, 659
cogongrass, 446 and Asiatic colubrina, 495
common reed, 450 Flowered sage. See Lantana
crimson fountain grass, 462 Fluazifop, chemical control (plants)
giant reed, 465 cogongrass, 447
gorse, 511 jubata grass, 477
Johnsongrass, 472 pampas grass, 477
jubata grass, 477 quackgrass, 488
kikuyugrass, 480 Fluridone, chemical control (plants)
lantana, 521 Eurasian watermilfoil, 324
medusahead, 484 giant salvinia, 329
melaleuca, 560 hydrilla, 334
prickly Russian thistle, 412 Fluroxypry, chemical control (plants)
Russian olive, 572 lantana, 521
Firetree. See Fire Tree Flying carp. See Silver Carp
Fire Tree, 248, 554–57 Forbs, 344–431
ISSG 100 worst invaders, 710 American species invasive abroad, 695
noxious designation, 666, 667, 693 ISSG 100 worst invaders, 710–11
pathways of introduction, 673, 676, noxious designation, 690–91
Fireweed, 414, 651 Formosan Subterranean Termite, xix (v. 1),
Fish, 157–201 xxiv (v. 1), xxvt (v. 1), 131–34
American species invasive ISSG 100 worst invaders, 134, 711
abroad, 697 state-by-state occurrences, 295–98, 300,
ISSG 100 worst invaders, 711 302, 305, 307, 308
Fishhook water flea, 97, 711 Fosamine, chemical control (plants)
Five-leaf akebia. See Chocolate Vine multiflora rose, 525
Five-stamen tamarisk, 579, 580, 581. Fountain grass, impacts, 679, 683
See also Tamarisk noxious designations, 667, 691.
Flannel leaf. See Common Mullein See also Crimson Fountain Grass
Flannel mullein. See Common Mullein Fragrant honeysuckle. See winter honeysuckle
Flaxweed. See Yellow Toadflax French broom, 496, 498, 499
Flies, biological control (plants) noxious designation, 666, 667, 670, 692.
Brazilian peppertree, 548 See also Brooms
Canada thistle, 348 French tamarisk, 579, 580, 582.
climbing ferns, 601 See also Tamarisk
common reed, 451 Freshwater herring, 157
hydrilla, 334 Fringecup, 370, 651
lantana, 521 Frog’s-bit family, 331
melaleuca, 561 FST. See Formosan Subterranean Termite
mile-a-minute, 629 Fuitour’s-grass. See Leafy Spurge
musk thistle, 403 Fund for Animals, animal welfare group, 271
Russian knapweed, 421 Fungi, 11–35
strawberry guava, 578 American species invasive abroad, 697
velvet tree, 592 Fungi, biological control (plants)
waterhyacinth, 342 Asiatic sand sedge, 439
yellow Himalayan raspberry, 539 Australian pine, 543
yellow starthistle, 431 buffelgrass, 439
 INDEX n 741

Canada thistle, 349 Giant African land snail. See Giant African Snail
cheatgrass, 443 Giant African Snail, 64–67
chocolate vine, 596 state-by-state occurrences, 298
cogongrass, 447 Giant air plant, 600, 651
common mullein, 357 Giant Asian dodder. See Japanese Dodder
common reed, 447 Giant cow-parsnip. See Giant Hogweed
cordgrasses, 458 Giant Hogweed, 372–76
dyer’s woad, 365 impacts, 678, 681, 682
Eurasian watermilfoil, 325 noxious designation, 665, 666, 668, 669,
fire tree, 557 670, 671, 690
Japanese hops, 621 pathways of introduction, 673, 676
kikuyugrass, 481 Giant knotweed, 388, 651,
Koster’s curse, 518 noxious designation, 666, 670, 671, 690
kudzu, 625 Giant Reed, 449, 462–66
paper mulberry, 564 impacts, 679, 681, 683, 686
silk tree, 575 ISSG 100 worst invaders, 710
tree of heaven, 589 noxious designation, 671, 691
tropical soda apple, 533 pathways of introduction, 673, 675
velvet tree, 592 uses of, 465. See also Common Reed
waterhyacinth, 342–43 Giant reedgrass. See Common Reed
West Indian marshgrass, 492 Giant Salvinia, 326–30
winter creeper, 643–44 impacts, 681, 682, 685
yellow starthistle, 431 noxious designation, 665, 666, 668, 669,
Furze. See Gorse 670, 671
Fusarium nivale, biological control (plants) pathways of introduction, 674, 677
cheatgrass, 443 Giant tree frog. See Cuban Treefrog
Fusarium oxysporum, biological control (plants) Giant whiteweed. See Perennial Pepperweed
tree of heaven, 589 Giant wild pine, 600, 651
Fusarium oxysporum f. perniciosum. Gill-over-the-ground. See ground ivy
See mimosa wilt Ginger family, 391
Ginger-lily. See white ginger
Galarhoeus esula, biological control (plants). Ginseng family, 602
See Leafy Spurge Gizzard Shad, 164, 165, 168–71
Galerucella species, biological control (plants) state-by-state occurrences, 295, 296, 299,
purple loosestrife, 417 300, 302, 303, 306, 308, 310
water chestnut, 338 Glacier National Park, and white pine blister
Gallerucida bifasciata, biological control (plants) rust, 34
Japanese knotweed, 390 Glassy-Winged Sharpshooter, xxvit (v. 1),
Gallina de palo, 221 134–38
Gambusia affinis. See Mosquitofish state-by-state occurrences, 296
Gambusia holbrooki. See Mosquitofish Globe artichoke, 651
Garlic Mustard, 51, 369–72, 507 and Canada thistle, 348
impacts, 682 Gloger’s Rule, xxxi (v. 1)
noxious designation, 665, 666, 668, 669, Glorybush, 515, 590, 657
670, 671, 690 and Koster’s curse, 515
pathways of introduction, 675, 676 Glut herring. See Alewife
Gateway National Recreation Area, 435 Glyphosate, chemical control (plants)
Genista junceum. See Brooms Asiatic sand sedge, 435
Genista monspessulana. See French Broom Australian pine, 543
Geomyces destructans. See Bat White-Nose Brazilian peppertree, 547
Syndrome Fungus brooms, 501
Georgia bully, 600, 657 buffelgrass, 439
Geraldton carnation weed, 396, 651 Canada thistle, 348
German trout. See Brown Trout carrotwood, 550
742 n INDEX
cheatgrass, 443
chinaberry, 554
Chinese lespedeza, 352
chocolate vine, 596
climbing ferns, 601
cogongrass, 446
common mullein, 356
common reed, 451
English ivy, 605
exotic bush honeysuckles, 506
field bindweed, 609
fig buttercup, 368
fire tree, 557
garlic mustard, 371
giant hogweed, 375
giant reed, 465
giant salvinia, 329
gorse, 511
goutweed, 379
halogeton, 382
ice plant, 386
Japanese barberry, 514
Japanese honeysuckle, 617
Japanese hops, 621
Japanese knotweed, 390
Japanese stiltgrass, 469
Johnsongrass, 472
jubata grass, 477
kikuyugrass, 481
Koster’s curse, 518
kudzu, 625
lantana, 521
leafy spurge, 398
medusahead, 484
melaleuca, 561
mile-a-minute, 628
multiflora rose, 525
musk thistle, 403
Oriental bittersweet, 632
pampas grass, 477
papermulberry, 564
perennial pepperweed, 409
porcelainberry, 635
prickly Russian thistle, 413
princess tree, 568
purple loosestrife, 417
quackgrass, 488
rattlebox, 529
silk tree, 575
swallow-worts, 640
tamarisk, 584
toadflax, 426
tree of heaven, 588
tropical soda apple, 534
waterhyacinth, 342
West Indian marsh grass, 492
winter creeper, 643
wisteria, 647
yellow Himalayan raspberry, 538
Glysine sinensis. See Chinese Wisteria
Goats
and brooms, 501
and fire tree, 557
and gorse, 511
plants toxic to, 360
Gold clam. See Asian clam
Golden Apple Snail, xxiiit (v. 1), 67–70
ISSG 100 worst invaders, 69, 711
state-by-state occurrences, 296, 297, 298, 308
Golden shad. See Alewife
Golden starthistle. See Yellow Starthistle
Golden star tunicate, 40
Golpar, 374
Gooseberries, White Pine Blister Rust alternate
host, 30, 32, 34
Goosefoot family, 379, 409
Gopher plant. See caper spurge
Gopher tortoise, 228, 663
and Australian pine, 543
and cogon grass, 446
Gorse, 508–12
and brooms, 498
impacts, 680, 683
ISSG 100 worst invaders, 710
noxious designation, 666, 667, 670, 671, 692
pathways of introduction, 673, 675
Goutweed, 376–79
impacts, 682
noxious designation, 666, 668, 671, 690
pathways of introduction, 673
Gracula religiosa, 233
Graminoids, 432–92
American species invasive abroad, 696
herbicides, xx (v. 2)
ISSG 100 worst invaders, 710
noxious designation, 691–92
Grape family, 633
Grape honeysuckle, 505, 657
Grape industry threat, Glassy-Winged
Sharpshooter, 137
Grass Carp, 164, 172–75
Hydrilla control, 334
state-by-state occurrences, 296, 300, 302,
307, 308
Grass Carp, biological control (plants)
Eurasian watermilfoil, 325
hydrilla, 334
waterhyacinth, 342
 INDEX n 743

