GUNTHER KOHLER

Reptiles of
Central America --~~ .
Impressum

Cover photos:
Front cover, left: Dactyloa microtus (Fortuna, Panama). Photo: S. Lotzkat
Right top: Iguana iguana (Belize). Photo: G. Kohler
Right bottom: Lampropeltis triangulum (Rio Patuca, Honduras). Photo: G. Kohler

Photo p. 4: Cloud forest (Cerro Kilambe, Jinotega, Nicaragua). Photo: G. Kohler

Back cover: Sceloporus teaniocnemis (Chiapas, Mexico). Photo: G. Kohler

Kinosternon angustipons (Nicaragua). Photo: J. Sunyer

Kohler, Gunther
Reptiles of Central America
2nd edition
Offenbach: Herpeton Verlag, 2008
ISBN 3-936180-28-8

Editorial Board: AJ Gutman, West Hartford; James Randy McCranie, Miami;

Peter Stafford, London; Harlan Walley, Dekalb; Larry David Wilson, Miami.

All rights reserved.

No part of this publication may be reproduced or transmitted in any form or by any
means, electronic or mechanical, including photocopy, recording, or any infor-
mation storage and retrieval system, without permission in writing from the
publisher.

No liability is assumed with respect to the use of information contained herein.

© 2008, 2003 Herpeton, Verlag Elke Kohler,

Rohrstr. 22, D-63075 Offenbach, Germany
Photos and drawings: Gunther Kohler, if not otherwise indicated
Layout: Elke Kohler, Offenbach
Contents
Foreword ............................................................ 6 Worm Lizards (Am.phisbaenia) .. 178
Introduction ...................................................... 9
Comments on the Usage of this Book ..... 10
Classification of the Reptiles of
Central America ............................................. 12
The Environment ........................................... 16
Geological History and Zoogeography
of the Reptile Fauna ..................................... 26
Crocodilians (Crocodylia) .................. 30
Turtles (Testudines) ................................ 35
Chelydridae ...................................................... 36
Dermatemydidae ............................................ 37
Emydidae ........................................................... 38
Kinosternidae .................................................. 43
Testudinidae ..................................................... 49
Cheloniidae, Dermochelyidae ................... 50
Lizards (Sauria) ......................................... 57
Anguidae ............................................................ 58
Eublepharidae ................................................. 71
Gekkonidae ....................................................... 72
Gymnophthalmidae ....................................... 84
Helodermatidae ............................................... 93
lguanidae ........................................................... 95
Scincidae ............................................................ 159
Teiidae ................................................................. 166
Xantusiidae ....................................................... 174
Xenosauridae .................................................... 176
Snakes (Serpentes) ............................ 180
Blind Snakes (Scolecophidia) ............ 181
Anomalepididae ......... :............................. 181
Leptotyphlopidae ..................................... 183
Typhlopidae ............................................... 185
Macrostomata ........................................... 186
Boidae ........................................................... 186
Loxocemidae .............................................. 191
Tropidophiidae .......................................... 191
Colubridae .................................................. 193
Elapidae ...................................................... 305
Viperidae ..................................................... 317
Table of Scalation Characters in
Snakes .......................................................... 335
Snakebite - Prevention and
First Aid ...................................................... 342
Mimicry in Snakes ................................. 346
Acknowledgments ................................... 350
Literature Cited ....................................... 352
Glossary ....................................................... 390
Index ............................................................. 393
5
 Foreword
Foreword
Middle America constitutes one of the bio-
diversity hotpots of the world-a region
characterized by a high level of species
richness and endemism. It is understand-
able, thus, that this area, considered to
extend from the Isthmus ofTehuantepec in
Mexico to the southern extent of the
Panamanian isthmus for the purposes of
this book, should have attracted consider-
able interest in this era of biodiversity
decline. The realization among natural
scientists that the Earth's biota is under
attack by forces most humans would
describe as involving simply the business
of living has generated an unprecedented
interest in the cataloguing of the extent of
that biota before it disappears.
The herpetofauna of Middle America, too,
is being studied by a multinational group
of specialists, including herpetologists
from the length and breadth of Mexico and
Central America, the United States, and,
in the case of the present work, Germany.
This study has resulted in the publication
within the last two decades of a number of
faunal works by L. Lee Grismer, Oscar
Flores-Villela and coworkers, Julian C.
Lee, Peter Stafford and John R. Meyer,
Jonathan A. Campbell, James R. McCranie
and coworkers, Josiah Townsend and me,
Gunther Kohler and coworkers, and Jay M.
Savage. Two important changes are occur-
ring in the study of the diverse Middle
American herpetofauna.
One is that more and more herpetologists
raised and educated in the area itself are
undertaking more of the study. The list is
becoming increasingly longer and involves
too many names to mention here. Suffice it
to say, however, that the era of domination
of herpetological studies in the area by for-
eigners is morphing into one increasingly
coming under the influence of native her-
petologists. The other important change is
6
that more and more attention is being
given to questions of the conservation sta-
tus of the members of the highly diverse
Middle American herpetofauna. These two
developments are both highly desirable
and offer the exciting opportunity for
increasing cooperation amo9g groups of
herpetologists from differing backgrounds
in the development of strategies for the
further study and conservation of these
intriguing organisms.
The present new edition is a substantial
and complete revision of an influential pre-
vious edition published in 2003. The fact
that a new edition is called for within only
five years is a testament to the huge
amount of research work that has occurred
in this region in the ensuing period of time.
Not only have new species of reptiles been
described apace, but notable changes have
occurred at higher taxonomic levels as
well. The most significant of these changes
have been at the highest level, necessita-
ting a major reexamination of the meaning
of the word "reptile." Both herpetologists
and ornithologists have been slow to
embrace these changes, but, nonetheless,
cladistic systematic theory requires that
birds be considered the sister group to
modern crocodilians, that crocodilians be
considered eusuchians, and that turtles be
considered members of a separate class
from the lizards, snakes, and amphisbaeni-
ans. This new edition treats 557 species, an
increase of 20 species over the 537 species
of amphisbaenians, crocodilians (eusuchi-
ans), turtles, lizards, and snakes covered in
the 2003 edition. This number will contin-
ue to rise based on studies already in
progress and additional work that will be
undertaken in the years to come. How
many additional reptile species remain to
be discovered in Middle America depends
on the results of continued field and labo-
ratory work, some of which is now under-

way. Nonetheless, it is evident that the
pace of discovery in Middle America
exceeds that occurring in the United
States and Canada. When I wrote the fore-
word to the first edition, I indicated that
the Center for North American Herpe-
tology website listed 592 amphibian and
reptile species for all of the U.S. and
Canada. As I write this, the number now
stands at 598, an increase of only six
species in the last five years! Thus, the rep-
tile diversity of what amounts to an isth-
mus between North and South America is
almost equivalent to all the herpetofaunal
diversity in the two huge nations that
make up the majority of the North
American continent.
This present work continues the publish-
ing tradition begun by Dr. Kohler in 1993
with a book on basilisks of publishing his
major works under the banner of Herpeton
Verlag Elke Kohler, the publishing compa-
ny operated by his spouse. In the last one
and one-half decades, Dr. Kohler has pro-
duced, sometimes alone, sometimes with
coauthors, an impressive series of colorful
books of both scholarly and popular inter-
est on subjects as broad-ranging as reptile
egg incubation, systematics of spiny
lizards (Sceloporus) and spiny-tailed igua-
nas (Ctenosaura), and the systematics and
distribution of the members of the
Nicaraguan herpetofauna, as well as the
present book. In addition, under another
banner, Dr. Kohler has coauthored a full-
Fig. 1. Review Workshop for the Global
Amphibian Workshop at La Selva, Costa Rica,
in November 2002. Photo: G. Kohler
Foreword
scale treatment of the herpetofauna of El
Salvador.
Gunther Kohler is Head of the Department
of Herpetology at the Research Institute
Senckenberg in Frankfurt am Main,
Germany, and Director of the Conservation
Project Utila Iguana, Honduras. He is con-
tinuing a tradition that extends back to the
days of Oskar Bottger and, more recently,
Robert Mertens. Dr. Kohler is continuing
and expanding on the interest Mertens
had in the herpetofauna of Middle
America. Kohler's books on the herpetofau-
nas of El Salvador, Nicaragua, and the Bay
Islands and Cayos Cochinos of Honduras
highlight this interest. Since 1995, when
he was awarded his Ph.D. and began work
at the Research Institute Senckenberg, he
has continued to furbish his considerable
reputation as a major player in the studies
on the herpetofauna of megadiverse
Middle America. This assessment is evi-
denced by the 15 books, 145 popular arti-
cles, and 110 scientific papers on reptiles
and amphibians he has produced to date.
On a more personal note, it continues to be
my pleasure to call Gunther Kohler both
colleague and friend. When I wrote the
foreword for the first edition of this book,
Gunther and I recently had participated in
the Mesoamerican Review Workshop for
the IUCN-supported Global Amphibian
Assessment (GAA), which occurred at the
La Selva Biological Station in eastern
Costa Rica. The results of the GAA have
now been published and have produced an
alarming evaluation of the precarious
environmental status of the world's
amphibians. The GAA, as I write, is being
followed by the Global Reptile Assessment,
with the expectation that reptiles will be
found to be in a similarly threatened po-
sition across the world. Thus, I expect that
Dr. Kohler and I will find ourselves, once
again, at a meeting of herpetologists who
have devoted their careers to the Middle
American herpetofauna at which we will
be considering the probable fate of "our
creatures." As was my impression gained
7
 Foreword
Fig. 2. Norops datzorum (adult male; La
Nevera , Panama). Photo: S. Lotzkat
at the La Selva meeting, I continue to be
impressed with Dr. Kohler's enthusiasm,
dedication, perspicacity, and breadth of
knowledge about the Middle American her-
petofauna, including the reptile segment.
In addition, his academic credentials are
also impressive. It is not many in our field
who hold both Ph.D. and DVM degrees!
The book for which this is the foreword is a
masterful synthesis of information from
not only the previous edition, but also a
broad array of scientific literature that has
appeared both before and after the publi-
cation of its predecessor, producing a
Literature Cited section of 1188 entries.
The species lists and keys for identification
have all been revised and updated and will
prove useful to anyone who has a doubt
about what creature is in hand. The text is
accompanied by professional quality color
photographs of about 466 of the 557
species and expertly done line drawings of
diagnostic features. The percentage repre-
sentation has improved as well, increasing
from 74 in the first edition to 84 in the pre-
sent one, with the representation especi-
ally improving in certain genera such as
Atractus, Omoadiphas, and Trimetopon . In
addition, there are many attractive habitat
photos and generalized distribution maps
that will broaden the reader's understand-
8
ing of the places these creatures inhabit.
The 6 and 1/2 by 9 inch size is also conve-
nient for portage into the field (encased
within a plastic bag, of course).
In my earlier foreword, I wrote that, "Thus,
finally, after more than three decades, we
have a fitting modern treatment of the her-
petofauna of Middle America, which has
been wanting since the classic work by
James L. Peters and his co-workers, cover-
ing only the squamate reptiles, appeared
in 1970 and quickly grew out of date." Now,
Dr. Kohler's first edition of this book has
grown out of date as well, necessitating the
present revised edition, providing a testa-
ment to the continuing work of herpetolo-
gists who have dedicated their professional
lives to advancing our understanding of
the important reservoir ofherpetodiversity
that is Middle America. Given that this
work demonstrates that we still have only
a partial understanding of the extent of
this herpetodiversity, this second edition
will also provide the same sort of snapshot
as did the first, but a snapshot that will be
consulted by students of Middle American
herpetology for years to come. In my own
case, I have carried the previous edition
into remote locations all over Honduras,
the country representing the focus of my
research. I expect to do the same with the
present edition.
This foreword was written while I was in
the field working on a grant from Critical
Ecosystem Partnership Fund to study the
conservation status and management of
the critically endangered amphibian fauna
of Honduras awarded to Josiah H.
Townsend, Max A. Nickerson, and James
D. Austin of the University of Florida.
Larry David Wilson
Tegucigalpa, Honduras
12 September 2008