Grass family, 435, 439, 443, 447, 452, 458, Green peach aphid, 663
462, 466, 469, 473, 478, 481, and tropical soda apple as host, 533–34
485, 489 Green sea turtle, 286, 543, 663
Grasshoppers, biological control (plants) Grey worms. See European Earthworms
cheatgrass, 443 Grinnell, Joseph, xxxi (v. 1)
Chinese lespedeza, 353 Ground elder. See Goutweed
Gratiana boliviana, biological control (plants) Ground ivy, 367, 651
tropical soda apple, 534 Gully-bean. See turkey berry
Gray, Asa GWSS. See Glassy-winged Sharpshooter
water chestnut cultivation, 336 Gypsy Moth, xix (v. 1), xxivt (v. 1), xxvt (v. 1),
Gray herring. See Alewife 138–42
Gray thistle. See wavyleaf thistle ISSG 100 worst invaders, 711
Grazing, physical control (plants) state-by-state occurrences, 295–97, 299,
brooms, 501 300, 301, 303, 304, 305, 307–10
giant hogweed, 375 Gymnancyla canela, biological control (plants)
kudzu, 625 prickly Russian thistle, 413
medusahead, 484 Gymnaetron tetrum, biological control (plants)
spotted knapweed, 420 common mullein, 357
tree of heaven, 588 Gynerium argentium. See Pampas Grass
yellow starthistle, 430 Gynerium jubatum. See Jubata Grass
Greater celandine. See celandine
Great Lakes, xxiv (v. 1), xxvii (v. 1), xxix (v. 1), Hack-and-squirt, herbicide application,
xxx (v. 1) xx (v. 2)
alewives, 157, 158, 159 Haghighat, Sahar, exotic bush honeysuckle
Eurasian watermilfoil, 325 study, 507
gizzard shad, 169 Hairy lespedeza. See Chinese Lespedeza
impacts of sea lamprey, 193 Hairy wheatgrass, 487, 654
musk thistle, 402 Hairy whitetop, 406, 651
New Zealand mud snail, 73, 74 noxious designation, 665, 666, 667, 670,
quagga mussel, 76, 77, 78 671, 672, 690
round goby, 187, 189, 190, 193 Haleakala National Park, and Argentine ants,
sea lamprey, 191 110, 112
spiny water flea, 96, 97 Halogeton, 379–83
threat from bighead carp, 165, 166 impacts, 678, 680, 682
tubenose goby, 188 noxious designation, 665, 666, 667, 669,
water chestnut, 335, 336, 337 670, 690
zebra mussel, 82, 83, 86 pathways of introduction, 677
Great shipworm. See Naval Shipworm and Russian thistle, 412
Great Smoky Mountains National Park, and Halogeton glomeratus. See Halogeton
hemlock woolly adelgid, 143, 145 Haloragaceae. See watermilfoil family
and feral pigs, 279 Halloween lady beetle. See Multicolored Asian
Green anaconda, 220 Lady Beetle
Green comet milkweed, and white Hall’s honeysuckle, 614, 660
swallow-wort, 637, 651 Halyomorpha halys. See Brown Marmorated
Green Crab, xxxi (v. 1), 90–92 Stink Bug
state-by-state occurrences, 296, 297, 300, Harmonia axyridis. See Multicolored Asian
301, 304, 305, 306, 307, 309 Lady Beetle
Greenheart. See coffee colubrina Harold L. Lyon Arboretum, velvet tree, 590
Greenhouse frog, 208–9 Harris mud crab, 87
Green Iguana, 221–25 Hart’s tongue fern, 639, 651
state-by-state occurrences, 297, 298, Hawaiian blackberry, 536, 657
308, 310 Hawai’i Volcanoes National Park
Green-lipped mussel. See Asian Green Mussel and feral goats, 270
Green mussel. See Asian Green Mussel and fire tree, 555, 556
744 n INDEX
and kahili ginger, 392, 393, 394
and yellow Himalayan raspberry, 538
Hawkmoth, biological control (plants)
leafy spurge, 398
Heartleaf horsenettle, 532, 651
Heart-podded hoary cress. See Hoary Cress
Hedera helix. See English Ivy
Hedge false bindweed, 607, 660
Hedge garlic. See Garlic Mustard
Hedychium gardnerianum. See Kahili Ginger
Hemlock Woolly Adelgid, xix (v. 1), xxvit (v. 1),
142–45
state-by-state occurrences, 297, 298, 300,
301, 307–9
Hemp family, 618
Hemp sesbania, 528
noxious designation, 665, 690, 692
Herbicide control. See species entries
Herbicides
application methods, xx (v. 2)
major categories, xx (v. 2)
Heracleum mantegazzianum. See Giant Hogweed
Heracleum species. See Giant Hogweed
Herpestes javanicus. See Indian Mongoose
Herpetogramma licarsicalis, biological control
(plants)
kikuyugrass, 481
Heteranthera formosa. See Waterhyacinth
Heterodera sinensis, biological control (plants)
cogongrass, 447
Heteroperreyia hubrichi, biological control
(plants)
Brazilian peppertree, 548
Hexazinone, chemical control (plants)
Brazilian peppertree, 547
buffelgrass, 439
crimson fountain grass, 462
giant salvinia, 329
Highway ice plant. See Ice Plant
Hildebrand, Dr. William, 233
Hill Myna, 233
Himalayan blackberry, 537, 657
noxious designation, 670, 692
pathways of introduction, 676
Himalayan bush clover. See Chinese Lespedeza
Histoplasma capsulatum, Rock Pigeon
as host, 258
HIV, xix (v. 1)
Hive death, honeybee tracheal mite as
cause, 99, 100
Hoary Cress, 404–9
pathways of introduction, 677
“Hogzilla,” Georgia wild pig, 277
Hoh River, Japanese knotweed, 389
Hojo-e, Buddhist captive animal release
ceremony, 165
Holcus halapensis. See Johnsongrass
Hollyhock bindweed. See mallow bindweed
“Hollywood Finches,” 241
Homalodisca vitripennis. See Glassy-Winged
Sharpshooter
Homogenization, of biota, xxviii–xix (v. 1)
Honey bee, 663
and gorse, 511. See also Asian honey bee
Honeybee Tracheal Mite, xix (v. 1), 99–102
state-by-state occurrences, 295–310
Honeycreepers, Hawaii, xxvii (v. 1), 1–3, 217,
247, 261
Honeylocust, 574, 659
Honeysuckle family, 502, 614
Honeysuckle. See Exotic Bush Honeysuckles;
Japanese Honeysuckle
Honeyvine, 637, 660
Hordeum caput-medusae. See Medusahead
Horn nut. See devil pod
Horned water chestnut. See Water Chestnut
Horses, plants toxic to
buffelgrass, 438
common St. Johnswort, 360
field bindweed, 609
Johnsongrass, 472
rattlebox, 529
yellow starthistle, 430
Horsetail tree. See Australian Pine
Hottentot fig. See Ice Plant
House cat. See Feral Cat
House Finch, xiii (v. 1), 240–43
state-by-state occurrences, 295–310
House Mouse, 281–83
ISSG 100 worst invaders, 712
state-by-state occurrences, 295–10
House Myna. See Common Myna
House rat. See Black Rat
House Sparrow, xviii (v. 1), xxiiit (v. 1),
xxxi (v. 1), 1, 238, 241, 243–46
state-by-state occurrences, 295–310
Human factor in species invasions, xxxii (v. 1)
Humboldt Bay, smooth cordgrass
eradication, 458
Humboldt Bay owl’s clover, 457, 651
Humulus japonicus. See Japanese Hops
Humulus scandens. See Japanese Hops
Hyadaphis tatariacae, biological control
(plants)
exotic bush honeysuckles, 507
Hybridization, impacts, xxviii (v. 1)
Hydrellia species, biological control (plants)
hydrilla, 334
 INDEX n 745

Hydrilla, xviii (v. 1), xxx (v. 1), 173, pampas grass, 477
331–35, 336 perennial pepperweed, 409
impacts, 681, 682, 685 silk tree, 575
noxious designation, 665, 66, 667, 668, 669, tamarisk, 584
670, 671, 689 tropical soda apple, 534
pathways of introduction, 674, 677 west Indian marshgrass, 492
Hydrilla verticillata. See Hydrilla yellow Himalayan raspberry, 539
Hydrocharitaceae. See frog’s-bit family Imperata arundinaceae. See Cogongrass
Hyles euphorbiae. See hawkmoth Imperata cylindrica. See Cogongrass
Hylobius transversovittatus, biological control Imperata cylindrica var. major. See Cogongrass
(plants) Indian lilac. See Chinaberry
purple loosestrife, 417 Indian Mongoose, xviii (v. 1), xxiiit (v. 1),
Hymenachne acutigluma. See West Indian xxvi (v. 1), 284–86
Marsh Grass ISSG 100 worst invaders, 285, 712
Hymenachne amplexicaulis. See West Indian state-by-state occurrences, 298, 310
Marsh Grass Indian Myna. See Common Myna
Hypena srigata, biological control (plants) Indian snakewood. See Asiatic Colubrina
lantana, 521 Indian star vine. See Hydrilla
Hypericum perforatum. See Common Indigo snake, 446, 663
St. Johnswort Infectious diseases, xxiiit (v. 1)
Hyperparathyroidism disease, in horses, 438 Influenza, xix (v. 1)
Hypophthalmichthys molitrix. See Silver Carp Injurious animal species, definition, xv (v. 1)
Hypophthalmichthys nobilis. See Bighead Carp Insects, 106–56
American species invasive abroad, 696
‘Inia. See Chinaberry Insects, biological control (plants), xviii (v. 2),
Iberian knapweed, 419, 651 xix (v. 2)
noxious designation, 665, 666, 669, Australian pine, 543
670, 690 Brazilian peppertree, 547–48
Iberian starthistle, 428, 651 brooms, 501–2
Ice Plant, 383–87 Canada thistle, 348–49
impacts, 683 climbing ferns, 601
noxious designation, 690 common mullein, 357
pathways of introduction, 675 common reed, 451
Ice plant scale insects, biological control (plants) common St. Johnswort, 361
ice plant, 386 Eurasian watermilfoil, 324–25
Iguana iguana. See Green Iguana field bindweed, 609
Imazapic, chemical control (plants) fire tree, 557
cheatgrass, 443 garlic mustard, 372
dyer’s woad, 365 giant hogweed, 375
Japanese stilt grass, 469 giant reed, 466
Imazapyr, chemical control (plants) giant salvinia, 330
Brazilian peppertree, 547 gorse, 511
chinaberry, 554 halogeton, 383
climbing ferns, 601 hoary cress, 409
cogongrass, 446 hydrilla, 334
common reed, 447 ice plant, 386
cordgrasses, 457 Japanese hops, 621
exotic bush honeysuckles, 506 Japanese knotweed, 390
hoary cress, 409 kikuyugrass, 481
jubata grass, 477 Koster’s curse, 518
kahili ginger, 394 kudzu, 625
lantana, 521 lantana, 521
mile-a-minute, 628 leafy spurge, 398–99
melaleuca, 561 mile-a-minute, 628–29
746 n INDEX
melaleuca, 561
multi-flora rose, 526
musk thistle, 403
perennial pepperweed, 409
prickly Russian thistle, 413
purple loosestrife, 417
rattlebox, 529
spotted knapweed, 421
strawberry guava, 578–79
swallow-wort, 640
tamarisk, 585
toadflax, 426
tropical soda apple, 534
waterhyacinth, 342–43
west Indian marsh grass, 492
yellow starthistle, 43–31. See also flies,
beetles, butterflies, midges, mites, wasps,
and weevils
Intentional pathways of introduction,
xxiiit–xxivt (v. 1)
Intermediate wheatgrass, 484, 654
International Biosphere Reserve,
Everglades, 560
International Maritime Organization, 708
International Plant Protection Convention
(IPPC), 700, 707
Introduced species, xiv (v. 1)
Invasion process, xx–xxii (v. 1)
Invasion science, xxx–xxxi (v. 1)
Invasive plants
American species abroad, 695–96
impacts of, xviii (v. 2), 678–86
introduction of, xvii–xviii (v. 2), 673–77
management, xix (v. 2)
noxious designation, 665–71, 689–93
organizations and publications concerning,
687–88
problem of, xviii (v. 2)
reproduction/dispersal, xvii (v. 2)
Invasive species
animal problem extent, xviii–xx (v. 1)
definitions, xiii–xv (v. 1)
ecological impacts, xxvi–xxvii (v. 1)
economic impacts, xxix–xxxi (v. 1)
federal legislation and agreements pertaining
to, 699–706
human factor, xxxii (v. 1)
international agreements and conventions
pertaining to, 707–9
natural ecosystem impacts,
xxvii–xxix (v. 1)
pathways of introduction, xxii–xxvi (v. 1)
plant problem extent, xv–xviii (v. 1)
process, xx–xxii (v. 1)
public health and well-being
impacts, xxx (v. 1)
Invasive Species Specialist Group (ISSG)100
worst invasive alien species, 710–12
Invertebrates, 36–156
American species invasive abroad, 696–97
Annelid Worms, 48–53
Bryozoan, 36–38
Cnidarian, 45–48
ISSG 100 worst invaders, 711
Mollusks, 53–86
Tunicates, 39–45
Ioxodes scapularis. See Black-legged tick
Ipomoea species. See pipevine and
Dutchman’s pipe
Irish furze. See Gorse
Irish ivy. See Atlantic ivy
Ironwood. See Australian Pine
Isatis tinctoria. See Dyer’s Woad
Ischnodemus variegatus. See myakka bug
Island applesnail, 67
Italian thistle, 346, 401, 651
noxious designation, 665, 666, 667, 670,
671, 690
Jack-by-the-hedge. See Garlic Mustard
Jacob’s ladder. See Yellow Toadflax
Jacob’s staff. See Common Mullein
Jamaican giant anole, 223
Japanese arrowroot. See Kudzu
Japanese bamboo. See Japanese Knotweed
Japanese barberry, 51, 512–15
impacts, 683
noxious designation, 666, 668, 692
pathways of introduction, 673, 674
Japanese Beetle, xxvit (v. 1), 146–48
state-by-state occurrences, 295–310
Japanese blood grass. See Cogongrass
Japanese brome, 440, 654, 692
Japanese Climbing Fern, 597–602
noxious designation, 665, 667, 693
pathways of introduction, 673
Japanese creeper. See Boston ivy
Japanese Dodder, 610–14
in medicine, 613
impacts, 678, 684, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671
Japanese fleece flower. See Japanese Knotweed
Japanese Honeysuckle, 614–18, 641–42
impacts, 684, 685
noxious designation, 666, 668, 669, 671, 693
pathways of introduction, 673
Japanese hop. See Japanese Hops