Introduction
The present book, which is based on
KOHLER (2000a, 2001a, 2003a), attempts
to provide a summary of the widely scat-
tered information on Central American
reptiles. The region that is covered by this
book includes the seven countries that
constitute political Central America
(Belize, Guatemala, El Salvador,
Honduras, Nicaragua, Costa Rica, and
Panama), as well as the southeastern part
of Mexico (the area east of the Isthmus of
Tehuantepec, including the Yucatan
Peninsula; Fig. 3). This part of Mexico
belongs to the zoogeographically impor-
tant Central American landbridge (SAVAGE
1982). Although the Swan Islands belong
politically to Honduras, they are not
included because they are biogeographi-
cally part of the West Indies, instead of
Central America.
The history of herpetological research in
Central America dates back to the early
nineteenth century. Early milestones in
the history of Central American herpetolo-
gy were the "Herpetologia Mexicana" of
Arend F. A. WIEGMANN (1834), the
"Biologia Centrali-Americani" of Albert C.
L. G. GUNTHER (1885-1902) and the
"Mission Scientifique au Mexique et dans
l 'Amerique Centrale" of DUMERIL,
BOCOURT & MOCQUARD (1870-1909). Early
milestones of Central American snake
literature include BOULENGER's Cata-
logues on the turtles, lizards and snakes
in the British Museum (BOULENGER 1885,
1889, 1893, 1894, 1896a). In 1970,
important works such as the two volume
"Catalogue of Neotropical Squamata" of
James A. PETERS and co-workers appeared.
In the interval of about 30 years since
these two publications, a large body of
papers dealing with the various aspects of
Central American reptiles has been
published. This body includes works on
certain genera or larger groups of turtles
Introduction
Fig. 3. The zoogeographic region of Central
America (shaded).
(e.g., ERNST & BARBOUR 1989, IVERSON
1992), lizards (e.g. UZZELL 1965a, 1965b,
ECHTERNACHT 1971, 0FTEDAL 1974, HARRIS
& KLUGE 1984, HARRIS 1994), snakes (e.g.,
DUELLMAN 1958, DOWNS 1967, MYERS
1974, CAMPBELL & LAMAR 1989, CAMPBELL
& BRODIE 1992, ROZE 1996, SAVAGE &
SLOWINSKI 1996, CAMPBELL & LAMAR 2004,
SOLORZANO 2004). Additionally, a number
of regional checklists and compilatory
works (MEYER & WILSON 1973, H . SMITH &
SMITH 1973, 1976, 1977, 1979, SAVAGE &
VILLA 1986, CAMPBELL & VANNINI 1989,
KOHLER 1999c, 200lb, PEREZ SANTOS
1999), as well as thorough monographs of
selected countries, have been published
(TAYLOR 1951, 1954, LEE 1996, CAMPBELL
1998a, STAFFORD & MEYER 2000, SAVAGE
2002, MCCRANIE et al. 2005, 2006, KOHLER
et al. 2006). In 1999, a masterful taxono-
mic reference for the snakes of the world
(Colubridae not treated in tlie first volu-
me) was published (MCDIARMID et al.
1999). I generally follow the taxonomic
arrangement proposed in that work.
Central America is the link between North
and South America and is of great zoogeo-
9
 Introduction

graphic importance because its fauna con- Comments on the Usage of

tains species from both continents, in addi-
tion to a high number of endemic species.
As presently understood, the reptile fauna
of Central America consists of 557 species
representing 153 genera and 29 families
(Table 1). This enormous diversity is due
to the exceptional vegetational variation
found in the region, which makes it a
mosaic of rain forests, pine forests, cloud
forests, dry forests, cactus forests, savan-
nas, swamps, and mangroves. This diversi-
ty, and the fact that some areas have so far
not been scientifically explored, renders
Central America a great challenge for her-
petological research. Much research
remains to be done on the diversity,
systematics, and ecology of Central
American reptiles. Recent discoveries have
necessitated the description of new species
(e.g., STAFFORD 2004, MCCRANIE 2006a,b,
2007, TOWNSEND & WILSON 2006, KOHLER
et al. 2007' KOHLER & SMITH 2008) and
even new genera (CAMPBELL & SMITH 1998,
KOHLER et al. 2001b). However, both scien-
tific investigation and efforts to preserve
biodiversity are in a race against time as
the pace of habitat destruction increases
in Central America.
this Book
The intent of this work is to provide an
identification guide for both scientists and
travelers to the numerous reptile species
found in Central America. Many Central
American species of reptiles are so distinc-
tive in coloration, as well as body size and
shape, that an animal in question often
can readily be identified by photographs or
drawings alone. If the identification remains
questionable by this method, identification
keys should be consulted. Additionally, the
distribution maps can. be included in the
identification process, because many
species have very restricted distributions.
The identification keys included in the
present book consider all families, genera, ·
and species that are known to occur in
Central America, even though this is not
explicitly stated in the respective key hea-
dings. Subspecies are mentioned in the
species accounts, but are not considered in
the identification keys or in the distribution
maps. The keys are dichotomous, meaning
that they provide pairs of contrasting
alternatives. Such a key works using step
by step elimination of alternatives until

Group Families Genera Species

Crocodiles 2 2 3

Turtles 7 14 25

Lizards 10 50 236

Amph isbaenians 1 1 3

Snakes 9 86 290

Total 29 153 557

Table 1. Taxonomic composition of the reptile fauna of Central America.

10

only a single alternative remains. It is
important to remember when using the
identification key that the juveniles of
many lizard and snake species have dif-
ferent coloration and pattern than the
adults. Any key based on distinguishing
coloration will not be able to definitively
identify the juveniles. The technical terms
and morphological character designations
used in the keys can be frustrating to non-
specialists, but I am optimistic that anyone
willing to invest some effort in learning the
terms will be able to use the keys with con-
fidence and success. The glossary included
in the book will be helpful in this regard, as
will the illustrations accompanying the
keys.
Following the introduction to each major
group of Central American reptiles, the fol-
lowing information is provided in the spe-
cies accounts: the current scientific name
with author and year of description, follow-
Fig. 4. Juvenile Enyalioides heterolepis (near Santa Fe, Panama).
Introduction
ed by the citation of the original descrip-
tion and type locality; the maximum size
(carapace length, total length [TL] or
snout-vent length [SVL]), and the known
geographic distribution. For some species,
additional notes on ecology and reproduc-
tion are included. Detailed anatomical
descriptions are beyond the scope of the
book. The recommendations for further
reading at the end of each generic account
serve the reader as a guide to the addition-
al references. The index at the end of the
book will help in finding the pages where
the keys, species accounts, photographs
(bold page numbers), and distribution
maps (italic page numbers) of the respec-
tive species are located. The keys and maps
included in this book are based mostly on
the information provided in the publicati-
ons listed at the end of each generic
account supplemented by my own unpu-
blished data. Drawings by the author if not
indicated otherwise.
Photo: S. Lotzkat
11
 Classification of the Reptiles of Central America

Classification of the Reptiles of Central America

Crocodylia (Crocodiles) Testudines (Turtles)

Dermatemydidae Emydidae
Dermatemys (1) Rhinoclemmys (6)
Terrapene (1) Kinosternon (5)
Trachemys (1) Staurotypus (2)

Cheloniidae Dermochelyidae
Caretta (1) Dermochelys (1)
Chelonia (1)
Eretmochelys (1)
Lepidochelys (2)

The taxonomic levels:

• Class Family
• Order Genus (in parentheses is the number of species
Suborder that occur in Central America)

12
 Classification of the Reptiles of Central America

Squamata (Squamates)

Sauria (Lizards) Amphisbaenia (Worm Lizards) Serpentes (Snakes)

Anguidae Gekkonidae
Abronia (18) Aristelliger ( 1)
Celestus (8) Gehyra (1)
Coloptychon (1) Gonatodes (1)
Diploglossus (3) Hemidactylus (5)
Gerrhonotus (1) Lepidoblepharis (2)
Mesaspis (2) Lepidodactylus (1)
Phyllodactylus (3)
Sphaerodactylus (9)
Thecadactylus (1)

Iguanidae Scincidae Teiidae

Anolis (1) Eumeces (1) Ameiva (6)
Basiliscus (4) Mabuya (1) Aspidoscelis (6)
Corytophanes (3) Mesoscincus (2) Cnemidophorus (1)
Ctenonotus (1) Scincella (1)
Ctenosaura (12) Sphenomorphus (4)
Dactyloa (8)
Enyalioides (1)
Iguana (1)
Norops (77)
Laemanctus (2)
Morunasaurus (1)
Phrynosoma (1)
Polychrus (1)
Sceloporus (18)
Urosaurus (1) Heloderma (1)

13
 Classification of the Reptiles of Central America

Amphisbaenia
(Worm Lizards)
Serpentes (Snakes)

Anomalepididae Typhlopidae
Anomalepis (1) Ramphotyphlops (1)
Helminthophis (1) Typhlops (4)
Liotyphlops (1)

Viperidae
Agkistrodon (1)
Atropoides (2)
Bothriechis (9)
Bothrops (2)
Cerrophidion (2)
Crotalus (1)
VENOMOUSI Lachesis (3)
Porthidium (7)

Macrostomata

Boidae
Boa (1)
Corallus (2)
Epicrates (1)

14
 Classification of the Reptiles of Central America

Colubridae

Colulirinae DipsaClinae Natricinae enoilontinae

Chironius (3) Coniophanes (9) Nerodia (1) Adelphicos (6)
Coluber ( 1) Dipsas (8) Storeria (1) Amastridium (2)
Dendrophidion (4) Imantodes (5) Thamnophis (4) Atractus (5)
Drymarchon (1) Leptodeira (5) Chapinophis (1)
Drymobius (4) Ninia (8) Clelia (3)
Elaphe ( 1) Pliocercus (2) Erythrolamprus (2)
Ficimia (2) Rhadinaea (21) Geophis (19)
Lampropeltis (1) Sibon (11) Hydromorphus (2)
Leptodrymus (1) Trimetopon (6) Family status Liophis (2)
Leptophis (7) Tropidodipsas (3) unclear Manolepis (1)
Masticophis (1) Urotheca (5) Conophis (2) Omoadiphas (2)
Mastigodryas (3) Crisantophis (1) Oxyrhopus ( 1)
Oxybelis (4) Diaphorolepis ( 1) Phimophis (1)
Pituophis ( 1) Enuliophis (1) Pseudoboa (1)
Pseustes ( 1) Enulius (3) Siphlophis ( 1)
Salvadora (1) Nothopsis (1) Tretanorhinus (2)
Senticolis (1) Rhinobothryum (1) Tripanurgos (1)
Symphimus (2) Scolecophis (1) Xenodon (1)
Scaphiodontophis (2)
Spilotes (1)
Stenorrhina (2)
Tantilla (23)
Tantillita (3)
Trimorphodon (2)

The taxonomic levels:

Suborder
D Infraorder
Family
• Subfamily
Genus (in parentheses is the number of species
that occur in Central America)

15
 The Environment

The Environment

The Environment
A chain of roughly 100 volcanoes stretches
along the Pacific coast of Central America
from eastern Chiapas, Mexico, to central
Panama. Of these, about 25 retain signs of
activity. The volcanic peaks are highest in
the north, where two in Guatemala reach
elevations of more than 4000 m . The
various mountain complexes that form the
physiogeography of Central America can
be divided into two major massifs. The
northern block includes the Mesa Central
of Chiapas, the highlands of central and
southern Guatemala, and those of Honduras,
El Salvador, and northern Nicaragua.
These are separated from the southern Fig. 5. Sandy beach on Isla de Utila
highlands by the Nicaraguan Depression. (Honduras). Photo: G. Kohler
The southern highlands include the
Cordilleras de Guanacaste, Central, and
Talamanca. The two large Nicaraguan Sandy beaches
lakes (Lagos de Managua and Nicaragua) Sandy beaches (Fig. 5) extend intermit-
are the largest inland bodies of water in tently along much of both the Pacific and
Central America, with surface areas of the Caribbean coast, especially in bays
1035 km2 and 8157 km2, respectively. The that are protected from the breakers by
diverse landscape of Central America sup- offshore reefs. Commonly, a narrow strip of
ports very different regional climatic coconut palms occurs along the beach.
situations (DUELLMAN 1966, STUART 1966). Dunes support the growth of scrubby xero-
On the Pacific versant of nuclear Central phytic vegetation, such as low shrubs, aga-
America the dry season extends from ves, and matted cacti (Rhipsalis sp.).
January through April. A subhumid corri- Among others, the heliothermic race
dor, made up predominantly of dry forest runners (Cnemidophorus) , spiny-tailed
or the remains thereof, stretches along the iguanas (Ctenosaura), and spiny lizards
Pacific versant of Central America from (Sceloporus) inhabit the sandy beaches.
the Isthmus of Tehuantepec through These exposed sandy habitats are also
northwestern Costa Rica. Generally, the important nesting sites for sea turtles and
Caribbean versant experiences an equa- spiny-tailed iguanas (Ctenosaura).
ble, tropical climate with relatively mesic
conditions throughout the year. Mangroves
Exceptions are the northern part of the Mangroves (Fig. 6) are found along the
Yucatan Peninsula, as well as some interi- coastline at many localities in Central
or valleys, which experience a more arid America. The red mangrove (Rhizophora
climate. The wettest regions of Central mangle) is more salt water tolerant, and
America are found in southeastern usually occurs immediately along the
Nicaragua and northeastern Costa Rica, edges of lagoons and bays. Behind the
which receive an excess of 6000 mm preci- stands of red mangrove, the much taller
pitation per year. black mangrove (Avicennia germinans)
16
 The Environment

prolonged drought periods. Dry forests

consist of a thick ground cover of succu-
lents and drought-resistant shrubs. Tree
heights generally do not exceed 20 m, and
many of the trees are armed with thorns.
A curious aspect of this formation is the
presence of tree cacti, which often reach
heights equal to that of trees (Fig. 7).

Savanna
The savanna that occurs in Central
America includes wet savanna (e.g., on the
Honduran island of Utila) and pine savan-
na ("Sabanas de Pinos Caribe"). Pine

Fig. 6. Mangrove with epiphytes on Isla de

Utila, Honduras. Photo: G. Kohler

and white mangrove (Laguncularia race-

mosa) often form dense swampy forests. A
tangle of aerial and adventitious roots
characterizes this ecosystem. Mangrove
swamps provide habitat for the American
crocodile (Crocodylus acutus), the Utila Fig. 7. Dry forest at Tehuantepec, Mexico
anole (Norops utilensis), and the Utila iguana (400 m elevation). Photo: G. Kohler
(Ctenosaura bakeri).

Dry forest
Dry forests are typical of the Pacific ver-
sant of Central America and a few interior
valleys of Honduras and Guatemala. The
climate is characterized by low precipita-
tion (500 to 1200 mm) that falls primarily
during the rainy season from June
through November. During the dry season,
when precipitation is negligible, a majori-
ty of the trees shed their leaves, permit-
ting sunlight and wind to desiccate the
ground layer. Only the large rivers contain
water throughout the year, whereas smal-
ler streams are intermittent or dry during
the dry season. Plants and animals of this Fig. 8. Pine savanna at Alamikamba,
extreme habitat are well adapted to the Nicaragua (100 m elevation). Photo: G. Kohler
17
 The Environment

savanna is found on the Caribbean ver-

sant of Honduras (from the Laguna Ebano
southeastward) and Nicaragua (from the
Rio Grande de Matagalpa northward).
This habitat is dominated by open stands
of Caribbean pine (Pinus caribaea) that
abruptly change over to broadleaf forest.

Rain forest
Evergreen tropical rain forest is among
the most diverse ecosystems on the planet,
and in many regions its scientific explora-
tion is still in the beginning stages. In the
northern parts of Central America, rain
forest is found mainly in the Caribbean
lowlands. In Costa Rica and Panama, it
also covers large areas of the Pacific ver-
sant. During the day, temperatures in the
tropical rain forest reach 30-35°C. During
the night, temperatures drop to about
25°C. Relative humidity is very high (80-
·:·
·:'l.J
Fig. 10. Buttress tree in the rain forest at Bar-
tola, Rio San Juan, Nicaragua (30 m elevation)
Photo: G. Kohler.
100%). Rainfall tends to be considerable
throughout the year, and heavy rains are
most common in the early afternoon.
Several levels of vegetation can be disting-
uished in tropical rain forest, from the
dimly lit forest floor, the scrub level, the
trees of different sizes, up to the tallest
trees of the forest, which reach high above
the canopy. The diversity of tree species is
extraordinary and can exceed 50-100 per

Fig. 9. Rain forest at Bartola, Rio San Juan, Fig. 11. Cloud forest at Volcan Mombacho,
Nicaragua (30 m elevation). Photo: G. Kohler Nicaragua (1100 m elevation). Photo: G. Kohler
18
 The Environment

Fig. 12. View of the cloud forest on Cerro Kilambe, Nicaragua. Photo: G. Kohler

hectare. Many reptiles occur exclusively in than that of the lowlands, but there is a
primary lowland and premontane rain high number of endemic species, i.e., spe-
forests and disappear with deforestation. cies that only occur in one particular area,
such as a summit or a mountain chain.
Cloud forest
A world of its own, cloud forest is the type
of natural vegetation that can be found
above 1200 m elevation in many of the
mountain ranges of Central America. In
this type of habitat, temperatures are rela-
tively low (5-15 °C, depending on altitude)
and humidity is constantly high. The trees
in cloud forest are densely covered with
epiphytes, such as bromeliads, orchids,
ferns, and mosses. Tree ferns, which can
reach several meters in height, are also
found frequently. Small streams channel
the water on its way to the lowlands. At
very high altitudes along exposed ridges, a
special sub-type of cloud forest grows, the
so-called elfin forest. Subject to low tem-
peratures, high precipitation, and high wind
speeds, trees in the elfin forest usually
attain less than 1-3 min height. The fauna Fig. 13. A cloud forest trail at Monteverde,
of cloud forests tends to be less diverse Costa Rica (1500 m elevation).Photo: G. Kohler

19
 Fig. 14. Wet savanna near Hopkins, Belize. Photo: G. Kohler

Fig. 15. Waterfall in the rain forest at Cerro San Gil, Izabal, Guatemala. Photo: G. Kohler

20
Fig. 16. Aquatic habitat between Hattleville and Belize City, Belize. Photo: G. Kohler

Fig. 17. Dry forest near Arenal, Motagua Valley, Guatemala. Photo: G. Kohler
21
Nl
Nl

~
~ =-
~ ('D

~ trj
< =
<
~ a·
Mesa Central de Chiapas
~·
0 §
('D

=
s==
....
\ ~
~

Cordillera Nombre de Dias

Sierra Madre de Chiapas

Guatemalan Plateau

Nicaraguan Depression
D 0-1000 m
[ 1000-2000 m Cordillera Central

II >2000 m Cordillera de Talamanca

iJ

Cordillera de Guanacaste
 II Marshes, swamp ~
~
~
II Mangrove 0
II Seasonal rain forest ~

II Evergreen rain forest (Caribbean versant) ~

~
II Evergreen rain forest (Pacific versant) {'D
"""'
~
II Cloud forest """'
1-1•
0
=
~
~

N
0
=
{'D
00.

Dry forest
L

Iii Pine-oak forest

L Pine savanna
~
~
trj

....=
<
'"'l

~
a
t:-:>
w
 The Environment

Climate in Central America

The climate in Central America is high-
ly location dependent, with the yearly
cycle largely characterized by variations
in precipitation levels. On the
Caribbean side, the heaviest rains fall
during December and January, while
the Pacific side receives virtually no rain
during the northern winter (the dry sea-
son on the Pacific side lasts from
January until April). In general, the
Caribbean side of Central America is
more humid and receives more precipi-
tation than the Pacific side.

The following climate diagrams illus- Belize City, Belize

trate the yearly climate in selected E 500 - 35 G
....
areas. The mean monthly precipitation .sc
0 450
is indicated by bars. The mean daily "'.~
0. 400
·;;
temperature can be read from the upper £ 350
curve and the mean nightly temperature
from the lower curve (data from MOLLER 300
20
1996, except for Utila, which was collec- 250

ted personally). 200

15

150 10

100

50

J F M A M J J A S 0 N D

Merida, Mexiko Guatemala City, Guatemala

E 500 - - ls E E 500 35
E
.sc .s e
0 450 ~ ~ 450 =
30 ~
"'.~ ~
30
~0.
0.
·;; 400 E :~ 400 E
e
0. 25
~ e 25
~
350 a. 350

300 300
20 20

250 250
15 15
200 200

150 10 150 10

100 100

50 50

J F M A M J J A s 0 N D J F M A M J J A s 0 N D

24
 The Environment

Utila, Honduras Tegucigalpa, Honduras

e
E.
500 - - 3S
E E.e soo - 3S 0
'--
c 4SO
E E
0 il ~ 450 il
30
~
30
5 !'!
·a..
5
·u
400
~ ·u
400
~
E ~ E ~
0.
3SO 2S a. 350 2S

300 300
20 20

2SO 2SO

1S 1S
200 200

1SO 10 1SO 10

100 100

so so

J F M A M J J A s 0 N D J F M A M J J A s 0 N D

San Salvador, El Salvador Managua, Nicaragua

E soo - 3S
0 e soo - 3S
0
E. '-- E. '--
c 450
0
30
~ c
:8
4SO
30
E
il
i·u 400
5
0.
E
!'!
·a.
·u
400
5
~
2S ~ 2S ~
E E
0. 3SO 0. 3SO

300 300
20 20

2SO 2SO

1S 1S
200 200

1SO 10 1SO 10

100 100

so so

J F M A M J J A s 0 N D J F M A M J J A S 0 N D

San Jose, Costa Rica Christobal, Panama

soo
"'--E eE.
- 500 - 35 - 3S -
E
~

--
E

- - ---.
-; 450 c 4SO E
~ ~
~ 30
~
:8 30
·a. 400 0.
!'!
·a. 400
~

~
·u E
·~ E
E
0. 3SO
25 ~ 0. 3SO 25 ~

300 300
20 20

2SO 2SO

1S 1S
200 200

1SO 1SO 10
10

100 100

so

F M A M J J A s 0 N D
so
~ nn
F M A M J J A S 0
J J N D

25
 Geological History
Geological History and Zoogeography of
the Reptile Fauna of Central America
SAVAGE (1966, 1982, 2002) provided
masterful analyses and discussions on the
composition, origins, and history of the
Central American herpetofauna and
developed a theory explaining the present
distribution pattern of the amphibians
and reptiles. Earlier biogeographers have
regarded Central American as a faunal
transition zone between the American con-
tinents (e.g., SIMPSON 1950, K. SCHMIDT
1954, DARLINGTON 1957, STUART 1964).
However, SAVAGE (1966, 1982, 2002)
demonstrated that the Central American
herpetofauna forms a distinct zoogeo-
graphic entity, both in regard to its history
and present composition. The following
discussion is based largely on the latter
three publications, supplemented with
information from works on the geology of
the region by MALFAIT & DINKELMAN
(1972) and MARSHALL et al. (1979).
The Central American herpetofauna is
composed of genera with an origin in the
region (autochthonous genera), as well as
those that dispersed from North and
South America into Central America.
SAVAGE (1966, 1982) concluded that three
major and one minor historical source
units had contributed to the Central
American herpetofauna.
The most important historical unit is the
"Middle American Element" (SAVAGE 1966,
1982) that is made up of genera that are
primarily tropical Mesoamerican in distri-
bution (many Central American endemics)
and have their closest relatives either in
Central or South America. The ancestors
of this historical unit usually evolved in
the region itself or in South America, and
reached far into North America during the
early Tertiary period when a tropical
mesic climate with warm temperatures
prevailed. When the climate generally
26
became cooler and dryer during the late
Cenozoic era, they became restricted to
tropical Mesoamerica. Examples of the
"Middle American Element" include the
Kinosternidae, Helodermatidae, Xantusi-
idae, and Xenosauridae. These families
are all known as fossils from early
Tertiary deposits in North America near
40° latitude north (SAVAGE 1966).
A second major historical unit is the "Old
Northern Element" (SAVAGE 1966, 1982),
which is composed of genera with a pri-
marily extratropical distribution in
Eurasia and/or North America. During the
early Tertiary period, this group's ances-
tors experienced a more or less continuous
and circumpolar distribution on the
northern landmasses. The gradual clima-
tic changes during the Cenozoic era forced
this group's ancestors southward and
caused their fragmentation into geograph-
ic isolates. As a result, this stock contains
a unique Mesoamerican component of
endemic taxa that have evolved with the
autochthonous "Middle American Element"
from the Eocene epoch onward.
A third major historical unit is the "South
American Element" (SAVAGE 1966, 1982),
which is composed of genera with primari-
ly South American distributions and rela-
tionships. Their ancestors evolved in
South America and migrated various
distances into Central America after the
closure of the Panamanian portal during
the Pliocene. Consequently, this unit is a
rather recent contributor to the current
Central American herpetofaunal diversity.
Finally, a fourth less important historical
unit, the "Young Northern Element"
(SAVAGE 1966, 1982) can be differentiated.
This unit is represented by a few genera
(including Sceloporus and Urosaurus) with

a primarily extratropical distribution
associated with semiarid to arid regions in
southwestern North America.
A brief summary of the geological history
of Central America facilitates the under-
standing of the current distribution pat-
terns of reptiles in the region. The land
connection that existed between the
American continents during the late
Cretaceous and early Tertiary periods (70
to 50 million years ago) broke up in the
Eocene epoch (54 to 35 million years ago)
due to inundation of the Isthmian Link
(= lower Central America). This so-called
Panamanian portal separated Nuclear
Central America from South America
during a period of time that lasted for
about 45 million years. It was not before
the early Pliocene (some 5 million years
ago) that the closing of the Panamanian
portal reconnected the American conti-
nents. The uplift of the Central American
mountain massifs began in the Pliocene
and continues today. The major physiogeo-
graphic changes caused significant clima-
tic shifts. During the Pliocene, the climate
became generally cooler and more arid.
Additionally, the Pleistocene glacials and
interglacials were extremely important
events.
In the early Tertiary period (e.g., during
the Paleocene epoch, 65 to 54 million years
ago), the North and South American conti-
nents were connected by a land bridge
with low relief that supported a general-
ized tropical reptile fauna that extended
northward to at least 40° latitude. It can
be assumed that all families of reptiles
that currently inhabit the Neotropics were
already present at that time. During the
Eocene, flooding of the southern parts of
the intercontinental land bridge (from
what is now central Nicaragua to north-
ern Colombia) eliminated the connection
of North and South America. Soon, two
groups were present in the northern re-
gion: the "Middle American Element" and
a subunit of the Old Northern Element,
the so-called "Central American Complex"
Geological History
(SAVAGE 1966, 1982). The latter group
reached Middle America in the Eocene
through migration southward along the
developing Rocky Mountains and became
isolated from their congeners in the
eastern and western areas of what is now
the United States by the gradual develop-
ment of a more arid climate across north-
ern Mexico. In the Oligocene epoch, condi-
tions remained similar to those of the
Eocene in Nuclear Central America, al-
though some uplift of the northern moun-
tain massifs may have occurred.
The Miocene epoch (25 to 5 million years
ago) was the time of greatest change
during the Tertiary period in Central
America, both in regard to climate and
vegetation, as well as changes to the land
relief In the Pliocene, with the disappear-
ance of the Panamanian portal, a contin-
uous land connection between the conti-
nents was re-established for the first time
since the early Eocene, making faunal
exchange possible.
The South American influence is present-
ly restricted mostly to lower Central
America (i.e., the region of what is today
Costa Rica and Panama). Only a few spe-
cies of South American origin have reached
northern Central America (e.g., Gymnoph-
thalmus speciosus), whereas most South
American elements dispersed only into
Panama (e.g., the genera Morunasaurus
and Trachyboa) or Costa Rica (e.g., the
genera Anadia, Dactyloa, and Neu-
sticurus). Many species of Central or
North American origin have migrated far
into South America (e.g., the genera
Crotalus, Iguana, Norops).
The Pliocene (5 to 1.8 million years ago)
was the most important period in the evo-
lution of the Central American reptile
fauna that led to the modern species com-
position. At the beginning of this epoch,
representatives of both historical units
(Middle American Element and Central
American Complex) were widely distrib-
uted over much of the Central American
27
 Zoogeography of the Reptile Fauna