Japanese Hops, 618–21
impacts, 681, 684
noxious designation, 666, 668, 693
pathways of introduction, 673, 675
Japanese Knotweed, 387–91
impacts, 681, 682, 685
ISSG 100 worst invaders, 710
noxious designation, 665, 666, 668, 669,
670, 671, 690
pathways of introduction, 673, 675
uses of, 390
Japanese lady beetle. See Multicolored Asian
lady beetle
Japanese millet, biological control (plants)
purple loosestrife, 417
Japanese mitten crab, 87
Japanese mystery snail. See Chinese
Mystery Snail
Japanese oyster, 40
Japanese sedge. See Asiatic Sand Sedge
Japanese Silver-eye, 556
Japanese Stilt Grass, 466–69
impacts, 683
noxious designation, 665, 666,
668, 691
pathways of introduction, 676
Japanese White-Eye, xviii (v. 1), 246–48
and kahili ginger, 393
state-by-state occurrence, 298
and strawberry guava, 578
and velvet tree, 591
Japanese wisteria. See Wisteria
Jenkin’s spire shell. See New Zealand
Mud Snail
Jesup’s milkvetch, 651
and swallow-worts, 639
Johnson, Velma B. (“Wild Horse
Annie”), 273
Johnson grass. See Johnsongrass
Johnsongrass, 469–72
impacts, 678, 679, 680, 681, 683, 685
noxious designation, 667, 668, 670
pathways of introduction, 675
Jointed grass, 467, 654
Jubata Grass, 472–77
impacts, 679, 683, 685
noxious designation, 667, 691
pathways of introduction, 673
Kahila garland lily. See Kahili Ginger
Kahili. See Kahili Ginger
Kahili Ginger, 391–94
impacts, 681, 682
ISSG 100 worst invaders, 710
INDEX n 747
noxious designation, 690
pathways of introduction, 673
Kariba weed. See Giant Salvinia
Karritree. See Princess Tree
Kelp destruction, Lacy Crust Bryozoan, 37
Kerr, Dr. Warwick, 106
Key West quail-dove, 285
Kiger mustang, 272
Kikuyugrass, 462, 478–81
impacts, 678, 679, 680
noxious designation, 665, 666, 668, 669,
670, 671, 690
pathways of introduction, 675
Killer bee. See Africanized Honey Bee
King, Helen Dean, 289
Klamath weed. See Common St. Johnswort
Knight anole, 223
Knobbed whelk, 80
Koa tree, 659
and fire tree, 556
and strawberry guava, 577
and velvet tree, 590
Kochia. See burning bush
Koi kandy. See Giant Salvinia
Koster’s Curse, 209, 515–18, 589
impacts, 683
ISSG 100 worst invaders, 710
noxious designation, 667, 692
pathways of introduction, 673, 674, 677
and velvet tree, 589
Kraunhia floribunda. See Japanese Wisteria
Kraunhia floribunda var. alba. See Japanese
Wisteria
Kraunhia floribunda var. sinensis. See Chinese
Wisteria
Kraunhia japonica. See Japanese Wisteria
Kraunhia sinensis. See Chinese Wisteria
Kudzu, 622–26
impacts, 684
ISSG 100 worst invaders, 710
and Japanese dodder, 611, 612
noxious designation, 666, 667, 668, 670,
671, 693
pathways of introduction, 674, 675
uses of, 624, 625
Kyack. See Alewife
Kyasuma grass, 459, 461, 654
noxious designation, 665, 666, 668, 669,
670, 671, 691
Lace bug, biological control (plants)
Canada thistle, 349
lantana, 521
Lacey Act, xvi (v. 1), 196, 200, 319, 699, 704
748 n INDEX
Lacy Crust Bryozoan, xv (v. 1), xxiv (v. 1),
36–38
state-by-state occurrences, 297, 300, 301,
303, 306
Lagurus cylindricus. See Cogongrass
Lake lamprey. See Sea Lamprey
Lamprey eel. See Sea Lamprey
Land invertebrate, ISSG 100 worst invaders, 711
Lantana, 234, 248, 518–22
impacts, 678, 679, 680, 681, 683, 685
ISSG 100 worst invaders, 710
noxious designation, 692
pathways of introduction, 674
uses of, 522
Lantana aculeate. See Lantana
Lantana camara. See Lantana
Lantana camara var. aculeata. See Lantana
Lantana camara var. nivea. See Lantana
Lantana wildtype. See Lantana
Lardizabala family, 594
Lardizabalaceae. See lardizabala family
Large St. Johnswort, 358, 651
Large-headed sedge, 432, 654
Largeleaf lantana. See Lantana
Larinus curtus, biological control (plants)
yellow starthistle, 431
Larinus minutus, biological control (plants)
spotted knapweed, 421
Larinus planus, biological control (plants)
Canada thistle control, 348
Latherleaf. See Asiatic Colubrina
Laycock, George, xxxi (v. 1)
Leaf-blotching bug, biological control (plants)
melaleuca, 561
Leaf rust, biological control (plants)
pepperweed, 409
Leaf worms. See European Earthworms
Leafy Spurge, 395–99
impacts, 678, 680, 681, 682
ISSG 100 invasive worst, 710
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 677
Least Bell’s Vireo, 662
and giant reeds, 464
and Japanese dodder, 612
Least Tern, 255, 662
and Asiatic sand sedge, 434
Legless red mealybug, biological control (plants)
common reed, 451
Legume family, 527. See also pea family
Leiopython albertisii, 220
Lens-pod hoary cress, 405, 651
noxious designation, 690
Lepidium draba. See Hoary Cress
Lepidium latifolium. See Perennial Pepperweed
Leptinotarsa texana, biological control (plants)
tropical soda apple, 534
Leptospirosis, Norway Rat as host, 290
Lespedeza cuneata. See Chinese Lespedeza
Lespedeza juncea var. sericea. See Chinese
Lespedeza
Lespedeza sericea. See Chinese Lespedeza
Lespedeza webworm, Chinese lespedeza
control, 353
Lesser celadine. See Fig Buttercup
Leucantha solstitialis. See Yellow Starthistle
Leucoptera spartifolilella, biological control
(plants)
brooms, 501
Linaria dalmatica ssp. dalmatica. See Dalmatian
Toadflax
Linaria genistifolia ssp. dalmatica. See Dalmatian
Toadflax
Linaria vulgaris. See Yellow Toadflax
Linepithema humile. See Argentine Ant
Linnet. See House Finch
Lionfish, xxiiit (v. 1), 175–78
state-by-state occurrences, 297,
298, 305, 307
Liothrips urichi, biological control (plants)
Koster’s curse, 518
Lipara species, biological control (plants)
common reed, 451
Lithobates catesbeianus. See American Bullfrog
Lithracus atronotatus, biological control (plants)
Brazilian peppertree, 548
Little, Clarence Cook, 282
Littleleaf sensitive briar, 574, 660
Littoraria irrorata. See marsh periwinkle
Littorina littorea. See Common Periwinkle
Littorina saxatilis. See Rough Periwinkle
Lius peisodon, biological control (plants)
Koster’s curse, 518
Lumbricus rubellus. See European Earthworms
Lumbricus terretris. See European Earthworms
Lonicera insularis. See Exotic Bush Honeysuckles
Lonicera japonica. See Japanese Honeysuckle
Lonicera maackii. See Amur Honeysuckle
Lonicera morrowiii. See Morrow’s Honeysuckle
Lonicera sibirica. See Exotic Bush Honeysuckles
Lonicera tatarica. See Tatarian honeysuckle
Lonicera x bella. See Bell’s Honeysuckle
Lonicera japonica. See Japanese Honeysuckle
Loosestrife family, 414
Lophyrotoma zonalis, biological control (plants)
melaleuca, 561
Louis’ swallow-wort. See Black Swallow-Wort

Love-apple. See cockroach berry
Lygodiaceae. See climbing fern family
Lygodium japonicum. See Japanese
Climbing Fern
Lygodium microphyllum. See Old World
Climbing Fern
Lygodium scandens. See Old World
Climbing Fern
Lymantria dispar. See Gypsy Moth
Lymantria xylina, biological control (plants)
Australian pine, 543
Lyme Disease Bacterium, xix (v. 1),
xxx (v. 1), 3–7
state-by-state occurrences, 296, 298, 299,
300. 301, 304, 306–10
Lythraceae. See loosestrife family
Lythrum salicaria. See Purple Loosestrife
MacArthur, Robert, xxxi (v. 1)
Maccartney rose, 523, 657
Machineel, 495, 659
Madwoman’s milk, 396, 651
Magainins, African Clawed Frog, 204
Magur. See Walking Catfish
Mahogany family, 551
Mahogany flat. See Common Bed Bug
Mallow bindweed, 607, 661
Malta starthistle, 427, 651
noxious designation, 669, 690
Mammals, 259–94
American species invasive abroad, 697
ISSG 100 worst invaders, 712
Mangosteen family, 358
Mangroves, 548, 550
Marine vomit. See Colonial Tunicate
Marlahan mustard. See Dyer’s Woad
Marsh marigold, 367, 651
Marsh periwinkle, biological control (plants)
cordgrass, 458
Mecinnus janthirus, biological control (plants)
toadflax, 426
Medusae, jellyfish generation, 46, 47
Medusahead, 481–84
and cheatgrass, 442,
impacts, 679, 680, 683, 691
noxious designation, 666, 670, 671
pathways of introduction, 677
Medusahead wildrye. See Medusahead
Megamelus species, biological control
(plants)
waterhyacinth, 342
Megastigmus aculeatus var. nigroflavus,
biological control (plants)
multiflora rose, 526
INDEX n 749
Megastigmus transvaalensis, biological control
(plants)
Brazilian peppertree, 548
Me-jiro. See Japanese White-Eye
Melaleuca, 557–61
impacts, 680, 681, 684
ISSG 100 worst invaders, 710
noxious designation, 665, 666, 668, 669,
670, 671, 692
pathways of introduction, 674, 675, 666
Melaleuca quinquenervia. See Melaleuca
Melaleuca species. See Melaleuca
Melastoma elegans. See Koster’s Curse
Melastoma hirta. See Koster’s Curse
Melastoma hirtum. See Koster’s Curse
Melastomataceae. See melastome family
Melastome family, 515, 589
Meliaceae. See mahogany family
Melia species. See Chinaberry
Melia azedarach. See Chinaberry
Melopsitticus undulates. See Budgerigar
Membranipora membranacea. See Lacy Crust
Bryozoan
Mesembryanthemum crystallinum. See Crystalline
Ice Plant
Mesembryanthemum edule. See Ice Plant
Metriona elatior, biological control (plants)
tropical soda apple, 534
Metrosideros quinquenervia. See Melaleuca
Metsulfuron, chemical control (plants)
climbing ferns, 601
common St. Johnswort, 361
dyer’s woad, 365
halogeton, 382
hoary cress, 409
Japanese hops, 621
yellow Himalayan raspberry, 539
Metzneria paucipunctella, biological control (plants)
spotted knapweed, 421
Mexican bamboo. See Japanese knotweed
Mexican garter snake, 207
Mexican water-fern, 327, 649
Miconia. See Velvet Tree
Miconia calvescens. See Velvet Tree
Miconia magnifica. See Velvet Tree
Microorganisms, 1–10
ISSG 100 worst invaders, 710
Microstegium imberbe. See Japanese Stilt Grass
Microstegium vimineum. See Japanese Stilt Grass
Midges, biological control (plants)
cogongrass, 447
common reed, 451
Eurasian watermilfoil, 324, 325
leafy spurge, 399
750 n INDEX