region, with speciation centers in the are effective for lowland species: highland
uplifting mountain massifs. The moun- massifs and climatically unsuitable areas.
tains formed a climatic divide that led to An example of the latter type of barrier is
the differentiation of species communities the xeric northern part of the Yucatan
adapted to the more arid conditions on the Peninsula that prevented mesic-adapted
Pacific versant, as well as those associated species from moving from the Peten
with the more mesic habitats on the forests into the region. An area of very
Caribbean versant. complex lowland climatic filters occurs
along the Pacific versant of Costa Rica and
These modern species dispersed along western Panama.
routes that were controlled by climatic
and physiogeographic factors (Fig. 18). From the late Pliocene onward, the overall
Mountains restricted the migration oflow- picture of relief and climate in Central
land species and provided dispersal routes America resembled, for the most part, the
for highland species. Two types of barriers present physiogeography, with the excep-
tion of the marine embayment of
Nicaragua and the still flooded northern
part of the Yucatan Peninsula, both of
which became more and more dry due to
D 0 0-1000 m decreasing sea levels.
• 1000-2000
• ~2000 m During the glacial periods of the
...._. dispersal routes Pleistocene (1.8 million years ago to the
mesic climate
dispersal routes
present), the summits of the highest
arid climate Central American mountains were cov-
__.- climatic barriers ered with glaciers. Based on recent studies
on fossil pollen profiles (COLINVAUX 1993,
1996), it is assumed that the average
temperature in Central America dropped
6 to 8°C below the present level during
lowland species glacial periods, resulting in environments
in the lowlands most closely resembling
those of midelevation forests (500 to 1600
m) at present (SAVAGE 2002). During these
D 0 0-1000 m
periods the distributions of the highland
• 1 000-2000 m species were compressed downward (by
• ~2000 m
about 800 m), where they co-occurred with
....... dispersal routes
lowland species (SAVAGE 2002). During
interglacial periods, the highland species
__.- climatic barriers moved back up the cordilleras as tem-
peratures returned to levels equivalent to
those at present (STUART 1951, CAMPBELL
1984, SAVAGE 2002). It is assumed that the
cooling and warming cycles in the latter
part of the Cenozoic, in association with
highland species the continued uplift of the cordilleras,
provided a major impetus for speciation
Fig. 18. Principal dispersal routes of amphibi- (SAVAGE 2002).
ans and reptiles in Central America during
the past two million years. After SAVAGE During the turbulent geological history of
(1966). Central America, several effects have con-

28

Miocene
tributed to the evolution of the region's
reptile fauna (SAVAGE 1966):
Peninsular effects
A characteristic of peninsulas is that spe-
cies can invade, but are hindered from
further migration by the surrounding sea.
Nuclear Central America formed a pen-
insula during most of the Tertiary period
and accumulated successive invaders from
the north. The Panamanian portal served
as an effective barrier, preventing these
invaders from migrating southward. From
the Eocene through the Pliocene epochs,
this peninsula served as a center of spe-
cies differentiation.
Marine portal effects
The formation of marine portals can frag-
ment the former continuous distribution
of a species, thus stimulating independent
differentiation of the isolated populations.
With the inundation of the Isthmian Link,
species that were formerly continuously
distributed in South and North America
became isolated on the two sides of the
Panamanian portal, with differentiation
Zoogeography of the Reptile Fauna
Oligocene _ ,,_ _ _..___.....
• Old Northern Element
• Generalized Tropical Fauna
Middle American Element
South American Element
Fig. 19. Geological history of
Central America since the
Paleocene. After SAVAGE (1966) .
proceeding in Nuclear Central America
independently from that in South
America. Examples for such sister groups
are the genera Ameiva I Cnemidophorus,
Caiman I Crocodylus, and Dactyloa I
Norops.
Insular effects
Due to isolation, islands stimulate specia-
tion. During the Tertiary period, particu-
larly in the Miocene epoch (24-5 million
years ago), the numerous islands associa-
ted with the Isthmian Link I Panamanian
portal zone of instability supported insu-
lar evolution.
Land bridge effects
The formation of land bridges makes pos-
sible the invasion of species previously
hindered from immigration by the exist-
ence of marine portals. After the re-estab-
lishment of the Isthmian Link in the
Pliocene, invasion from the south by spe-
cies previously excluded from Central
America began and continues to the pres-
ent.
29
 Crocodylia
Crocodilians (Crocodylia)
The order Crocodylia is represented in
Central America by three species in two
families. Typical of all species of the order,
these are large, robust, semiaquatic rep-
tiles. They inhabit lakes, lagoons, swamps,
and large rivers. These strictly carni-
vorous reptiles feed on invertebrates and
vertebrates and exhibit necrophagy and
even cannibalism (LEE 1996).
Fig. 20. Adult Crocodylus acutus (Rio San
Juan, Nicaragua). Photo: G. Kohler
Fig. 21. Adult Morelet's crocodile (Crocodylus
moreletii). Photo: R. Cedeno V.
30
In Central America, the American croco-
dile (Crocodylus acutus) inhabits predom-
inantly brackish coastal environments
(lagoons, marshes, and mangrove swamps)
with individuals penetrating further
inland along major rivers. Morelet's
crocodile (Crocodylus moreletii) and the
spectacled or brown caiman (Caiman
crocodilus) live primarily in freshwater
habitats, such as lakes, ponds, and rivers.
Fig. 22. Spectacled caiman (Caiman crocodilus) at
Caiio Negro, Costa Rica. Photo: H. Maass
Fig. 23. Spectacled caiman (Caiman crocodilus) at
Caiio Negro, Costa Rica. Photo: H. Maass
 Crocodylia
Crocodilians (Crocodylia)
The order Crocodylia is represented in
Central America by three species in two
families. Typical of all species of the order,
these are large, robust, semiaquatic rep-
tiles. They inhabit lakes, lagoons, swamps,
and large rivers. These strictly carni-
vorous reptiles feed on invertebrates and
vertebrates and exhibit necrophagy and
even cannibalism (LEE 1996).
Fig. 20. Adult Crocodylus acutus (Rio San
Juan, Nicaragua). Photo: G. Kohler
Fig. 21. Adult Morelet's crocodile (Crocodylus
moreletii). Photo: R. Cedeno V
30
In Central America, the American croco-
dile (Crocodylus acutus) inhabits predom-
inantly brackish coastal environments
(lagoons, marshes, and mangrove swamps)
with individuals penetrating further
inland along major rivers. Morelet's
crocodile (Crocodylus moreletii) and the
spectacled or brown caiman (Caiman
crocodilus) live primarily in freshwater
habitats, such as lakes, ponds, and rivers.
Fig. 22. Spectacled caiman (Caiman crocodilus) at
Caiio Negro, Costa Rica. Photo: H. Maass
Fig. 23. Spectacled caiman (Caiman crocodilus) at
Caiio Negro, Costa Rica. Photo: H. Maass

Fig. 24. Crocodylus moreletii (Peten,
Guatemala). Photo: G. Kohler
All crocodilians are oviparous and lay
hard-shelled eggs (usually seasonally, and
a single clutch per year). The spectacled
caiman and Morel et, s crocodile build
mound-type nests made of dead vegetation,
sticks, and mud for their eggs, whereas the
American crocodile excavates a shallow
hole in the ground. In Chiapas, Mexico, the
egg-laying season of the American croc-
odile is during the months of March, April,
and May with individual nests also
observed as early as February (ALVAREZ
DEL TORO 1974). The female digs a hole,
approximately 70 cm in diameter and 20
to 50 cm deep. Preferably, nests are made
on sandy embankments above the water-
line or high tide mark. Crocodylus acutus
produces 15 to 105 (usually 20 to 60) eggs
per clutch that hatch after an incubation
period of 85 to 111 days (KOHLER 1997).
The mound-type nests of Caiman crocodilus
can reach a height of one meter and a
diameter of more than two meters (STATON
& DIXON 1977). After oviposition, the fe-
male closes the nest cavity and crawls over
it several times to condense the material.
However, air pockets prevail in the nest
throughout the incubation period. Due to
fermentation and decomposition of the
plant material, constant humid tempera-
tures of 32-34 °C are produced within the
mound. Caiman crocodilus (egg-laying
Crocodylia
(/J
.....a
- .....
"t:I
0
~
e
u
Fig. 25. Juvenile Spectacled caiman (Bartola,
Nicaragua). Photo: G. Kohler
season from early August through early
November; GRENARD 1991) produces clut-
ches of 10 to 23 eggs that hatch after an
incubation period of 88 to 104 days. In
Crocodylus moreletii (egg-laying season
from April through June), the clutch size
varies from 20 to 45 eggs and incubation
takes 78 to 98 days (ALVAREZ DEL TORO
1974, KOHLER 1997).
As far as we know, all crocodilians exhibit
parental care (GREER 1971, KUSHLAN &
SIMON 1981). During the incubation peri-
od, the female remains in the immediate
vicinity of the nest and attacks potential
egg predators (MODHA 1967, COTT 1971).
The female responds to vocalizations
made by juveniles from within the nest by
opening it and transporting the hatchlings
to water (POOLEY 1977).
In crocodilians, the incubation tempera-
ture determines the sex of the hatchlings.
In most species, high and low tem-
peratures produce male hatchlings, where-
as intermediate temperatures produce
females (DEEMING & FERGUSON 1988,
KOHLER 1997).
In Central America, all three species are
endangered due to overhunting and
destruction of habitat. National laws to
protect crocodilians exist in all countries.
31

Fig. 24. Crocodylus moreletii (Peten,
Guatemala). Photo: G. Kohler
All crocodilians are oviparous and lay
hard-shelled eggs (usually seasonally, and
a single clutch per year). The spectacled
caiman and Morelet's crocodile build
mound-type nests made of dead vegetation,
sticks, and mud for their eggs, whereas the
American crocodile excavates a shallow
hole in the ground. In Chiapas, Mexico, the
egg-laying season of the American croc-
odile is during the months of March, April,
and May with individual nests also
observed as early as February (ALVAREZ
DEL TORO 1974). The female digs a hole,
approximately 70 cm in diameter and 20
to 50 cm deep. Preferably, nests are made
on sandy embankments above the water-
line or high tide mark. Crocodylus acutus
produces 15 to 105 (usually 20 to 60) eggs
per clutch that hatch after an incubation
period of 85 to 111 days (KOHLER 1997).
The mound-type nests of Caiman crocodilus
can reach a height of one meter and a
diameter of more than two meters (STATON
& DIXON 1977). After oviposition, the fe-
male closes the nest cavity and crawls over
it several times to condense the material.
However, air pockets prevail in the nest
throughout the incubation period. Due to
fermentation and decomposition of the
plant material, constant humid tempera-
tures of 32-34 °C are produced within the
mound. Caiman crocodilus (egg-laying
Crocodylia
l:ll
.....~
-
.....
"ti
0
c,)
e
0
Fig. 25. Juvenile Spectacled caiman (Bartola,
Nicaragua). Photo: G. Kohler
season from early August through early
November; GRENARD 1991) produces clut-
ches of 10 to 23 eggs that hatch after an
incubation period of 88 to 104 days. In
Crocodylus moreletii (egg-laying season
from April through June), the clutch size
varies from 20 to 45 eggs and incubation
takes 78 to 98 days (ALVAREZ DEL TORO
1974, KOHLER 1997).
As far as we know, all crocodilians exhibit
parental care (GREER 1971, KUSHLAN &
SIMON 1981). During the incubation peri-
od, the female remains in the immediate
vicinity of the nest and attacks potential
egg predators (MODHA 1967, COTT 1971).
The female responds to vocalizations
made by juveniles from within the nest by
opening it and transporting the hatchlings
to water (POOLEY 1977).
In crocodilians, the incubation tempera-
ture determines the sex of the hatchlings.
In most species, high and low tem-
peratures produce male hatchlings, where-
as intermediate temperatures produce
females (DEEMING & FERGUSON 1988,
KOHLER 1997).
In Central America, all three species are
endangered due to overhunting and
destruction of habitat. National laws to
protect crocodilians exist in all countries.
31
 Crocodylia
Fig. 26. Female Spectacled caiman with neonates (Bartola, Nicaragua).
Furthermore, Crocodylus acutus and C.
moreletii are listed in Appendix 1, and
Caiman crocodilus in Appendix 2 of the
Convention on International Trade in
Endangered Species (CITES).
Fig. 27 Juvenile American crocodile
(Crocodylus acutus) at Rio Patuca, Honduras.
Photo: G. Kohler
32
Photo: G. Kohler
Alligatoridae
Caiman crocodilus (LINNAEUS 1758), Systema
Naturae, ed. 10: 206; type locality: unknown.
Maximum total length 2 m . Southern Mexico
along most of the coasts of Central America
into South America, sea level to 200 m eleva-
tion. Recently, BUSACK & PANDYA (2001)
demonstrated that no subspecies of C. croco-
dilus should be recognized. The same authors
raised the former subspecies yacare to full
species status.
Crocodylidae
Crocodylus acutus CUVIER 1807, Ann. Mus.
Hist. Nat., Paris, 10: 55; type locality: Santo
Domingo. Maximum total length 4 m (rarely
up to 6 m). From southern Florida, USA,
across the Greater Antilles, as well as from
Sinaloa, Mexico to South America (Colombia
and Venezuela), sea level to 500 m elevation.
Crocodylus moreletii DUMERIL & BIBRON
1851, Cat. Method. Coll. Rept.: 28; type locali-
ty: Lake Peten, Guatemala. Maximum total
length 3 m (rarely up to 4 m). Tamaulipas,
Mexico, across most of the Yucatan Peninsula
to southern Belize, sea level to 500 m eleva-
tion.
 Crocodylia
Fig. 26. Female Spectacled caiman with neonates (Bartola, Nicaragua).
Furthermore, Crocodylus acutus and C.
moreletii are listed in Appendix 1, and
Caiman crocodilus in Appendix 2 of the
Convention on International Trade in
Endangered Species (CITES).
Fig. 27 Juvenile American crocodile
(Crocodylus acutus) at Rio Patuca, Honduras.
Photo: G. Kohler
32
Photo: G. Kohler
Alligatoridae
Caiman crocodilus (LINNAEUS 1758), Systema
Naturae, ed. 10: 206; type locality: unknown.
Maximum total length 2 m. Southern Mexico
along most of the coasts of Central America
into South America, sea level to 200 m eleva-
tion. Recently, BUSACK & PANDYA (2001)
demonstrated that no subspecies of C. croco-
dilus should be recognized. The same authors
raised the former subspecies yacare to full
species status.
Crocodylidae
Crocodylus acutus CUVIER 1807, Ann. Mus.
Hist. Nat., Paris, 10: 55; type locality: Santo
Domingo. Maximum total length 4 m (rarely
up to 6 m). From southern Florida, USA,
across the Greater Antilles, as well as from
Sinaloa, Mexico to South America (Colombia
and Venezuela), sea level to 500 m elevation.
Crocodylus moreletii DUMERIL & BIBRON
1851, Cat. Method. Coll. Rept.: 28; type locali-
ty: Lake Peten, Guatemala. Maximum total
length 3 m (rarely up to 4 m). Tamaulipas,
Mexico, across most of the Yucatan Peninsula
to southern Belize, sea level to 500 m eleva-
tion.
 Crocodylia

rJJ

...=
-
... i:TJ

"O
0
~

e
u

Fig. 28. Crocodylus acutus submerged in a

small stream (Rio Patuca, Olancho, Honduras).
Photo: G. Kohler

Fig. 29. Crocodylus acutus (Caii.on de Sumidero,

Caiman crocodilus
Chiapas, Mexico). Photo: A Ramirez

Crocody/us acutus

33
 Crocodylia

I'll

a
.....
......
.....
"ti
0
CJ
e
0

Fig. 28. Crocodylus acutus submerged in a

small stream (Rio Patuca, Olancho, Honduras).
Photo: G. Kohler

Fig. 29. Crocodylus acutus (Canon de Sumidero,

Caiman crocodilus
Chiapas, Mexico). Photo: A Ramirez

• Crocodylus acutus

33
 Crocodylia

nuchals

Caiman crocodilus Crocodylus acutus

Fig. 30 Characteristics important in the identification of crocodylians. Drawings by M. Vesely.

Key to crocodilians
1 a A transverse ridge just anterior to eyes on
top of head (Fig. 30); no teeth of lower jaw
visible when mouth closed .. .................. .. .
.. .. .. .. .. .. .. .. .. .. .. ......... Caiman crocodilus
b No transverse ridge just a.nterior to eyes
on top of head (Fig. 30); 4th tooth oflower a. Crocody/us acutus
jaw lies laterally in .an open groove and is
visible when mouth is closed ................... 2
2 a Primary subcaudal whorls not interupted
by incomplete secondary whorls (Fig. 31a);
snout relatively slender, about twice as
long as wide ............ ...Crocodylus acutus
b Primary subcaudal whorls interupted by
incomplete secondary whorls (Fig. 31b);
snout relatively broad, length about 1.5
times width ........... Crocodylus moreletii

b. Crocody/us moreletii
Further Reading
WERMUTH & MERTERNS 1961, NEILL 1971,
BRAZAITIS 1973, ALVAREZ DEL TORO 197 4; Ross &
Ross 1974; BUFFETAUT 1979, Ross 1987,
DENSMORE & OWEN 1989, Ross & GARNETT
1989, GRENARD 1991, PEREZ-HiGAREDA et al. Fig. 31. Ventral surface of tail. Incomplete
1991, ERNST et al. 1999, CEDENO-VAzQUEZ et al. secondary subcaudal whorls in Crocodylus
2006 moreletii shaded.

34
 Crocodylia

nuchals

Caiman crocodi/us Crocody/us acutus

Fig. 30 Characteristics important in the identification of crocodylians. Drawings by M. Vesely.

Key to crocodilians ~-\

1 a A transverse ridge just anterior to eyes on

top of head (Fig. 30); no teeth oflower jaw )
visible when mouth closed ...................... .
. .... ... . . . . . . . . . . . . . . . ........ Caiman crocodilus
b No transverse ridge just anterior to eyes
on top of head (Fig. 30); 4th tooth of lower a. Crocodylus acutus
jaw lies laterally-in an open groove and is
visible when mouth is closed ............... .... 2
2 a Primary subcaudal whorls not interupted
by incomplete secondary whorls (Fig. 31a);
snout relatively slender, about twice as ~- ·
long as wide .......... .... .Crocodylus acutus
b Primary subcaudal whorls interupted by
incomplete secondary whorls (Fig. 31b); I
snout relatively broad, length about 1.5 \
times width ........... Crocodylus moreletii

b. Crocodylus moreletii
Further Reading
WERMUTH & MERTERNS 1961, NEILL 1971,
BRAZAITIS 1973, ALVAREZ DEL TORO 1974; Ross &
Ross 1974; BUFFETAUT 1979, Ross 1987,
DENSMORE & OWEN 1989, Ross & GARNETT
1989, GRENARD 1991, PEREZ-HIGAREDA et al. Fig. 31. Ventral surface of tail. Incomplete
1991, ERNST et al. 1999, CEDEiiro-VAzQUEZ et al. secondary subcaudal whorls in Crocodylus
2006 moreletii shaded.

34
 Testudines

Turtles (Testudines)

b.

marginal plates

axillar plate inguinal plate

c. Fig. 32. Characteristics
important in the identifi·
cation of turtles
a. Lateral view
chin barbels b. Dorsal view;
c. Ventral view;
M=marginal plate;
humeral plate P=pleural plate;
V=vertebral plate.

abdominal plate

Turtles are represented by 25 species in 14

genera and 7 families in Central America,
the kinosternids and emydids being the
most speciose with 8 species each.

Key to genera of turtles

1 a Forelimbs modified as oar-like flippers; Fig. 33. Plastrons.
two or fewer claws on each foreleg; oceanic
species Sea Turtles (see key on page 56) After WERMUTH &
MERTENS (1961).
b Forelimbs not modified as oar-like flip- a. Dermatemys
pers; more than two claws on each foreleg;
species living on land or in fresh or
brackish water habitats .... ..... .... .... ... ..... 2
2 a Plastron with 11 or fewer plates ........... .. 3
b Plastron with 12 plates ..... ..... .. ..... ........ . 7
3 a Plastron not reduced and not cruciform .. 4
b Plastron very reduced and cruciform (Fig.
33b-d) ·· ····· ··· ········ ····· ·· ········ ·················· 5 b. Chelydra c. Staurotypus d. Claudius

35
 Testudines

Turtles (Testudines)

marginal plates

axillar plate inguinal plate

c. Fig. 32. Characteristics
important in the identifi-
cation of turtles
a. Lateral view
chin barbels
b. Dorsal view;
c. Ventral view;
M=marginal plate;
P=pleural plate;
V=vertebral plate.

abdominal plate

Turtles are represented by 25 species in 14

genera and 7 families in Central America,
the kinosternids and emydids being the
most speciose with 8 species each.

Key to genera of turtles

1 a Forelimbs modified as oar-like flippers; Fig. 33. Plastrons.
two or fewer claws on each foreleg; oceanic
species Sea Turtles (see key on page 56) After WERMUTH &
MERTENS (1961).
b Forelimbs not modified as oar-like flip- a. Dermatemys
pers; more than two claws on each foreleg;
species living on land or in fresh or
brackish water habitats ......................... 2
2 a Plastron with 11 or fewer plates ............. 3
b Plastron with 12 plates .......................... 7
3 a Plastron not reduced and not cruciform .. 4
b Plastron very reduced and cruciform (Fig.
33b-d) ·········· ·············· ··························· 5 b. Chelydra c. Staurotypus d. Claudius

35
 Testudines

4 a Plastron with two moveable hinges (Figs.

32c and 34); chin barbels present
......... ............ .............. ........ ... Kinosternon
b Plastron rigid, without moveable hinges
(Fig. 33a ); chin barbels absent ....... ............ .
......................... ..... Dermatemys (in part)
5 a Tail as long as carapace or longer Chelydra
b Tail shorter than carapace ....................... 6
6 a Plastron rigid, without moveable hinges
(Fig. 33d); bridge connected to carapace by
ligaments .. ..... ................ ......... Claudius
b Plastron with moveable hinge ; bridge
connected to carapace by an osseous suture
.... ....... .... .. .. ......... .............. Staurotypus
7 a Plastron with moveable hinge; plastron
and carapace in contact, bridge absent ..... .
............ ..... ............. ........ .... ... Terrapene
b Plastron rigid, without moveable hinge;
bridge present ............. ................ ............. 8
Fig. 35. Chelydra serpentina (Rio San Juan,
8 a Hind limbs elephantine (columnar); cara- Nicaragua). Photo: G. Kohler
pace dome-shaped; without webbing bet-
ween toes ..... ........ ... .... ....... . Geochelone
b Hind limbs not columnar; carapace rela-
tively flat, only slightly convex; webbing
between toes more or less developed ....... 9
9 a Limbs unicolor dark gray; snout slightly Chelydridae
upturned, somewhat tubular ........... .......... .
................ ... .. ...... .. Dermatemys (in part) Chelydra
b Limbs with some kind of pattern (mostly
pale stripes); snout not as above ........... 10
The snapping turtle (Chelydra serpentina)
lOa Pattern of carapace consists of yellow can reach 4 7 cm carapace length with a
rings with dark center; hind feet with weight of 20 kg and should be handled
extensive webbing ........ ......... Trachemys
with great care. It is able to deliver quick
b Pattern of carapace without yellow rings,
or, if yellow rings are present, then h ind strikes with its powerful jaws. Snapping
feet with little or no webbing .................... . turtles inhabit ponds, lakes, and slow-
.......... . . .. ....... ....... .............. Rhinoclemmys moving rivers with muddy bottoms .
Mostly nocturnal, these animals usually
rest in shallow water with only the
nostrils and eyes protruding above the
surface during the day. At night, this turt-
le walks slowly on the bottom of its watery
habitat searching for food. Snapping turt-
les feed on almost anything they can swal-
low, particularly insects, shrimp, crabs,
fish, and frogs, but also crocodile and turt-
le hatchlings, snakes, birds, and small
mammals, as well as plant matter such as
algae and other water plants and fruits .
Fig. 34. All species of Kinosternon have a plas-
tron with hinges that enable a complete occlu- The female digs a nest for her eggs (in
sion of the shell (K. leucostomum). April through June) on a sandy embank-
Photo: G. Kohler ment above the waterline, preferably in a
36
 Testudines

4 a Plastron with two moveable hinges (Figs.

32c and 34); chin barbels present ...... .
...... ............. .. ...................... ... Kinosternon
b Plastron rigid, without moveable hinges
(Fig. 33a); chin barbels absent ................... .
......... .... ...... ........... Dermatemys (in part)
5 a Tail as long as carapace or longer Chelydra
b Tail shorter than carapace ....................... 6
6 a Plastron rigid, without moveable hinges
(Fig. 33d); bridge connected to carapace by
ligaments .... ... ... ............... ....... Claudius
b Plastron with moveable hinge; bridge
connected to carapace by an osseous suture
.. .. ........................... ........... Staurotypus
7 a Plastron with moveable hinge; plastron
and carapace in contact, bridge absent ......
.. .... ... ....... ...... ....................... Terrapene
b Plastron rigid, without moveable hinge;
bridge present .................... ..... .. ............... 8
Fig. 35. Chelydra serpentina (Rio San Juan,
8 a Hind limbs elephantine (columnar); cara- Nicaragua). Photo: G. Kohler
pace dome-shaped; without webbing bet-
ween toes .... .................... .... Geochelone
b Hind limbs not columnar; carapace rela-
tively flat, only slightly convex; webbing
between toes more or less developed ....... 9
9 a Limbs unicolor dark gray; snout slightly Chelydridae
upturned, somewhat tubular ... ...................
..... ........................ Dermatemys (in part) Chelydra
b Limbs with some kind of pattern (mostly
pale stripes); snout not as above .. ......... 10
The snapping turtle (Chelydra serpentina)
lOa Pattern of carapace consists of yellow can reach 47 cm carapace length with a
rings with dark center; hind feet with weight of 20 kg and should be handled
extensive webbing ...... ........... Trachemys
with great care. It is able to deliver quick
b Pattern of carapace without yellow rings,
or, if yellow rings are present, then hind strikes with its powerful jaws. Snapping
feet with little or no webbing ....... ............. . turtles inhabit ponds, lakes, and slow-
............. ............. .. ...... ....... . Rhinoclemmys moving rivers with muddy bottoms .
Mostly nocturnal, these animals usually
rest in shallow water with only the
nostrils and eyes protruding above the
surface during the day. At night, this turt-
le walks slowly on the bottom of its watery
habitat searching for food. Snapping turt-
les feed on almost anything they can swal-
low, particularly insects, shrimp, crabs,
fish, and frogs, but also crocodile and turt-
le hatchlings, snakes, birds, and small
mammals, as well as plant matter such as
algae and other water plants and fruits.
Fig. 34. All species of Kinosternon have a plas-
tron with hinges that enable a complete occlu- The female digs a nest for her eggs (in
sion of the shell (K. leucostomum). April through June) on a sandy embank-
Photo: G. Kohler ment above the waterline, preferably in a
36