Migratory Bird Treaty Act (2001), 255 Mosquitos

Mikania micrantha, 626, 710 Asian Tiger, 116–20
Mile-A-Minute, 626–29 as avian malaria vectors, 1, 3
impacts, 678, 684, 685 as tularemia vector, 290
noxious designation, 665, 666, 669, 670, as West Nile virus vector, 8, 9, 119
693. See also Mikania micrantha Yellow fever mosquito, 117
Mile-a-minute weed. See Mile-A-Minute Moths, biological control (plants)
Milfoil midge, biological control (plants) Australian pine, 543
Eurasian watermilfoil, 324 bindweed, 609
Military grass. See Cheatgrass Brazilian peppertree, 547–48
Milk thistle, 346, 651 brooms, 501
Milkweed family, 636 chocolate vine, 596
Millettia japonica. See Japanese Wisteria climbing ferns, 601
Mimosa. See Silk Tree common mullein, 357
Mimosa arborea. See Silk Tree common reed, 451
Mimosa julibrissin. See Silk Tree common St. Johnswort, 361
Mimosa wilt, biological control (plants) Eurasian watermilfoil, 324–25
silk tree, 575 field bindweed, 609
Missiongrass, 459–60, 655 fire tree, 557
Mites, biological control (plants) giant reed, 466
climbing ferns, 601 gorse, 511
field bindweed, 609 halogeton, 383
gorse, 511 hydrilla, 334
multiflora rose, 526 Japanese hops, 621
prickly Russian thistle, 413 Koster’s curse, 518
waterhyacinth, 342 leafy spurge, 398
Mollusks, 53–86 mile-a-minute, 629
American species invasive abroad, 696 prickly Russian thistle, 413
Mompha trithalama, biological control Russian knapweed, 421
(plants) sawtooth blackberry, 539
Koster’s curse, 518 spotted knapweed, 421
Monarch butterflies, 663 toadflax, 426
and Asiatic sand sedge, 434 velvet tree, 592
and swallow-worts, 639 waterhyacinth, 342
Money monitor. See Nile Monitor Moth mullein, 354, 651
Monk Parakeet, xxvt (v. 1), xxix (v. 1), Mowing. See cutting
248–52 Mud shad. See Gizzard Shad
state-by-state occurrences, 295, 297–300, Mulberry family, 562
304–8, 310 Mulching, tarping, or solarization, physical
Monogynella japonica. See Japanese Dodder control (plants)
Monopterus albus. See Asian Swamp Eel chocolate vine, 596
Montana Dyer’s Woad Cooperative Project, 365 common reed, 451
Mooney, H. A., xxxi (v. 1) cordgrasses, 457
Moraceae. See mulberry family English ivy, 605
Morella faya. See Fire Tree field bindweed, 609
Morning glory family, 606 goutweed, 379
Morrow’s honeysuckle, noxious designation, Japanese knotweed, 390
666, 668, 669, 671, 692 Johnsongrass, 472
and Japanese honeysuckle, 614. See also kikuyugrass, 481
Exotic Bush Honeysuckles West Indian marsh grass, 491
Mosquito fern, 327, 329, 650 winter creeper, 643
Mosquitofish, xxiiit (v. 1), 178–82 Mulhaden. See Alewife
ISSG 100 worst invaders, 181, 711 Mullein moth, biological control (plants)
state-by-state occurrences, 295–306, 307–10 mullein, 347

Multicolored Asian Lady Beetle, xxiiit (v. 1),
xxx (v. 1), 148–52
state-by-state occurrences, 307
Multiflora Rose, 522–26
impacts, 678, 680, 683, 685
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 692
pathways of introduction, 674, 675, 676
Musk Thistle, 345, 348, 399–404
impacts, 680, 683
noxious designation, 666, 667, 668, 669,
670, 671, 672, 690
pathways of introduction, 677
Mus musculus. See House Mouse
Mustang. See Feral Horse
Mustard family, 362, 369, 404
Mute Swan, 252–56
state-by-state occurrences, 297, 300, 301,
304, 305, 309
Myakka bug, biological control (plants)
West Indian marsh grass, 492
Myiopsitta monachus. See Monk Parakeet
Myocastor coypus. See Nutria
Mycoplasmal conjunctivitis
House Finch decline, 241–42
spread, 243
Mylopharyngodon piceus. See Black Carp
Myrica faya. See Fire Tree
Myricaceae. See sweet gale family
Myriophyllum spicatum. See Eurasian
Watermilfoil
Myriothecium verrucaris, biological control
(plants)
kudzu, 625
Myrtaceae. See eucalyptus family; myrtle
family
Myrtle family, 557, 576
Mytella charruana. See Charru mussel
Myxobolus cerebralis. See Whirling Disease
Nanday Conure (parakeet), 250
Nandayus nenday. See Nanday Conure
Nanny shad. See Gizzard Shad
Nanophyte species, biological control (plants)
water chestnut, 338
Nasturtium draba. See Hoary Cress
National Gypsy Moth Slow the Spread (STS)
program, 142
National Invasive Species Act, 703
Native species, definitions, xiii (v. 1)
Natural selection, and brown anoles, 217
Naval Shipworm, 70–72
Neochetina species, biological control (plants)
waterhyacinth, 342
INDEX n 751
Neodiplogrammus quadrivittatus. See sesbania
stem borer
Neogobius melanostomus. See Round Goby
Neomusotima conspurcatalis, biological control
(plants)
climbing ferns, 601
Neomusotima fuscolinealis, biological control
(plants)
climbing ferns, 601
Nepalese browntop. See Japanese Stilt Grass
New Zealand Mud Snail, xxvt (v. 1), 73–76
state-by-state occurrences, 295, 296, 298,
302, 303, 306, 308–10
Nicosulfuron, chemical control (plants)
quackgrass, 488
Nightcrawler, xxiiit (v. 1), xxxv (v. 1). See also
European Earthworms
Nigropalallidal encephalomalacia disease, in
horses, 430
Nigua. See Koster’s Curse
Nile Monitor, xxivt (v. 1), 225–28
state-by-state occurrences, 297
Nintooa japonica. See Japanese Honeysuckle
Niphograpta albiguttalis. See waterhyacinth moth
Nitrogen fixer
Australian pine, 541
brooms, 497, 500
fire tree, 556
gorse, 509–10
kudzu, 623, 627
Russian olive, 571–72
silk tree, 575
Nodding thistle. See Musk Thistle
Nonindigenous Aquatic Nuisance Prevention
and Control Act (1990), xv (v. 1),
85, 702
Nonnative species, xv (v. 1)
Norops sagrei. See Brown Anole
Norops sagrei sagrei, 214
Norops sagrei ordinates, 214
North Cascades National Park, and white pine
blister rust, 34
Northern African python, 220
Northern pearly eye butterfly, and Japanese stilt
grass, 468, 664
Northern Snakehead, xix (v. 1), xxiiit (v. 1),
182–85
state-by-state occurrences, 301, 306, 309
Northern watermilfoil, 321–22, 324, 325, 649
Northern wheatgrass, 487, 655
Norway Rat, xviii (v. 1), xxvit (v. 1), 259, 260,
262, 283, 287–90
state-by-state occurrences, 295–310
Noxious species, definition, xv (v. 1)
752 n INDEX
Noxious Weed Control and Eradication Act,
704–5
Nutria, xviii (v. 1), xxiiit (v. 1), xxvi (v. 1),
290–94
ISSG 100 worst invaders, 293, 712
state-by-state occurrences, 295– 298, 300–2,
304–6, 307, 308, 309
Oberea erythrocephala, biological control (plants)
leafy spurge, 398
Ohia tree, 556, 659
and strawberry guava, 577, 579
and velvet tree, 591
Old World Climbing Fern, 597–602
impacts, 684
noxious designation, 665, 667, 693
pathways of introduction, 674, 679
Octagonal-tail earthworm. See European
Earthworms
Oleaster. See Russian Olive
Oleaster family, 568
Oncideres cingulata. See twig girdler
Onopordum acanthium. See Scotch thistle
Oncorhynchus mykiss. See Rainbow Trout
Ophideres fullonica, biological control (plants)
chocolate vine, 596
Ophioglossum japonicum. See Japanese
Climbing Fern
Ophiomyia lantanae, biological control (plants)
lantana, 521
Ophiostoma novo-ulmi. See Dutch Elm Disease
Ophiostoma ulmi. See Dutch Elm Disease
Oppositeleaf Russian thistle, 412, 651
noxious designation, 666, 690
Orangeberry nightshade, 532, 661
Orange sheath tunicate. See Chain tunicate
Orchids, 495, 661
Orconectes rusticus. See Rusty Crayfish
Orconectes virilis. See Virile Crayfish
Organ Pipe Cactus National Monument
buffelgrass, 437, 438
Oriental Bittersweet, 629–33
impacts, 681, 684
noxious designation, 666, 668,
669, 671, 693
pathways of introduction, 674
and winter creeper, 641
Orseolia javanica, biological control (plants)
cogongrass, 447
Orthogalumna terebrantis, biological control
(plants)
waterhyacinth, 342
Osteopilus septentrionalis. See Cuban Treefrog
Ossabaw Island hog, 280
Oxyops vitiosa, biological control (plants)
melaleuca, 561
Oyster thief (green algae), 37
Pacific mosquito fern, 327, 649
Painted butterfly, biological control (plants)
Canada thistle, 348
Pale Swallow-Wort, 636–40
impacts, 678, 680, 684
noxious designation, 666, 668, 669,
671, 693
pathways of introduction, 674
Palila, and gorse, 510, 662
Pampas Grass, 460, 473–78
impacts, 679, 683, 685
noxious designation, 691
pathways of introduction, 674
Panicum amplexicaule. See West Indian Marsh
Grass
Paper Mulberry, 562–65
impacts, 681, 684
noxious designation, 693
pathways of introduction, 674
Paperbark tea tree. See Melaleuca
Paperbark tree. See Melaleuca
Papyrius papyriferus. See Paper Mulberry
Parapoynx diminutalis, biological control (plants)
hydrilla, 334
Paraquat, chemical control (plants)
cheatgrass, 443
medusahead, 484
Parrotfeather, 322, 649
noxious designation, 665, 666,
668, 671, 689
Partridge pea, 574, 651
Pasilla. See Chinaberry
Passer domesticus. See House Sparrow
Passer montanus, 244
Pasto buffel. See Buffelgrass
Pasture rose, 524, 657
Pathways of introduction, xxii–xxvi (v. 1)
intentional, xxiiit–xxivt (v. 1)
unintentional, xxiiit (v. 1)
Paulownia. See Princess Tree
Paulownia elongata. See elongate paulownia
Paulownia fortunei. See white-flowered
paulownia
Paulownia imperialis. See Princess Tree
Paulownia tomentosa. See Princess Tree
Peacock cichlid, 198
Pea family, 349, 496, 508, 527, 572, 622, 644
Peacock fly, biological control (plants)
yellow starthistle, 431
Pearl millet, 462, 655
 INDEX n 753

Pempelia genistella, biological control (plants) Phragmites. See Common Reed

gorse, 511 Phragmites australis ssp. australis.
Pennisetum ciliare. See Buffelgrass See Common Reed
Pennisetum clandestinum. See Kikuyugrass Phragmites communis. See Common Reed
Pennisetum conchroides. See Buffelgrass Phragmites communis var. berlandieri.
Pennisetum inclusum. See Kikuyugrass See Common Reed
Pennisetum incomptum. See Buffelgrass Phragmites phragmites. See Common Reed
Pennisetum longstylum. See Kikuyugrass Phyllachora species, biological control (plants)
Pennisetum longstylum var. clandestinum. West Indian marsh grass, 492
See Kikuyugrass Phyllocoptes fructiphilus, biological control (plants)
Pennisetum macrostachyon. See Crimson multiflora rose, 526
Fountain Grass Phyllonorycter myricae, biological control
Pennisetum rueppelianum. See Crimson (plants)
Fountain Grass fire tree, 557
Pennisetum ruppelii. See Crimson Fountain Grass Phyllorhiza punctata. See Australian Spotted
Pennisetum setaceum. See Crimson Fountain Jellyfish
Grass Phyllotreta ochripes, biological control (plants)
Peppervine, 633, 634, 660, 661 garlic mustard, 372
noxious designation, 693. See also Phymatototrichum omnivorum, biological
Porcelainberry control (plants)
Pepperweed whitetop. See Hoary Cress common mullein, 357
Perennial peppergrass. See Hoary Cress winter creeper, 643
Perennial Pepperweed 404–9 Physical control (plants). See species entries
impacts, 680, 683 Phytophthora ramorum. See Sudden Oak Death
noxious designation, 665, 666, 667, 668, Piaropus crassipes. See Waterhyacinth
669, 670, 671, 672 Piaropus mesomelas. See Waterhyacinth
pathways of introduction, 677 Pierce’s disease, Glassy-Winged Sharpshooter
Perennial snapdragon. See Yellow Toadflax as vector, 137
Perna viridis. See Asian Green Mussel Pickerelweed, 342, 649
Persian ivy. See colchis ivy Pickerelweed family, 339
Persian lilac. See Chinaberry Picloram, chemical control (plants)
Persicaria arifolium var. perfoliatum. See Australian pine, 543
Mile-A-Minute chinaberry, 554
Persicaria perfoliata. See Mile-A-Minute common St. Johnswort, 361
Peruvian peppertree, 545, 659 gorse, 511
Pet trade release Japanese honeysuckle, 617
boa constrictors, 220 Japanese knotweed, 390
Burmese Python, 219 leafy spurge, 398
Common Myna, 233 multiflora rose, 525
Nile Monitor, 226 musk thistle, 403
Petromyzon marinus. See Sea Lamprey spotted knapweed, 420
Phakopsora apoda, biological control (plants) strawberry guava, 578
kikuyugrass, 481 tropical soda apple, 534
Phalaris setaceum. See Crimson Fountain Grass wisteria, 647
Phomopsis arnoldiae, biological control (plants) Pigface fig. See Ice Plant
Russian olive tree, 572 Piggyback plant, 370, 651
Phomopsis broussonetiae, biological control Pilewort. See Fig Buttercup
(plants) Pili grass, 438, 655
paper mulberry, 564 Pimienta de Brazil. See Brazilian Peppertree
Phothedes dulcis, biological control (plants) Pineapple guava. See Strawberry Guava
giant reed, 466 Pine bark adelgid, 31
Phragmataecia castaneae, biological control Pineland lantana, 519, 657
(plants) Pineland verbena. See pineland lantana
common reed, 451 Pine straw, and climbing ferns, 600, 601
754 n INDEX
Pineus strobi. See Pine bark adelgid
Pineywoods rooter. See Feral Pig
Pink-flowered tamarisk. See Tamarisk
Pink pampas grass. See Jubata Grass
Pink sand verbena, 386, 651
Pink snow mold, biological control (plants)
cheatgrass, 443
Pipevine, 594, 661
Piping Plover, 662
and Asiatic sand sedge, 434
and oriental bittersweet, 632
Plains bristlegrass, 437, 655
Planthopper, biological control (plants)
cordgrass, 458
waterhyacinth, 342
Plant Protection Act, 704
Plasmodium relictum capistranoae. See Avian
Malaria
Platycephala planifrons, biological control
(plants)
common reed, 451
Pleuropterus cuspidatus. See Japanese Knotweed
Pleuropterus zuccarinii. See Japanese Knotweed
Plowing, physical control (plants). See tilling
Plumeless thistle, 401, 652
noxious designation, 665, 666, 667,
668, 669, 670, 671, 672, 691.
See also Musk Thistle
Poaceae. See grass family
Poison hemlock, 374, 652
Poison ivy, 547, 661
and English ivy, 604
and kudzu, 623
Polygonaceae. See buckwheat family
Polygonum cuspidatum. See Japanese Knotweed
Polygonum perfoliatum. See Mile-A-Minute
Polygonum zuccarinii. See Japanese Knotweed
Pomacea canaliculata. See Golden Apple Snail
Pomacea insularum. See Island applesnail
Pondweed, 333–34, 649
Pontederiaceae. See pickerelweed family
Pontederia crassipes. See Waterhyacinth
Poor-man’s mustard. See Garlic Mustard
Popillia japonica. See Japanese Beetle
Porcelain ampelopsis. See Porcelainberry
Porcelainberry, 633–36
and English ivy, 603
impacts, 684
noxious designation, 666, 668, 693
pathways of introduction, 674
Porcelain vine. See Porcelainberry
Portuguese broom, noxious designation, 670,
692. See also Brooms
Postemergent herbicides, xx (v. 2)
Potamopyrgus antipodarum. See New Zealand
Mud Snail
Potato family, 530
Potato x disease, field bindweed as host, 609
Powderpuff tree. See Silk Tree
Proterorhinus marmoratus. See Tubenose goby
Preemergent herbicides, xx (v. 2)
Prickly glasswort. See Prickly Russian Thistle
Prickly lantana. See Lantana
Prickly rose, 524, 657
Prickly Russian Thistle, 409–14
and halogeton, 380
impacts, 678, 679, 683, 685
noxious designation, 665, 666, 667, 669, 691
pathways of introduction, 677
Prickly sage. See Lantana
Pride of India. See Chinaberry
Princess flower. See glorybush
Princess Tree, 121, 565–68
impacts, 684
noxious designation, 666, 693
pathways of introduction, 674, 676
uses of, 567
Prokeleisa marginata. See planthopper
Prostrate tickrefoil, 623, 652
Pseudocercospora humuli, biological control (plants)
Japanese hops, 621
Pseudodaleta unipuncta. See grass army worms
Pseudomonas bacteria, biological control (plants)
cheatgrass, 443
kudzu, 625
Pseudophilothrips ichini, biological control (plants)
Brazilian peppertree, 548
Pseudorabies, Wild Pig as carrier, 280
Psidium cattleianum. See Strawberry Guava
Psidium guajava. See common guava
Psidium littorale var. longipes. See Strawberry
Guava
Psylliodes chalcomera, biological control (plants)
musk thistle, 403
Pterois miles. See devil firefish. See also Lionfish
Pterois volitans. See Lionfish
Pt. Reyes bird’s beak, 457, 652
Public health and well-being impacts, xxx (v. 1)
Australian pine, 543
chinaberry, 553
English ivy, 605
giant hogweed, 375
Japanese hops, 620
Johnsongrass, 472
leafy spurge, 397
melaleuca, 560
paper mulberry, 563
Public Land Management Act of 2009, 706

Puccinia juncea var. solstitialis, biological control
(plants)
yellow starthistle, 431
Puccinia lygodii, biological control (plants)
climbing ferns, 601
Puccinia punctiformis, biological control (plants)
Canada thistle, 349
Pueraria montana. See Kudzu
Pueraria species. See Kudzu
Puerto Rican treefrog. See Coqui
Pulvinariella species. See ice plant scale insects
Punk tree. See Melaleuca
Purple African nightshade, 532, 657
Purpleflowering raspberry, 524, 657
Purple guava. See Strawberry Guava
Purple Loosestrife, 414–17
impacts, 681, 683
ISSG 100 invasive worst, 711
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 691
pathways of introduction, 674, 675, 677
Purple lythrum. See Purple Loosestrife
Purple pampas grass. See Jubata Grass
Purple plague. See Velvet Tree
Purple sesbane. See Rattlebox
Purple starthistle, 419, 428, 652
noxious designation, 665, 666, 669, 670,
671, 691
Purplestem angelica, 373, 652
Purple strawberry guava. See Strawberry Guava
Pyricularia grisea, biological control (plants)
buffelgrass, 439
kikuyugrass, 481
Python molurus bivittatus. See Burmese Python
Python sebae, 220
Quackgrass, 485–89
impacts, 678, 680, 683
noxious designation, 665, 666, 667, 670,
671, 672, 691
pathways of introduction, 677
Quagga Mussel, xxvt (v. 1), 76–79, 83
state-by-state occurrences, 295, 296,
298–303, 305, 306
Quaker Conure. See Monk Parakeet
Quaker Parrot. See Monk Parakeet
Quassia family, 585
Queen Anne’s lace, 373, 652
Rabbit flower. See Yellow Toadflax
Rainbow Trout, xviii (v. 1), xxivt (v. 1),
xxviii (v. 1), 166, 185–87
ISSG 100 worst invaders, 187, 711
state-by-state occurrences, 295–310
INDEX n 755
Rainbow weed. See Purple Loosestrife
Rajania quinata. See Chocolate Vine
Ralstonia solanacearum, biological control (plants)
kahili ginger, 394
tropical soda, 534
Rambler rose. See Multiflora Rose
Ramorum blight. See Sudden Oak Death
Rana platernera. See Cuban Treefrog
Range expansion, xv (v. 1)
Range maps, xiv (v. 1)
Ranunculuaceae. See buttercup family
Ranunculus ficaria. See Fig Buttercup
Rapana venosa. See Veined Rapa Whelk
Rattlebox, 527–30
impacts, 679, 681, 682, 683, 685, 692
noxious designation, 692
Rattus norvegicus. See Norway Rat
Rattus rattus. See Black Rat
Razorback. See Feral Pig
Red baron. See Japanese Blood Grass
Redbelly tilapia, 196
Red-billed Leiothrix, and kahili ginger, 393, 662
and velvet tree, 591
Redcoat. See Common Bed Bug
Red honeysuckle, 505, 657
Red Imported Fire Ant, xxvit (v. 1), xxviii (v. 1),
152–56
ISSG 100 worst invaders, 711
state-by-state occurrences, 295–98, 305,
307–10
Red lionfish. See Lionfish
Red mulberry, 562, 659
Red sage. See Lantana
Red sesbania. See Rattlebox
Red sheath tunicate. See Chain Tunicate
Red starthistle. See purple starthistle
Red swamp crayfish, 89
Red wigglers. See European Earthworms
Red worms. See European Earthworms
Rehsonia floribunda. See Japanese Wisteria
Rehsonia sinensis. See Chinese Wisteria
Reptiles, 214–28
American species invasive abroad, 697
ISSG worst invaders, 712
Residuals, herbicides, xx (v. 2)
Reticulated python, 220
Reynoutria. See Japanese Knotweed
Reynoutria japonica, 387
Rhamnaceae. See buckthorn family
Rhamnus asiatica. See Asiatic Colubrina
Rhamnus splendens. See Asiatic Colubrina
Rhinocyllus conicus, biological control (plants)
Canada thistle, 348
musk thistle, 403
756 n INDEX