spot with some protection by low vegeta-
tion. Twenty to 30 eggs (exceptionally up
to 83 eggs in very large females) are laid
per clutch. The eggs measure 23 to 33 mm
(weight 7 to 15 g) and the juveniles (cara-
pace length 24 to 31 mm) hatch after an
incubation period of 55 to 125 days (ERNST
& BARBOUR 1989).
Chelydra serpentina (LINNAEUS 1758),
Systema Naturae, ed. 10: 199; type locality:
"Calidis regionibus". Carapace length to 47
cm. Widely distributed across southeastern
U.S.A., Mexico, and Central America to
Ecuador, sea level to 1160 m elevation. The
subspecies Chelydra s. acutirostris PETERS
1862 (northern Honduras across lower
Central America to northern South America)
and Chelydra s. rossignonii (BOCOURT 1868)
(central Veracruz, Mexico, across the base of
the Yucatan Peninsula to western Honduras)
occur in Central America.
Further Reading
ERNST & GIBBONS 1988
• Chelydra serpentina
• Dermatemys mawii
Dermatemydidae
Dermatemydidae
Dermatemys
The family Dermatemydidae consists of a
single genus with one species. Dermat-
emys mawii is found in lagoons, rivers, and
lakes from southeastern Mexico through
Belize. The species also tolerates brackish
water. These nocturnal turtles are highly
adapted to their aquatic environment, and
are capable of performing prolonged dives.
On land, however, locomotion is difficult,
as is lifting the head. Adult Central
American river turtles feed almost
exclusively on plant material (algae and
other water plants and fruits that fall into
the water). The egg-laying period lasts
from April through September with the
females burying their clutches of 6 to 20
eggs (57-70 x 30-34 mm) on sandy
embankments near the shoreline.
Dermatemys mawii GRAY 1847, Proc. Zool.
Soc. London 1847: 55; type locality: unknown.
Carapace length to 65 cm. From central
Veracruz, Mexico, through Tabasco, northern
Chiapas, Mexico, and northern Guatemala to
Belize, sea level to 300 m elevation.
Further Reading
IVERSON & MITTERMEIER 1980
Fig. 36. Dermatemys mawii in Peten,
Guatemala. Photos: G. Kohler
37
 Emydidae

Emydidae Carapace length to 290 mm. Southeastern

Rhinoclemmys
The species of Rhinoclemmys are distribu-
ted across the tropics and subtropics of
Mexico, Central America, and northern
South America. Some species (e.g., R. fune-
rea) are highly aquatic, whereas the
majority of Neotropical wood turtles have
a semiaquatic or even terrestrial lifestyle.
In the Biosphere Reserve, Bosawas,
Nicaragua, I encountered several R. annu-
lata within the rain forest, some 100
meters from the nearest body of water. On
the other hand, I have recorded R. funerea
in the same general area, but exclusively
in rivers. There, it can frequently be seen
basking on partially submerged logs.
Panama to northwestern Ecuador, sea level to
400 m elevation.
Rhinoclemmys pulcherrima (GRAY 1855), Cat.
Shield Rept. Brit. Mus. 1: 25; type locality:
Mexico. Carapace length to 200 mm. Sonora,
Mexico, to Costa Rica, sea level to 1160 m ele-
vation. The subspecies Rhinoclemmys p. pul-
cherrima (Guerrero and Oaxaca, Mexico), R.
p. incisa (BOCOURT 1868) (Oaxaca, Mexico, to
northern Nicaragua), and R. p. manni DUNN
1930c (southern Nicaragua to Costa Rica)
occur in Central America.
Rhinoclemmys rubida (Cope 1870), Proc.
Amer. Philos. Soc. 11: 148; type locality:
Juchitan, Oaxaca, Mexico. Carapace length to
230 mm. Pacific coast of Mexico from Jalisco
to western Chiapas, sea level to 750 m. Only
the nominate subspecies occurs in Central
America.

Neotropical wood turtles are highly herbi-

vorous and only occasionally ingest in-
sects. They often forage on land at night,
feeding predominantly on broadleaf
plants, grasses, and fruits. Most Central
American species produce small clutches
with one or two eggs; only R. melanosterna
and R. pulcherrima lay larger clutches (up
to 5 eggs per clutch) (ERNST & BARBOUR
1989, KOHLER 1997).

Rhinoclemmys annulata (GRAY 1860), Proc.

Zool. Soc. London 1860: 231; type locality:
Esmeraldas, Ecuador. Carapace length to 200 Fig. 37. Rhinoclemmys rubida.
mm. Eastern Honduras to northwestern Photo: R. Wicker
South America, sea level to 920 m elevation.
Rhinoclemmys areolata (DUMERIL & BIBRON
1851), Cat. Method. Coll. Rept.: 10; type loca-
lity: El Peten, Guatemala. Carapace length to
200 mm. Southern Veracruz, Mexico, to extre-
me western Honduras, sea level to 300 m ele-
vation.
Rhinoclemmys funerea (COPE 1876), J. Acad.
Nat. Sci. Philadelphia (2) 8: 154; type locality:
Puerto Limon, Costa Rica. Carapace length to
325 mm. Rio Coco (border Honduras I
Nicaragua) to central Panama, sea level to
600 m elevation.
Rhinoclemmys melanosterna (GRAY 1861),
Proc. Zool. Soc. London 1861: 205; type locali-
ty: Cherunha, Gulf of Darien, Colombia. Fig. 38. Juvenile Rhinoclemmys funerea (Rio
San Juan, Nicaragua). Photo: G. Kohler
38
 Emydidae

---· • Rhinoclemmys funerea

• Rhinoclemmys melanosterna

Fig. 39. Rhinoclemmys pulcherrima (Rio

Patuca, Olancho, Honduras). Photo: G. Kohler
• Rhinoclemmys pulcherrima

• Rhinoclemmys annulata
• Rhinoclemmys areolata
v • Rhinoc/emmys rubida
Fig. 40. Rhinoclemmys areolata (Peten,
Guatemala). Photo: G. Kohler

Fig. 41. Rhinoclemmys areolata (Belize).

Photo: G. Kohler

Fig. 42. Rhinoclemmys annulata is more

terrestrial than other species of the genus.
Photo: G. Kohler
39
 Emydidae

Key to Rhinoclemmys
1 a Hind feet with extensive webbing ........ ... 2
b Hind feet with little or no webbing ......... 4
2 a Iris pale gray without transverse dark
bar; posterior surface of thigh with verti-
cal orange bars ...... ......... ............... .. .. . .
.... ... ......... Rhinoclemmys melanosterna
b Iris with transverse dark bar; pattern of
thigh not as above .... ....... .... .............. ...... 3
3 a One or two red supratemporal stripes
present on each side reaching snout; cara-
pace with or without ocellated pattern
.............. .. .. Rhinoclemmys pulcherrima
b No red supratemporal stripes; carapace
without ocellated pattern ............ ....... .....
........ ......... . . . ... . . . . Rhinoclemmys funerea Fig. 44. Terrapene carolina yucatana
4 a Dorsal surface of head with distinct large (Calakmul Biosphere Reserve, Campeche,
yellow marking (often horseshoe-shaped; Mexico). Photo: H . Bahena B.
Fig. 37); bridge darkly pigmented ........... .
..... ..... .. ......... ........ Rhinoclemmys rubida
b Dorsal surface of head not as above;
bridge pigmented or not .......... ............. .... 5
5 a Dorsal surface of head uniformly dark
brown; supratemporal stripe, if present,
indistinct; carapace with broad keel Terrapene
.... .... .. ......... .... Rhinoclemmys annulata
b One or two pale supratemporal stripes
present on each side of head posterior to Four species of the genus Terrapene are
orbit; keel on carapace, if present, narrow currently recognized, with only T. carolina
........................ Rhinoclemmys areolata occurring in southern Mexico (ERNST &
MCBREEN 1991). On the Yucatan
Further Reading Peninsula, this species inhabits wet
ERNST 1978, 1980a-c, 1981a-c, PEREZ-HIGAREDA savannas and pasture land, as well as
& SMITH 1987, MERCHAN FORNELINO 2005 thorn forest and evergreen rain forest,
with a preference for open habitats (LEE
1996). Although box turtles exhibit a ter-
restrial life style, individuals can be ob-
served sitting in shallow water. Juveniles
· appear to have a predominantly carni-
vorous diet, while adults are predominant-
ly herbivorous. The carnivorous diet of box
turtles includes snails, worms, insects, spi-
ders, salamanders, and frogs.

Copulation is preceded by an extensive

Fig. 43. Rhinoclemmys pulcherrima (near
Portegolpe, Guanacaste, Costa Rica).
Photo: H. Bringsoe
courtship, which can be divided into three
phases (EVANS 1953): (1) a circling, biting,
and shoving phase; (2) a preliminary
mounting phase; and (3) a copulatory
phase. The females produce clutches of 2
to 4 eggs that hatch after 50 to 70 days,
depending on ambient temperature
(KOHLER 1997).

40

Terrapene carolina (LINNAEUS 1758), Systema
Naturae, ed. 10: 198; type locality: "Carolina."
Carapace length to 200 mm. Widely distribu-
ted over the central and eastern U.S.A. with
isolated populations in Mexico (Tamaulipas,
San Luis Potosi, Veracruz, and the Yucatan
Peninsula), sea level to 100 m elevation. The
subspecies T. c. mexicana (GRAY 1849b) (Ta-
maulipas to Veracruz, Mexico) and T. c. yuca-
tana (BOULENGER 1895) (Yucatan Peninsula)
occur in the area covered by this book.
Further Reading
MILSTEAD 1969 (revision ofTerrapene); ERNST &
MCBREEN 1991, H. SMITH et al. 1996
Terrapene carolina
Fig. 45. Terrapene carolina yucatana
(Campeche, Mexico). Photo: J.C. Lee
Emydidae
Fig. 46. Juvenile Trachemys scripta
(Alamikamba, Nicaragua). Photo: G. Kohler
Trac hemys
The slider turtle (Trachemys scripta) has
the widest geographic distribution of any
non-marine turtle species. Due to its popu-
larity as a pet, it represents the most well
known turtle species. At the same time,
there is still considerable disagreement
among scientific authorities regarding its
taxonomy on both the generic and subspe-
cific level. I follow CARL H. ERNST (in
GIBBONS 1990: pp. 57-67), who places this
species in the genus Trachemys and recog-
nizes the subspecies T. s. venusta
(Caribbean versant of southern Mexico
and Yucatan Peninsula), T. s. grayi (Pacific
versant of the Isthmus of Tehuantepec to
western Guatemala), and T. s. ornata
(Honduras to Panama) in Central
America. The position of JOHN M. LEGLER
(in GIBBONS 1990: pp. 82-105), who assigns
the species to the genus Pseudemys, with
the following Central American subspecies
P. s. venusta (Caribbean versant from
41
 Emydidae

southern Mexico through most of Central

America), P. s. grayi (Pacific versant of the
Isthmus of Tehuantepec to border of El
Salvador), and P. s. emolli (Nicaraguan
lakes and adjacent Costa Rica), seems
unjustified. According to the studies of
SEIDEL & SMITH (1986) and SEIDEL &
JACKSON (1990), the generic name
Trachemys is correct for scripta and a few
more species from the Caribbean region.
Recently, SEIDEL (2002) proposed to recog-
nize some of the former subspecies of T.
scripta as distinct species. According to his
taxonomic arragement, two species of
slider turtles are recognized in Central
America: T. venusta and T. emolli (see also Fig. 47. Adult female Slider turtle (Trachemys
ERNST 2008). However, the taxonomy of scripta) at Rio Patuca, Honduras.
Central American slider turtles may be Photo: G. Kohler
subject to further revisions in the near
future. Therefore, I have adopted a conser-
vative approach and have used the name length of 200 to 240 mm (MOLL & LEGLER
T. scripta for the time being. 1971, ERNST & BARBOUR 1989, VOGT 1990).
Slider turtles are diurnal and exhibit a
very aquatic lifestyle. They occupy most Trachemys scripta (SCHOEPFF 1792), Hist.
freshwater habitats, with a preference for Testud.: 16; type locality: unknown. Carapace
length to 60 cm. Widely distributed across
quiet waters with soft bottoms, such as central and eastern U.S.A. and southward
ponds, lakes, and rivers. This turtle can through Mexico and Central America to
frequently be seen basking on partially Colombia and Venezuela, sea level to 1000 m
submerged rocks, roots, and logs. elevation. See text above for subspecies.
Juveniles feed mostly on fish and aquatic
invertebrates, such as insects, worms, and Further Reading
snails. The food preferences of these ani- MOLL & LEGLER 1971, GIBBONS 1990, SEIDEL
mals change with age toward a more herb- 2002
ivorous diet, including algae and other
water plants. Adults are virtually omni-
vorous. Slider turtles are preyed upon by
crocodilians and humans.