Rhinoncomimus latipes, biological control (plants) Running euonymus. See running

mile-a-minute, 628 strawberry bush
Rhinusa species, biological control (plants) Running strawberry bush, 641, 657
toadflax, 426 Rush wheatgrass, 487, 655
Rhizedra lutosa, biological control (plants) Russian knapweed, 418–19, 428, 652
common reed, 451 noxious designation, 665, 666, 667, 668,
Rhizobacteria, biological control (plants) 669, 670, 671, 672, 691
cheatgrass, 443 toxic, 430
Rhus terebinthifolia. See Brazilian Peppertree Russian Olive, 568–72, 705
Rhyssomatus marginatus. See sesbania impacts, 680, 682, 684, 686
seed weevil noxious designation, 666, 669, 693
Rice eel. See Asian Swamp Eel pathways of introduction, 674, 675, 676
Rice-paddy eel. See Asian Swamp Eel Russian thistle, 380. See Prickly Russian Thistle
Richard’s pampas grass. See toe toe Russian tumbleweed. See Prickly Russian Thistle
RIFA. See Red Imported Fire Ant Russian wheatgrass, 487, 655
Ringed Turtle-Dove, 235–36 Russian wild boar. See Feral Pig
Rio Grande ragweed, 438, 652 Rusty blackhaw, 641, 657
River herring. See Alewife Rusty Crayfish, xxiiit (v. 1), 93–95
Robust blackberry. See Yellow Himalayan state-by-state occurrences, 297, 299, 300,
Raspberry 301, 302, 304, 306, 309
Rock Dove. See Rock Pigeon Ryegrass, 487, 655
Rock Pigeon, xviii (v. 1), 256–59
state-by-state occurrences, 295–310 Sacramento Mountain thistle, and musk thistle,
Rocky Mountain National Park, chytrid frog 402, 652
fungus, 19 Safflower, 348, 652
quackgrass, 487 Salicaire. See Purple Loosestrife
Roof rat. See Black Rat Salmo trutta. See Brown Trout
Root rot Salsola species. See Prickly Russian Thistle
and Australian pine, 544 Salsola tragus. See Prickly Russian Thistle
and common mullein, 357 Saltcedar leaf beetle, biological control (plants)
and giant reed, 466 tamarisk, 585
and winter creeper, 644 Salt cedar. See Tamarisk
Rosa cathayensis. See Multiflora Rose Saltlover. See Halogeton
Rosaceae. See rose family Saltmarsh clubrush. See cosmopolitan bulrush
Rosa multiflora. See Multiflora Rose Salt marsh hay. See Cordgrasses and Their
Roseau. See Common Reed Hybrids
Roseau cane. See Common Reed Salt meadow cordgrass. See Cordgrasses and
Rose family, 522, 535 Their Hybrids
Rose stemgirdler, biological control (plants) Saltwort. See Prickly Russian Thistle
multiflora rose, 526 Salvinia. See Giant Salvinia
Round-leaved bittersweet. See Oriental Salvinia auriculata. See Giant Salvinia
Bittersweet Salvinia auriculata complex, 326
Rough Periwinkle, 61 Salviniaceae. See floating fern family
Round Goby, xxiv (v. 1), xxvt (v. 1), xxvii (v. 1), Salvinia molesta. See Giant Salvinia
187–90 Sameodes albiguttalis. See waterhyacinth moth
state-by-state occurrences, 299, San Clemente Island goats, 269, 270
301, 302, 305 Sand dune thistle, 402, 652
Royal paulownia. See Princess Tree Sapindaceae. See soapberry family
Rubus ellipticus. See Yellow Himalayan Sarcotheca bahiensis. See Brazilian Peppertree
Raspberry Sarthamnus scoparius. See Brooms
Rubus flavus. See Yellow Himalayan Raspberry Sassafras, 562, 659
Rubus gowreephul. See Yellow Himalayan Satintail. See Cogongrass. See also Brazilian
Raspberry satintail; California satintail
Rugosa rose, 524, 657 Sauce-alone. See Garlic Mustard
 INDEX n 757

Saussurea species, 402 Sericea bush clover. See Chinese Lespedeza

Sawbelly. See Alewife Sericothrips staphylinus, biological control (plants)
Sawtooth blackberry, 537, 539, 657 gorse, 511
noxious designation, 667, 692 Serrate spurge. See toothed spurge
Scaldweed. See common dodder Sesbania flower beetle, biological control (plants)
Scaly bark oak. See Australian Pine rattlebox, 529
Scarlet wisteria. See Rattlebox Sesbania puniceae. See Rattlebox
Scheiffelin, Eugene, 238 Sesbania seed weevil, biological control (plants)
Schinus. See Brazilian Peppertree rattlebox, 529
Schinus species. See Brazilian Peppertree Sesbania tripetii. See Rattlebox
Schinus terebinthifolius. See Brazilian Peppertree Sesbania stem borer, biological control (plants)
Schistocerca americana, biological control (plants) rattlebox, 529
Chinese lespedeza, 353 Sethoxydim, chemical control (plants)
Schizaphiz graminum, biological control (plants) quackgrass, 488
giant reed, 466 Seven-spotted lady beetle, 149
Schoenobius giganatella, biological control (plants) Sewer rat. See Norway Rat
common reed, 451 Shattercane. See sorghum
Scientific Committee on Problems of Sheep, plants toxic to
Environment (SCOPE), xxxi (v. 1) buffelgrass, 438
Sclerotinia sclerotiorum, biological control (plants) common St. Johnswort, 360
Canada thistle, 349 halogeton, 382
Scolytus multistriatus. See smaller European elm Johnsongrass, 472
bark beetle Shenandoah National Park, and
SCOPE, xxxi (v. 1) gypsy moth, 143
Scotch broom, 509 Ship rat. See Black Rat
noxious designation, 666, 667, 670, 671, Shortspike watermilfoil. See northern
692. See also Brooms watermilfoil
Scotch thistle, 401, 403, 652 Short-tip gall midge, biological control (plants)
noxious designation, 665, 666, 667, 668, leafy spurge, 399
669, 670, 671, 672, 691 Shrubby nightshade, 532, 657
Scrambling nightshade. See Tampico soda apple Shrubby Russian thistle, 412, 657
Scrophulariaceae. See figwort family noxious designation, 665, 666, 667, 668,
Seabeach amaranth, 434, 652 669, 670, 671, 692
Seabeach evening primrose, 434, 652 Shrubs, 493–39
Sea-coast marsh elder, 434, 652 American species invasive abroad, 695
Sea fig, 384, 652. See also Ice Plant ISSG 100 worst invaders, 710–11
Sea Lamprey, xxiv (v. 1), xxvt (v. 1), 158, 183, noxious designation, 692
190–93 Sicilian starthistle, 427, 652
state-by-state occurrences, 299, 301, 302, noxious designation, 665, 666, 691
305, 310 Silk Tree, 572–75
Sea sandwort, 434, 652 impacts, 684
Sea-side arrowgrass, 455, 457, 655 noxious designation, 693,
Seaside knotweed, 434, 652 pathways of introduction, 674, 676
Sedge family, 336, 432 Silky acacia. See Silk Tree
Sedges, 455, 546, 654, 655. See also Asiatic Silky bush clover. See Chinese Lespedeza
Sand Sedge Silver Carp, 164, 194–96
Segmented worms, 48 state-by-state occurrences, 299, 300
Selloa. See Jubata Grass Silverberry, 570, 655
Senna obtusifolia, biological control (plants) Silver buffaloberry, 570, 656
kudzu, 625 Silverleaf nightshade, 532, 657
Septoria hodgesii sp. nov, biological control noxious designation, 665, 666, 667, 669,
(plants) 670, 671, 692
fire tree, 557 Silverleaf whitefly, and tropical soda apple as
Sericea lespedeza. See Chinese Lespedeza host, 533–34
758 n INDEX
Silver pampas grass. See Pampas Grass
Silversword, 356, 657
Silverthorn, 570, 658
Simaroubaceae. See quassia family
Simberloff, Daniel, xxxi (v. 1)
Sipha species. See sugarcane aphids
Sisymbrium alliaria. See Garlic Mustard
Sisymbrium officinalis. See Garlic Mustard
Skipjack. See Gizzard Shad
Slender-flowered thistle, 401, 652
Slender lespedeza, 350–51, 652
Slender perennial peppercress. See Perennial
Pepperweed
Slender Russian thistle, 411, 652
noxious designation, 665, 666, 691
Slender seapurslane, 434, 652
Slippery mullein. See moth mullein
Slow paralysis virus (SPV), bee virus, 105–6
Small Asian mongoose. See Indian Mongoose
Small cordgrass, 452, 655. See also Cordgrasses
and Their Hybrids
Small crowfoot. See Fig Buttercup
Smaller European elm bark beetle, 22, 23, 24
Small-flowered morning glory. See Field
Bindweed
Small-flowered tamarisk. See Tamarisk
Small Indian mongoose. See Indian Mongoose
Small-leaf climbing fern. See Old World
Climbing Fern
Smooth brome, 440, 655
noxious designation, 692
Smooth cordgrass. See Cordgrasses and Their
Hybrids
Smooth cordgrass hybrid. See Cordgrasses and
Their Hybrids
Smooth rose, 524, 658
Snow-on-the-mountain. See Goutweed
Snowpeaks raspberry, 537, 658
noxious designation, 667, 692
Soapberry family, 548
Soap bush. See Koster’s Curse
SOD. See Sudden Oak Death
Sodom apple. See Tropical Soda Apple
Sod-web worm, Canada thistle host, 347
Soft brome, 440, 655
noxious designation, 692
Soft chess. See Cheatgrass
Soil Conservation Service. See U.S. Department
of Agriculture Soil Conservation Service
Solanaceae. See potato family
Solanum chloranthum. See Tropical Soda Apple
Solanum khasianum var. chatterjeeanum.
See Tropical Soda Apple
Solanum viarum. See Tropical Soda Apple
Solanum viridiflorum. See Tropical Soda Apple
Solarization, physical control (plants).
See mulching or tarping
Soldierwood, 493, 658
Solenopsis invicta. See Red Imported Fire Ant
Solenopsis richteri. See Black imported fire ant
Sonoran Desert weedwackers, 438
Sophonia rufostachia. See two-spotted leafhopper
Sorghum, 470, 472, 655
Sorghum halepense. See Johnsongrass
Sorghum miliaceum. See Johnsongrass
Southern house mosquito, as disease vectors, 1,
3, 8, 9
South Texas. See Rio Grande ragweed
Soybean loper, 664,
and tropical soda apple as host, 533–34
Spanish broom, noxious designation, 667, 670,
671, 692. See also Brooms
Spanish gold. See Rattlebox
Spanish mustang, 271–72
Spanish reed. See Giant Reed
Spartina species. See Cordgrasses and Their
Hybrids
Spartium junceum. See Brooms
Spartium scoparium. See Brooms
Speargrass. See Cogongrass
Species Survival Commission (SSC), ISSG 100
worst invasive species, 710–12
Sphenophorus entus vestitus, biological control
(plants)
kikuyugrass, 481
Spiked loosestrife. See Purple Loosestrife
Spike watermilfoil. See Eurasian Watermilfoil
Spiny Water Flea, xxiv (v. 1), 95–99
state-by-state occurrences, 301, 302, 305,
306, 309
Spirochetes bacterium, 3
Spissistulus festinus. See three-cornered alfalfa leaf
hopper
Spotted jellyfish. See Australian Spotted Jellyfish
Spotted knapweed, 417–21
impacts, 679, 680, 683
noxious designatioins, 665, 666, 667, 668,
669, 670, 671, 672, 691
pathways of introduction, 677
Spotted Tilapia, 196–98
state-by-state occurrences, 296, 297, 303.
See also European Starling
Spurge family, 395
Spurgia esulae. See short-tip gall midge
Spurious mullein. See moth mullein
Squarrose knapweed, 419, 652
noxious designation, 665, 666, 670, 671, 691
Staff-tree family, 629