During the egg-laying period, which is in

the dry season (December through May),
the females bury their clutches of 9 to 25
eggs in open places that are exposed to the
sun. The female digs a hole of 10 to 15 cm
depth that is usually somewhat wider at
the bottom. A female can produce up to six
clutches in one season. Juveniles (cara-
pace length 30-33 mm) hatch after an
incubation period of 65 to 85 days.
Females reach sexual maturity at an age
of five to seven years and a carapace • Trachemys scripta

42
 Kinosternidae

Kinosternidae reported to estivate sequestered in mud

during the dry season and resume activity
Claudius with the onset of the rainy season
(ALVAREZ DEL TORO 1983).
The genus Claudius is monotypic; the sole
species is C. angustatus, the chopontil or Claudius angustatus COPE 1865, Proc. Acad.
narrow-bridged musk turtle. Narrow- Nat. Sci. Philadelphia 17: 187; type locality:
bridged musk turtles are characterized by Tabasco, Mexico. Carapace length to 165 mm.
a small plastron that is connected to the Central Veracruz, Mexico, across northern
Guatemala to Belize, sea level to 200 m ele-
carapace by ligaments. These turtles are vation. Recently recorded on the north-
known for their pugnacious attitude, and western part of the Yucatan Peninsula
should be handled with great care in order (CALDERON MANDUJANO et al. 2001).
to avoid severe bites. Claudius angustatus
is an aquatic species that principally in- Further Reading
habits areas with seasonally flooded IVERSON & BERRY 1980
grasslands and forages in these grass-
lands during the flooding (FLORES-VILLELA
& Zua 1995). It is most active around
dawn. Claudius angustatus is strictly
carnivorous and preys on invertebrates
Claudius angustatus
and small vertebrates. During the rainy
season, some individuals migrate consider-
able distances and can then occasionally
be encountered on land during the day-
time (LEE 1996). After the floods, C. angus-
tatus reproduces. The clutch size varies
from 1 to 8 eggs, which hatch after an
incubation period of 94 to 150 days
depending on environmental conditions
(ERNST & BARBOUR 1989, KOHLER 1997).
The Narrow-bridged musk turtle is

Fig. 48. Claudius angustatus (Tabasco, Fig. 49. Claudius angustatus (Rio Hondo,
Mexico). Photo: J. Haft Quintana Roo, Mexico). Photo: H. Bahena B.

43
 Kinosternidae

Kinosternon
Mud turtles (genus Kinosternon ; about 20
species) are small to medium-sized, pre-
dominantly aquatic chelonians that are
widely distributed over North, Central,
and South America. They are character-
ized by one or two moveable hinges in the
plastron that enable complete closure of
the shell. The Central American species of
Mud turtles live aquatically, mostly in
ponds, rivers, and marshes from sea level
to slightly more than 1000 m elevation.
Frequently, individuals can be encoun-
tered on land when they wander from one Fig. 51. Kinosternon acutum (El Peten,
body of water to another. On the morning Guatemala). Photo: G. Kohler
of 20 April 1990, during a · stay at
Tortuguero, Costa Rica, I observed a spec-
imen of Kinosternon leucostomum in a
clearing within the rain forest. When a
mud turtle is lifted, it usually retreats into
its shell and closes the shell completely
with the aid of the plastron hinges (Fig.
34). After a while, some individuals might
stick out their heads and attempt to bite
whoever holding it. This is often accom-
panied by the excretion of a malodorous
fluid from their cloaca} bladders.

In regions with a pronounced annual cli-

matic cycle, mud turtles spend the dry sea-
son estivating while buried in mud (e.g.,
Fig. 52. Kinosternon acutum (El Peten,
Guatemala). Photo: G. Kohler

Kinosternon creaseri in the northern

Yucatan Peninsula; LEE 1996).
Kinosternids feed mostly on aquatic inver-
tebrates, such as snails, worms, and in-
sects. Some species (e.g., K leucostomum)
also consume a large quantity of plant
material (seeds, fruits, flowers, and leaves);
however, whenever available, they also
prey on small vertebrates (fishes, frogs,
lizards, and snakes) including carrion.
During the reproductive season, the fe-
male produces several clutches, which,
Fig. 50. Kinosternon angustipons (Los depending on the species, contains one to
Guatuzos, Nicaragua). Photo: J. Sunyer ten eggs (KOHLER 1997).
44
 Kinosternidae

Kinosternon acutum GRAY 1831, Synops.

Rept. Cataphracta 1: 34; type locality: un-
known. Carapace length to 120 mm. Central
Veracruz, Mexico, through northern
Guatemala to Belize, sea level to 200 m ele-
vation.
Kinosternon angustipons LEGLER 1965, Univ.
Kansas Puhl. Mus. Nat. Hist. 15 (13): 617;
type locality: Los Diamantes, Limon
Province, Costa Rica. Carapace length to 120
mm. Southeastern Nicaragua to northern
Panama, sea level to 260 m elevation.
Fig. 53. Kinosternon creaseri (Calakmul,
Kinosternon creaseri HARTWEG 1934, Occ. Campeche, Mexico). Photo: R. Cedeno V.
Pap. Mus. Zool., Univ. Michigan 277: 1; type
locality: "one mile south of the Hacienda,
Chichen Itza, Yucatan," Mexico. Carapace
length to 121 mm. Northern portion of
Yucatan Peninsula, sea level to 100 m eleva-
tion.
Kinosternon leucostomum DUMERIL & BIBRON
1851, Cat. Method. Coll. Rept. : 17; type locali-
ty: N. Orleans [in error]; Mexique; Rio
Usumacinta. Carapace length to 174 mm.
Central Veracruz, Mexico, through Central
America to northern South America, sea level
to 1500 m elevation. Two subspecies:
Kinosternon l. leucostomum (Veracruz ,
Mexico, to Nicaragua) and K. l. postinguinale Fig. 54. Kinosternon leucostomum (Rio San
COPE 1887 (Nicaragua to Ecuador). Juan, Nicaragua). Photo: G. Kohler
Kinosternon scorpioides (LINNAEUS 1766),
Systema Naturae, ed. 12: 352; type locality:
Surinam. Carapace length to 270 mm.
Southern Tamaulipas, Mexico, through
Central America to South America, sea level
to 1500 m elevation. The subspecies K. s. scor-
pioides (Panama southward), K. s. abaxillare
BAUR 1925 (Central Depression of Chiapas),
K. s. albogulare DUMERIL & BOCOURT 1870)
(Honduras to Panama), K. s. cruentatum
DUMERIL & B!BRON 1851 (Mexico to
Honduras) occur in Central America.

• Kinosternon creaseri

"" Kinosternon acutum

• Kinosternon scorpioides

45
 Kinosternidae

Fig. 55. Kinosternon scorpioides (near Porte- Fig. 56. Kinosternon leucostomum (Tortuguero,
golpe, Guanacaste, Costa Rica). Costa Rica). Photo: G. Kohler
Photo: H. Bringsoe

Key to Kinosternon
1 a Bridge very narrow, its width less than
21% of carapace length ......... ....... ............. .
. . . .. . . . . . . .. . . . .... Kinosternon angustipons anterior
b Bridge less narrow, its width more than moveable
21% of carapace length .... ............ ............ 2 lobe
2 a Length of gular scute about half the
length of anterior moveable lobe of plas- fixed lobe
tron (Fig. 57a,b) .......... .. ....... .... ....... ... . 3
b Length of gular scute distinctly less than
half the length of anterior moveable lobe posterior
moveable
ofplastron (Fig. 57c,d) .. ...... ......... ..... .... 4 lobe
3 a Anterior moveable lobe of plastron as long
as or shorter than fixed lobe (Fig. 57a) b. K. creaseri
a. K. acutum
... .. .... ..... ... ......... .. Kinosternon acutum
b Anterior moveable lobe of plastron con-
spicuously longer than fixed lobe (Fig.
57b) ... ....... ....... ... Kinosternon creaseri
4 a Anterior moveable lobe of plastron con-
spicuously longer than fixed lobe (Fig.
57c); axillary scute usually not in contact axillary scute
with inguinal scute; carapace never with
more than one keel; a pale postorbital
stripe usually present (Figs. 56 and 58) .....
..................... Kinosternon leucostomum inguinal scute
b Anterior moveable lobe of plastron as long
as fixed lobe (Fig. 57d); axillary scute
usually in contact with inguinal scute;
carapace usually with three keels; no pale c. K. Jeucostomum d. K. scorpioides
postorbital stripe ................... .. ...... ...... ... .
. . . .......... ... ..... . Kinosternon scorpioides
Fig. 57. Plastron of Kinosternon species.
Axillary and inguinal scutes orange.
Further Reading Adopted after LEE (1996).
IVERSON 1976 (Kinosternon in Belize); IVERSON
1980a, b, 1983b

46
 Kinosternidae

Fig. 58. Kinosternon leucostomum (Rio San Fig. 61. Kinosternon scorpioides (Drake Bay,
Juan, Nicaragua). Photo: J . Sunyer Panama). Photo: G. Kohler

Fig. 59. Kinosternon scorpioides (Morgans Fig. 62. Kinosternon scorpioides (Gualaca,
Rock, Nicaragua). Photo: J . Sunyer Chiriqui, Panama). Photo: M. Lundberg

Fig. 60. Ventral view of Kinosternon angustipons (left; Los Guatuzos, Nicaragua) and K scorpioides
(right, Morgans Rock, Nicaragua). Photos: J. Sunyer

47
 Kinosternidae

Staurotypus

The genus Staurotypus includes two spe-

cies, the Pacific coast giant musk turtle
(S. salvinii) and the Mexican giant musk
turtle (S. triporcatus), both of which occur
in Central America. These are aggressive
turtles that need to be handled with great
caution. With their strong, sharp jaws,
these animals can cause serious bite
wounds. The strongly developed dorsal
keels help to stabilize the carapace, there-
by making it more difficult for predators
(e.g., Crocodylus moreletii) to break the
strong shell. The species of this turtle
genus are strictly aquatic inhabitants of Fig. 64. Staurotypus triporcatus (Tabasco,
lakes and slow moving rivers with soft Mexico). Photo: J. Haft
bottoms and rich aquatic vegetation.
Giant musk turtles are not great swim-
mers; these animals prefer to walk around
on the bottom of their aquatic habitat. Staurotypus saluinii GRAY 1864, Proc. Zool.
They are carnivorous, feeding on crabs, Soc. London 1864: 127; type locality:
Huamuchil, Guatemala. Carapace length to
worms, snails, aquatic insects, amphib- 250 mm. Pacific versant of northern Central
ians, and fishes. Additionally, smaller America from Oaxaca, Mexico, to El Salvador,
turtles (especially of the genus Kino- sea level to 700 m elevation.
sternon) are preyed upon by the giant Staurotypus triporcatus (WIEGMANN 1828),
musk turtle and their shells broken with Isis von Oken 21: 364; type locality: Rio
the aid of their strong jaws (CAMPBELL Alvarado, Veracruz, Mexico. Carapace length
1998a). During the reproductive season, to 379 mm. On Caribbean versant from
Veracruz, Mexico, through the base of
the female buries several clutches, each Yucatan Peninsula to western Honduras, sea
with three to ten eggs, on the riverbank. level to 300 m elevation.
The hatchlings open the eggshells after
120 to 207 days of incubation. At that
time, they have a shell length of 25 to 31
mm (8. salvinii) or 30 to 35 mm (S. tripor- • Staurotypus triporcatus
• Staurotypus salvinii
catus) (ERNST & BARBOUR 1989, KOHLER
1997).

Fig. 63. Staurotypus saluinii (ventral view;

near lztapa, Escuintla, Guatemala).
Photo: G. Kohler

48

Key to Staurotypus
1 a Dorsolateral keels extend the length of
carapace from anterior to posterior mar-
ginals (Fig. 65a); anterior moveable lobe of
plastron shorter than posterior moveable
lobe ................. Staurotypus triporcatus
b Dorsolateral keels do not extend the
length of carapace, but end on 1st and 4th
pleurals (Fig. 65b); anterior moveable lobe
of plastron longer than posterior moveable
lobe .... ....... .......... . Staurotypus salvinii
Further Reading
DEAN & BICKHAM 1983, IVERSON 1983a, 1985
a. Staurotypus triporcatus b. Staurotypus sa/vinii
Abb. 65. Carapace with keels (shaded).
Fig. 66. Staurotypus salvinii (near Iztapa,
Escuintla, Guatemala). Photo: G. Kohler
Testudinidae
Fig. 67. Adult Geochelone carbonaria (Darien,
Panama). Photo: H. Birkahn
Testudinidae
Geochelone
The red-footed tortoise (Geochelone carbo-
naria), which occurs in Panama, is the
only South American tortoise that reaches
as far north as southern Central America
(LEGLER 1963). It is otherwise distributed
mostly in tropical South America. The spe-
cies was introduced by humans on several
Caribbean islands, such as Isla de Maiz
Grande, Nicaragua (PRITCHARD & TREBBAU
1984). In Panama, the red-footed tortoise
is found mostly in forested areas and does
not occur in the humid savannas (LEGLER
1963). These diurnal animals dig burrows
in the ground into which they retreat at
night. Red-footed tortoises feed mainly on
grasses, flowers, fallen fruit, and carrion.
In Panama, this tortoise is hunted and
eaten by the local people, as is the case in
other parts of its range to the south. The
egg-laying season is from June through
September. During this period, the fe-
males bury their clutches with 2 to 15 eggs
in the ground. Females probably produce
several clutches per season. The eggs are
somewhat elongate (40-59 mm x 34-48
mm) and have a rigid shell. After an incu-
bation period of 116 to 185 days, the ju-
49
 Testudinidae

veniles hatch; they have a shell length of

39 to 45 mm (ERNST & BARBOUR 1989,
KOHLER 1997). Reproductive maturity is
reached at a shell length of 200 to 250 mm
(LEGLER 1963).

Geochelone carbonaria (SPIX 1824), Anim.

Spec. Nov. Testud. Brasil.: 22; type locality:
"Habitat sub cognomine 'Capitary' (?) ad flu-
men Amazonum," South America. Carapace
length to 51 cm. Southeastern Panama
through most of the Amazon Basin south-
ward to Rio de Janeiro and westward to
Paraguay and to northern Argentina, sea
level to 300 m elevation.