Staff-vine family, 640
Staghorn sumac, 586, 658
Star thistles, 402. See also Yellow Starthistle
St. Augustine grass, 479, 655
St. Barnaby’s thistle. See Yellow Starthistle
Steele, Dr. Allen, 4
Steelhead (coastal rainbow trout), 185, 186
St. Johnswort. See Common St. Johnswort
Stinking shumac. See Tree of Heaven
Stone plant family, 383
Strangleweed. See Japanese Dodder
Strawberry Guava, 279, 576–79
impacts, 678, 682, 684, 686
ISSG 100 worst invaders, 711
noxious designation, 693
pathways of introduction, 674, 676
Streptopelia decaocto. See Eurasian Collared-Dove
Streptopelia risoria. See Ringed Turtle-Dove
Strophocaulos arvensis. See Field Bindweed
Sturnus vulgaris. See European Starling
Sudangrass, 470, 655
noxious designation, 667
Sudden Oak Death, xix (v. 1), 25–29
state-by-state occurrences, 296, 306
Sugarcane aphids, biological control (plants)
kikuyugrass, 481
Sulfometuron, chemical control (plants)
Japanese honeysuckle, 617
Japanese hops, 621
Sumac family, 544
Summer cypress. See burning bush
Sun sensitivity in humans and livestock
common St. Johnswort, 360
giant hogweed, 375
Sun spurge. See madwoman’s milk
Sunflower family, 344, 399, 417, 427
Surfactant, herbicides, xx (v. 2)
Surinam cherry, 659
and strawberry guava, 578
Sus scrofa. See Feral Pig
Susumber. See turkey berry
Swallow-worts, 636–40
impacts, 678, 680, 684
noxious designation, 666, 668,
669, 671, 693
pathways of introduction, 674
Swamp beaver. See Nutria
Swamp dodder. See common dodder
Swamp fly-honeysuckle, 505, 658
Swamp rose, 524, 658
Swamp verbena, 414, 652
Sweetberry honeysuckles, 505, 658
Sweet breath of spring. See winter honeysuckle
Sweetbriar rose, 524, 658
INDEX n 759
Sweet cicely, 370, 653
Sweet gale family, 554
Swine, plants toxic to, 360
Swine brucellosis, Wild Pig as carrier, 280
Synchytrium puerariae, biological control (plants)
kudzu, 625
Systemic herbicides, xx (v. 2)
Taeniatherum asperum. See Medusahead
Taeniatherum caput-medusae. See Medusahead
Taeniatherum crinitum. See Medusahead
Tahiti, 591, 592
Tall wheatgrass, 487, 655
Tall whitetop. See Perennial Pepperweed
Tamaricaceae. See tamarisk family
Tamarisk, 579–85
impacts, 682, 684, 686
ISSG 100 worst invaders, 711
noxious designation, 666, 668, 669, 670,
671, 672, 693
pathways of introduction, 674, 676
Tamarix chinensis. See Five-stamen Tamarisk and
Chinese Tamarisk
Tamarix gallica. See French Tamarisk
Tamarix parviflora. See Small-flowered Tamarisk
Tamarix ramossissima. See Pink-flowered
Tamarisk
Tamarix species. See Tamarisk
Tampico soda apple. See wetlands nightshade
Tanglehead. See pili grass
Taosa species, biological control (plants)
waterhyacinth, 342
Tatarian honeysuckle, noxious designation, 666,
668, 669, 671, 692
and Japanese honeysuckle, 614. See also
Exotic Bush Honeysuckles
Tebuthiuron, chemical control (plants)
buffelgrass, 439
Japanese honeysuckle, 617
Tectococcus ovatus, biological control (plants)
strawberry guava, 578
Teleonemia scrupulosa, biological control (plants)
lantana, 521
Teline monspessulanus. See Brooms
Tender fountain grass. See Crimson Fountain
Grass
Teredo navalis. See Naval Shipworm
Terellia ruficauda, biological control (plants)
Canada thistle, 348
Tetralopha scortealis. See lespedeza webworm
Tetranychus lintearis, biological control (plants)
gorse, 511
Tetranychus urticae, Japanese honeysuckle as
host, 617
760 n INDEX
Texas umbrella chinaberry, 552, 659
Texas umbrella tree. See Texas umbrella
chinaberry
Thai eggplant. See turkey berry
Thatch bromegrass. See Cheatgrass
Theodore Roosevelt National Park, and feral
horses, 271, 272, 273
Theory of Island Biogeography, The (MacArthur
and Wilson), xxxi (v. 1)
Thorny olive. See silverthorn
Three-cornered alfalfa leaf hopper, biological
control (plants)
Chinese lespedeza, 353
Three-leaf akebia, 594, 661
Thrypticus species, biological control (plants)
waterhyacinth, 342
Tick quackgrass, 487, 655
Tilapia, ISSG 100 worst invaders, 711
waterhyacinth control, 342. See also Spotted
Tilapia
Tilapia mariae. See Spotted Tilapia
Tilapia zilli. See Redbelly tilapia
Tilling, physical control (plants)
cheatgrass, 442
cogongrass, 446
dyer’s woad, 365
field bindweed, 609
medusahead, 484
musk thistle, 403
quackgrass, 488
Timandra convectaria, biological control (plants)
mile-a-minute, 629
Timandra griseata, biological control (plants)
mile-a-minute, 629
Tingis ampliata, biological control (plants)
Canada thistle, 349
Tinker’s penny, 359, 653
Tipton weed. See Common St. Johnswort
Tithymalus esula. See Leafy Spurge
Toadflax, 421–26
impacts, 678, 679, 680, 683, 686
noxious designation, 665, 666, 667, 668,
669, 670, 671, 672, 690
pathways of introduction, 674, 675
Tobacco budworm, 664
Japanese honeysuckle as host, 617
tropical soda apple as host, 533–34
Tobacco hornworm, 664
tropical soda apple as host, 533–34
Tobacco leaf curl, biological control (plants)
Japanese honeysuckle, 617
Tobacco mild green mosaic tobamovirus,
biological control (plants)
tropical soda apple, 534
Toe toe, 475, 655
Tomato hornworm, 664
tropical soda apple as host, 533–34
Tomato pinworm, 664
tropical soda apple as host, 533–34
Tomato spotted wilt, field bindweed as host, 609
Tomato weed. See silverleaf nightshade
Toothed spurge, 396, 653
Toothworts, 370, 371, 650, 653
Tordon, chemical control (plants)
gorse, 511
Torrey’s nightshade. See western horsenettle
Toxicodendron altissima. See Tree of Heaven
Toxic plants
brooms, 500
buffelgrass, 438
chinaberry, 553
common St. Johnswort, 360
English ivy, 605
field bindweed, 609
giant hogweed, 375
halogeton, 382
Johnsongrass, 472
kikuyugrass, 480
Koster’s curse, 517
lantana, 521
leafy spurge, 397–98
Oriental bittersweet, 632
rattlebox, 529
toadflax, 425
tropical soda apple, 531
yellow starthistle, 430
Trabutina mannipara, biological control (plants)
tamarisk, 585
Trapa bicornis. See devil’s pod
Trapa bispinosa. See Water Chestnut
Trapaceae. See water caltrop family
Trapa natans. See Water Chestnut
Trapa natans var. bispinosa. See Water Chestnut
Trapa natans var. natans. See Water Chestnut
Trapdoor snail. See Chinese Mystery Snail
Trap-neuter-release (TNR) programs,
feral cats, 268
Tree of Heaven, 248, 585–89
impacts, 684
noxious designation, 666, 668, 669, 671, 693
pathways of introduction, 674
uses of, 588
Trees, 540–93
American species invasive abroad, 695
ISSG 100 worst invaders, 710–11
noxious designation, 692–93
Trichapion latrivetre. See sesbania
flower beetle
 INDEX n 761

Trichomonas gallinae parasite, Eurasian Trumpet creeper, 645, 661

Collared-Dove as host, 236 Trumpet grass. See West Indian Marsh Grass
Trichosirocalus horridus, biological control (plants) Tubenose goby, 188
musk thistle, 403 Tuckeroo tree. See Carrotwood
Trichosporium visiculosum, biological control Tucson Mountain Park, 438
(plants) Tumbleweed. See Prickly Russian Thistle
Australian pine, 543 Tunicates, 39–45
Triclopyr, chemical control (plants) Turkey berry, 531, 658
Asiatic colubrina, 495 noxious designation, 665, 666, 667, 668,
Australian pine, 543 669, 670, 671, 692
Brazilian peppertree, 547 Turkeyfish, 175
brooms, 501 Turtles, and Australian pine, 543
carrotwood, 550 Tu Si Zi. See Japanese Dodder
chinaberry, 554 Twig girdler, biological control (plants)
Chinese lespedeza, 352 Australian pine, 543
chocolate vine, 596 Twinberry honeysuckle. See bearberry
common mullein, 36 honeysuckle
English ivy, 605 Twitchgrass. See Quackgrass
Eurasian watermilfoil, 324 2,4-D, chemical control (plants)
exotic bush honeysuckles, 506 common St. Johnswort, 361
giant hogweed, 375 dyer’s woad, 365
gorse, 511 Eurasian watermilfoil, 324
Japanese barberry, 514 field bindweed, 609
Japanese knotweed, 390 gorse, 511
Koster’s curse, 518 Japanese honeysuckle, 617
kudzu, 625 leafy spurge, 398
multiflora rose, 525 multiflora rose, 525
musk thistle, 403 musk thistle, 403
Oriental bittersweet, 632 prickly Russian thistle, 413
paper mulberry, 564 spotted knapweed, 420
porcelainberry, 635 strawberry guava, 578
prickly Russian thistle, 413 tamarisk, 584
princess tree, 568 velvet tree, 592
purple loosestrife, 417 water chestnut, 338
rattlebox, 529 waterhyacinth, 342
strawberry guava, 578 Twoleaf toothwort. See crinkleroot
swallow-worts, 640 Two-spotted leafhopper, biological control
tamarisk, 585 (plants)
tree of heaven, 588 fire tree, 557
tropical soda apple, 534 Tyta luctuosa, biological control (plants)
velvet tree, 592 field bindweed, 609
winter creeper, 643
wisteria, 647 Ugena microphylla. See Old World Climbing Fern
yellow Himalayan raspberry, 538, 539 Ulex europaeus. See Gorse
Triploid grass carp, 174 Umbrellatree. See Chinaberry
Triticum repens. See Quackgrass Unaspis euonumi, biological control (plants)
Triticum vaillantianum. See Quackgrass winter creeper, 643
Trompetilla. See West Indian Marsh Grass Unintentional pathways of introduction,
Tropical curlygrass fern, 600, 653 xxvt (v. 1), 676–77
Tropical Soda Apple, 530–35 Upland frog. See African Clawed Frog
impacts, 678, 680, 681, 684, 685 Urophora species, biological control (plants)
noxious designation, 665, 666, 668, 669, spotted knapweed, 421
670, 671, 692 yellow starthistle, 431
pathways of introduction, 677 Uruguayan pampas grass. See Pampas Grass
762 n INDEX

U.S. Department of Agriculture Soil Vitis brevipedunculata. See Porcelainberry

Conservation Service, xvi (v. 2), Viviparus georgianus, 59
xviii (v. 2), 437, 499, 524, 623 VOC. See volatile organic compound
Ustilagao bulleta, biological control (plants) Volatile organic compound, and kudzu, 624
cheatgrass, 443
Wahiawa Botanical Garden, and velvet tree, 590
Vaccinium flase bottom, field bindweed Waiawi. See Strawberry Guava
as host, 609 Walking Catfish, xviii (v. 1), 198–201
Vanessa cardui. See painted butterfly ISSG 100 worst invaders, 200, 711
Varanus niloticus. See Nile Monitor state-by-state occurrences, 297
Varanus salvator. See Asian water monitor Wall louse. See Common Bed Bug
Variable leaf waterhyacinth, 339–40, 649 Wasps, biological control (plants)
Variable watermilfoil, 322, 649 Australian pine, 543
noxious designation, 666, 667, 668, Brazilian peppertree, 548
671, 689 multiflora rose, 526
Varnish tree. See Tree of Heaven velvet tree, 592
Varroa destuctor. See Varroa Mite Water caltrop. See Water Chestnut
Varroa Mite, xix (v. 1), 102–5 Water caltrop family, 335
Africanized Honey Bee as host, 103, 108 Water Chestnut, 335–39
state-by-state occurrences, 295–310 impacts, 682, 685
Veined Rapa Whelk, 79–82 noxious designation, 665, 666, 667, 668,
state-by-state occurrences, 308 669, 670, 671, 689
Velvet dock. See Common Mullein pathways of introduction, 674, 677
Velvet Tree, 515, 589–93 Water clover, 327, 330, 650
impacts, 682, 684 Water fern. See Giant Salvinia
and Koster’s curse, 515 Waterhyacinth, xviii (v. 1), 173, 293, 336,
noxious designation, 667, 693 339–43, 522
pathways of introduction, 674 impacts, 680, 681, 682, 685
Verbascum thapsus. See Common Mullein ISSG 100 worst invasives, 710
Verbenaceae. See verbena family noxious designation, 665, 666, 668, 669,
Verbena family, 518 670, 671, 689
Vertebrates, 157–294 pathways of introduction, 674, 677
American species invasive abroad, 697 Waterhyacinth moth, biological control (plants)
amphibians, 201–14 waterhyacinth, 342
birds, 228–59 Water milfoil family, 322
fish, 157–201 Water spangles. See common salvinia
ISSG 100 worst invaders, 711–12 Water straw grass. See West Indian Marsh Grass
mammals, 259–94 Water thyme. See Hydrilla
reptiles, 214–28 Water velvet. See Giant Salvinia
Verticillium dahliae, biological control (plants) Wauke. See Paper Mulberry
tree of heaven, 589 Wavyleaf thistle, 345, 653
Vincetoxicum medium. See Pale Swallow-Wort Weevils, biological control (plants)
Vincetoxicum nigrum. See Black Swallow-Wort Australian pine, 543
Vincetoxicum rossicum. See Pale Swallow-Wort brooms, 501
Vines, 594–644 Canada thistle, 348, 349
ISSG 100 worst invaders, 710–11 common mullein, 357
noxious designation, 693 Eurasian watermilfoil, 324, 325
Violets, 370, 653 garlic mustard, 372
Violet tunicate, 39 giant salvinia, 330
Virginia creeper, 619, 661 gorse, 511
Virginia cutgrass, 467, 656 kudzu, 625
Virginia rose, 524, 658 mile-a-minute, 628–29
Virile crayfish, 93, 94 musk thistle, 403
Vitaceae. See grape family purple loosestrife, 417

rattlebox, 529
tropical soda apple, 534
spotted knapweed, 421
toadflax, 426
tropical soda apple, 534
velvet tree, 592
water chestnut, 338
waterhyacinth, 342
yellow starthistle, 430–31
West African pennisetum. See Kikuyugrass
Western horsenettle, 533, 653
noxious designation, 666, 691
Western morning glory, 607, 661
Western mosquito fish. See Mosquitofish
Western wheatgrass, 442, 487, 656
West Indian mahogany, 495, 659
West Indian Marsh Grass, 489–92
impacts 682, 683
noxious designation, 692
pathways of introduction, 675, 677
West Indian raspberry, 537, 658
noxious designation, 692
West Nile Virus, xix (v. 1), 7–10
mosquito as vector, 119
state-by-state occurrences, 295–310
West Virginia white butterfly, and garlic
mustard, 371
Wetlands nightshade, 531–32, 661
noxious designation, 665, 666, 667, 668,
669, 670, 671, 693
Wetlands soda apple. See wetlands
nightshade
Wharf rat. See Norway Rat
Whin. See Gorse
Whirling disease parasite, salmonids, 187
Whistling pine. See Australian Pine
White amur. See Grass Carp
White avens, 370, 653
White basswood, 562, 659
White bottlebrush tree. See Melaleuca
White cedar. See Chinaberry
White-flowered paulownia, 565–66, 659
White ginger, 391, 392, 393, 394, 653
White grass. See Virginia cutgrass
White herring. See Alewife
White horsenettle. See silverleaf nightshade
White lace bryozoan. See Lacy Crust Bryozoan
White leaf rust, biological control (plants)
perennial pepperweed, 409
White-lipped python, 220
White mulberry, 562, 659
White Pine Blister Rust, 29–35
state-by-state occurrences, 296–310
White ricefield eel. See Asian Swamp Eel
INDEX n 763
White-spotted jellyfish. See Australian
Spotted Jellyfish
White swallow-wort, 637, 661
noxious designation, 671, 693
White-tailed deer, 663
and earthworms, 51
White top. See Hoary Cress
White-winged parakeet, 250
Wild blackberry. See Yellow Himalayan
Raspberry
Wild cane. See Giant Reed
Wild cinnamon, 495, 659
Wild coffee. See Asiatic Colubrina
Wild cucumber, 619, 661
Wild donkey. See Feral Burro
Wild Free-Roaming Horse and Burro Act
(1971), 273, 274
Wild ginger. See Kahili Ginger
Wild grape, 603, 661. See also Porcelainberry
Wild horse. See Feral Horse
Wild oats. See Cheatgrass
Wild parsnip, 373, 653
Wild pigs. See Feral Pig
Wild raspberry. See Yellow Himalayan
Raspberry
Wild rye, 483, 656. See also Quackgrass
Wild snapdragon. See Toadflax
Wilelaiki. See Brazilian Peppertree
Willow Flycatcher, 662
and giant reeds, 464
Wilson, E. O., xxxi (v. 1)
Windwitch. See Prickly Russian Thistle
Winter creeper, 640–44
impacts, 684
noxious designation, 693
pathways of introduction, 674
Winter honeysuckle, 504–5, 658
Winter shad. See Gizzard Shad
Wiregrass. See Quackgrass
Wisteria, 644–47
impacts, 681, 684
noxious designation, 693
pathways of introduction, 674
Wistar Institute, Philadelphia, 289
Wisteria sinensis. See Chinese Wisteria
Wisteria floribunda. See Japanese Wisteria
Witchgrass. See Quackgrass
Witchweed. See Prickly Russian Thistle
WNV. See West Nile Virus
Wolf’s milk. See Leafy Spurge
Wolf’s primrose, 386, 653
Woman’s tongue, 572–73, 660
Woodbine. See Japanese Honeysuckle
Woolly. See Hemlock Woolly Adelgid
764 n INDEX
Wooly Dutchman’s pipe, 594, 661
and climbing ferns, 600
Wooly mullein. See Common Mullein
World Conservation Union (IUCN), ISSG 100
worst invasive alien species, 710–12
Wormwood, 434, 658
Wrinkle wrinkle. See Common Periwinkle
Xenopus laevis. See African Clawed Frog
Xylella fastidiosa, glassy-winged sharpshooter as
vector, 137. See also bacterial leaf scorch
Yellow anaconda, 220
Yellow-brown stink bug. See Brown Marmorated
Stink Bug
Yellow cane. See Common Reed
Yellow-chevroned parakeet, 250
Yellow cockspur. See also Yellow Starthistle
Yellow eel. See Asian Swamp Eel
Yellow fever mosquito, 117
Yellow ginger, 391–92, 393, 394, 653
Yellow Himalayan Raspberry, 535–39
impacts, 684
ISSG 100 worst invaders, 711
noxious designation, 667, 692
pathways of introduction, 676
Yellow honeysuckle, 505, 658
Yellow raspberry. See Yellow Himalayan
Raspberry
Yellow sage. See Lantana
Yellowspine thistle, 345, 653
Yellow Starthistle, 402, 427–31
impacts, 679, 680, 683, 685, 686
noxious designation, 665, 666, 667, 668,
669, 670, 671, 691
pathways of introduction, 677
Yellow toadflax. See Toadflax
Yersinia pestis, carried by black
rats’ fleas, 262, 290
Zacate buffel. See Buffelgrass
Zapata bladderpod, 438, 653
Zasloff, Dr. Michael, 204
Zauclophora pelodes, biological control (plants)
Australian pine, 543
Zebrafish. See Lionfish
Zebra Mussel, xix (v. 1), xxiv (v. 1), xxvt (v. 1),
xxvii (v. 1), xxviii (v. 1), xxix (v. 1), 58,
76, 77, 78, 79, 80, 82–86, 189, 190
ISSG 100 worst invaders, 711
state-by-state occurrences, 295–10
Zingiberaceae. See ginger family
Zonate leafspot, biological control (plants)
tree of heaven, 589
Zooids
bryozoan young, 36, 37
tunicate young, 39–0, 42
Zosterops japonicas. See Japanese White-Eye
Zyginidia guyumi, biological control (plants)
giant reed, 466
n About the Authors
SUSAN L. WOODWARD received her PhD in Geography—with a specialization in biogeog-
raphy—from the University of California, Los Angeles in 1976. Her doctoral research
included three years along the Lower Colorado River studying feral burros, considered by
some then and now to be an invasive species. When her work began, the burro (along with
the feral horse) had just been placed under the jurisdiction of the U.S. Bureau of Land
Management (BLM), which had a federal mandate to manage this living symbol of the Old
West. The results of her field work provided the BLM with some of its earliest baseline data
on burro population biology and ecology.
Dr. Woodward taught biogeography, physical geography, and human ecology for 22 years
at Radford University in Virginia, before retiring in 2006. She is the author of Biomes of Earth
(2003) and served as general editor and author of three volumes for Greenwood Guides to
Biomes of the World (2009).

JOYCE A. QUINN retired from California State University, Fresno as professor emerita after
21 years of teaching a variety of courses in physical geography and mapping techniques. She
earned an MA from the University of Colorado and a PhD from Arizona State University,
both in Geography, specializing in the effect of climate and soils on the distribution of
plants. She has traveled extensively throughout North America, Latin America, Europe,
northern and southern Africa, Uzbekistan, Nepal, China, Southeast Asia, Micronesia, and
elsewhere. She is a member of the Cactus and Succulent Society of America and the
California Invasive Plant Council and is the author of two volumes of Greenwood Guides to
Biomes of the World (2009).