Further Reading
LEGLER 1963; ERNST & LEUTERITZ 1999

Fig. 69. Hawksbill sea turtle (Eretmochelys

imbricata) at a coral reef. Photo: J . Pichler

Sea Turtles
(Cheloniidae, Dermochelyidae)
Sea turtles were already to be found around
the world during the Cretaceous period,
with fossil finds dating back at least 200
million years. Thus, the ancestors of our
modern-day sea turtles lived side-by-side
with the dinosaurs, but were able to
outlive the gigantic Plesiosaurus and
Ichthyosaurus. The modern species came
into existence around 10 - 60 million
years ago and comprise, along with marine
iguanas and sea snakes, the only marine
reptiles. All species of sea turtles have
forelimbs that have evolved into paddle-
shaped fins, with only one or two claws.
Further adaptations to the aquatic life
style are salt glands, by means of which
superfluous salt, which has been ingested
through feeding in sea water, is expelled.
Fig. 68. Juvenile Geochelone carbonaria These animals can reach adult weight of
(Cordoba, Colombia). Photo: M. Lundberg between 35 and 500 kg.
50
 Cheloniidae, Dennochelyidae

The females crawl up onto certain beaches

at dusk, in order to dig a nest and deposit
their eggs. After filling in the nest, the
female returns to the sea during the same
night. The young sea turtles hatch after an
incubation period of approximately two
months and abandon the nest together for
the sea during the night. After attaining
sexual maturity a number of years later,
the adult females return to the original
beach for nesting. However, this generali-
zation of a sea turtle's life cycle only applies
in a general way. The eight known species
of sea turtles vary considerably with Fig. 70. Green turtles (Chelonia mydas)
regard to their geographical location, copulating. Photo: J. Pichler
mating cycles, and behavior. Moreover, the
various populations of a particular species
exhibit a number of differences due to geo-
graphical distribution. The following in-
formation about the nesting seasons of the
individual species relates in each case to
Central America. Six species of sea turtles
from two families and five genera nest on
the beaches of Central America. There
used to be mass landings of sea turtles (so-
called "arribadas" or "arribazones") in
many Central American countries (Costa
Rica, Honduras, Nicaragua, and Panama
to name a few), in which thousands of
females of the species Lepidochelys oliva-
cea and others came onto land all at once
in order to lay their eggs on one beach. Fig. 71. Lepidochelys oliuacea nesting at Playa
This phenomenon only takes place in Bavaria, Guanacaste, Costa Rica.
Photo: G. Kohler
Costa Rica presently (CORNELIUS 1995).

Among the most important factors that

have led to the rapid decrease in sea turtle
populations are their massive commercial
use (the collection of eggs and killing of
adult turtles), the plundering of nests by
feral animals (dogs and pigs), shrimp
fishing, and sea pollution (cases of death
due to ingestion of tar and plastic refuse).
Turtle products are offered in numerous
tropical and subtropical countries as sou-
venirs for tourists. It is completely ir-
responsible and illegal to sell and abuse
sea turtles or products made from these
animals as souvenirs, in light of their
endangered status. Fig. 72. Sea turtle hatchlings on their
dangerous walk to the sea. Photo: J. Pichler
51
 Cheloniidae, Dennochelyidae
Fig. 73. Green turtle (Chelonia mydas) with
shark suckers. Photo: J. Pichler
The loggerhead sea turtle (Caretta
caretta) is found worldwide, primarily in
subtropical regions, and less in tropical cli-
mates. It is found primarily on the
Caribbean side of Central America. The
most important nesting grounds of this
species are located in Florida and South
Carolina in the U.S., while in Central
America few specimens come onto land to
lay their eggs, mostly in the northeastern
portion of the Yucatan Peninsula. In
Caretta caretta, egg laying has been docu-
mented from April to July. After a single
short mating phase, a female produces 2 -
5 clutches, each with 40 - 190 eggs at
intervals of approximately two weeks.
From these, the young will hatch after 55-
68 days of incubation (MAGNUSON et al.
1990, MARQUEZ 1990).
While it was earlier proposed that two spe-
cies of the genus Chelonia (green turtles)
populated the coastal regions of Central
America (C. mydas on the Caribbean side
and C. agassizii on the Pacific side), it is
currently believed that only a single spe-
cies is represented in the genus (ERNST &
BARBOUR 1989, KARL & BOWEN 1999). The
shell of C. mydas can measure up to 140
cm in length. Important nesting grounds
52
Fig. 74. Loggerhead sea turtle (Caretta
caretta). Photo: R. D. Bartlett
are located along the coast of the Mexican
state of Michoacan, as well as in El
Salvador (Jiquilisco), Guatemala, and
Costa Rica (Tortuguero). Single females
come on land in many areas along the
Central American Pacific and Caribbean
coasts to nest. During one nesting season,
which can last from August until January
(Pacific side) or April until October
(Caribbean side), a female can produce 2 -
5 clutches (rarely up to 8) at intervals of 12
- 14 days, each with 38 - 120 (rarely up to
195) eggs, from which the young hatch
after 50 - 75 days (MAGNUSON et al. 1990,
MARQUEZ 1990). The animals reach sexual
maturity between eight and thirteen years
of age. A female does not come on land
every year for nesting, rather every 2 - 5
years. Green turtles subsist mainly on
vegetarian fare (various sea algae and sea
grass), but also feed upon sponges, mol-
luscs, jellyfish, and fish.
The hawksbill sea turtle (Eretmochelys
imbricata) is found around the world in all
tropical seas and on both the Pacific, as
well as the Caribbean side of Central
America. Important Central American
nesting grounds for this species are found
on the coast of the Yucatan Peninsula, as

Fig. 75. Kemp's ridley sea turtle (Lepidochelys
kempii) swimming in shallow water.
Photo: R. D. Bartlett
well as near Tortuguero (Costa Rica),
while nesting specimens have only been
sporadically reported elsewhere. The egg-
laying season is from April until
November, but it varies according to re-
gion: Mexico and Belize - April to August;
Guatemala- May to November; Honduras
- June to September; Nicaragua - May to
October; Costa Rica - May to November
(MARQUEZ 1990). During one season, each
female comes on land once or twice to bury
clutches consisting of 93 - 223 eggs each.
A female only nests every 2 - 3 years. The
incubation period lasts 47 - 75 days and
the young have a shell-length of 38 - 46
mm (MARQUEZ 1990). The hawksbill sea
turtle is mainly carnivorous, and its food
ranges from crabs, sponges, sea anemones,
jellyfish, molluscs, squid, and fish to sea
algae. However, there are apparently
great geographical differences with regard
to its range of foods. According to the in-
vestigations by CARR & STANCYK (1975),
the hawksbill sea turtles in the Tortu-
guero region (Costa Rica) feed primarily
upon certain sponges.
Cheloniidae, Dermochelyidae
Both species of the genus Lepidochelys
(ridley sea turtle) are found on the
coasts of Central America, with the
Kemp's ridley sea turtle (L. kempii) only
found on the Yucatan Peninsula in this
region, and the olive ridley sea turtle (L.
olivacea) along the entire Pacific coast of
Central America. The latter is the most
common species of sea turtle on the Pacific
coast of Central America, with its most
important nesting sites in Nicaragua
(Masachapa, Pochomil, Boquita) and
Costa Rica (Nancite, Ostional) (CORNELIUS
1995). Playa Nancite is a beach in the
Santa Rosa National Park in Costa Rica
that is naturally protected by the sharp
rocks and thorny bushes that surround it.
There, one can experience the spectacular
"arribadas" from July until September, in
which up to 8000 L. olivacea females crawl
up onto the beach to lay their eggs within
a span of a few days. Important nesting
grounds for L. kempii are located along the
Gulf Coast, with the beaches near Rancho
Nuevo (Tamaulipas, Mexico) of by far the
greatest importance. In the 1940's, the
amateur filmmaker Andres Herrera docu-
mented a mass landing of approximately
40,000 L. kempii near Rancho Nuevo
(MAGNUSON et al. 1990). This type of mass
landing is known only in the case of the
species of the genus Lepidochelys. It is
assumed that the simultaneous hatching
of many young increases the chances that
the necessary number of turtles survive, in
spite of numerous predators. Lepidochelys
kempii also differs from the other sea turt-
les in that it comes onto the beach during
the day to lay its eggs, while the other spe-
cies nest at night. Also remarkable is the
fact that the females of both these species
nest nearly every year, producing 2 - 7
clutches, each with 50 - 140 eggs in one
season. After an incubation period of 45 -
65 days, the young hatch with a shell
length of 35 - 45 mm. (MARQUEZ 1990).
Ridley sea turtles feed on crabs and other
invertebrates (sponges, sea anemones, jel-
lyfish, and squid).
53
 Cheloniidae, Dermochelyidae
Fig. 76. Leatherback sea turtle (Dermochelys
coriacea) nesting at Playa Naranjo, Costa Rica.
Photo: M. Franzen
The largest turtle living today is the
leatherback sea turtle (Dermochelys
coriacea), with the largest specimen ever
measured having a shell length of 256.5
cm. and a mass of916 kg. (MARQUEZ 1990).
Most specimens, however, have shells with
a length of 150 - 170 cm. This species can
be found all over the world, as they are
able to populate cooler seas than other sea
turtle species. Leatherback sea turtles
usually stay in the open ocean and are
observed only occasionally near the coast.
They can submerge to great depths.
Important beaches for nesting in Central
America are located in Costa Rica (Playa
Naranjo, Playa Tamarindo, between
Tortuguero and Limon) and Panama.
While the nesting season on the
Caribbean side lasts from April to October,
the nesting season on the Pacific side lasts
from October to March. The females come
on land to nest every 2-3 years so that
they can bury 4-5 clutches of61-126 eggs
each in a single season. The incubation
period is 50-78 days and the young have a
shell length of 51-68 mm (MARQUEZ 1990).
Leatherback sea turtles are mainly carni-
vorous and feed on jellyfish and pelagic
crustaceans, and occasionally also on sea
algae.
54
Fig. 77. Chelonia mydas. Photo: J. Pichler
Cheloniidae
Caretta caretta (LINNAEUS 1758), Systema
Naturae, ed. 10: 197; type locality: "insulas
Americanas." Carapace length to 105 cm.
Worldwide; in Central America mostly in the
Caribbean.
Chelonia mydas (LINNAEUS 1758), Systema
Naturae, ed. 10: 197; type locality: "insulas
Pelagi: insulam Adscensionis." Carapace
length to 140 cm. Worldwide.
Eretmochelys imbricata (LINNAEUS 1766),
Systema Naturae, ed. 12: 350; type locality:
"Mari Americano, Asiatico." Carapace length
to 114 cm. Worldwide.
Lepidochelys kempii (GARMAN 1880), Bull.
Mus. Comp. Zool. 6: 123; type locality: Gulf of
Mexico. Carapace length to 75 cm. Gulf of
Mexico.
Lepidochelys olivacea (ESCHSCHOLTZ 1829),
Zool. Atlas 1: 3; type locality: Manila Bay,
Philippines. Carapace length to 75 cm. World-
wide; in Central America only in the Pacific.
Dermochelyidae
Dermochelys coriacea (LINNAEUS 1766),
Systema Naturae, ed. 12: 350; type locality:
"maris Tyrrheni oram in agro Laurentiano."
Carapace length to 256,5 cm. Worldwide.
 Cheloniidae, Dermochelyidae

Fig. 78. Eretmochelys imbricata. Photo: J. Pichler

"" Caretta caretta

• Lepidochelys kempii

"" Che/onia mydas

55
 Cheloniidae, Dennochelyidae

Key to sea turtles

1 a Shell covered with a continuous layer of
undivided skin (Fig. 80a) ... .. ........ ..... .... .. .
. . . . . .. ....... .. ........ .... Dermochelys coriacea
b Shell covered with distinct horny scutes
(Fig. 80b-f) ................................................ 2
a. Chelonia b. Eretmochelys
2 a Four pleural (= costal) scutes present on
each side (Fig. 80c,d); nuchal (cervical) Fig. 79. Lateral view of head (prefrontal scales
scute not in contact with first pleural pale brown). Note also the hawk-like beak in
scutes .............. ..... .............. ..................... 3 Eretmochelys.
b Five or more pleural(= costal) scutes pres-
ent on each side (Fig. 80e,f); nuchal
(cervical) scute in contact with first pleu-
ral scutes ........................ .... .................... 4 Carapace of sea turtles
3 a One pair of prefrontals; beak not hawk-
like (Fig. 79a); scutes of carapace non-
imbricate .. ................... Chelonia mydas
b Two pairs of prefrontals (Fig. 79b); beak
hawk-like; scutes of carapace imbricate
........ ............... Eretmochelys imbricata
4 a Bridge usually with three inframarginal
scutes, each without pores; three or more
inframandibular scales behind beak on
each side oflower jaw .... Caretta caretta
b Bridge usually with four inframarginal
scutes, each with a pore at its posterior a. Dermoche/ys coriacea b. Caretta caretta
border; single large inframandibular scale
behind beak on each side oflower jaw .... 5
5 a Usually 5 pleural scutes (Fig. 80e); ground
color gray .... ..... ..... Lepidochelys kempii
b Usually 6 or more pleural scutes (Fig. 80£);
ground color olive Lepidochelys olivacea

Further Reading
CARR 1967, HIRTH 1980a, b, PRITCHARD 1980,
MAGNUSON et al. 1990, MARQUEZ 1990, WILSON
& ZUG 1991, CORNELIUS 1995, ZUG & ERNST
1998 c. Chelonia mydas d. Eretmochelys imbricata

e. Lepidochelys kempii f. Lepidoche/ys olivacea

Fig. 80. Carapace of sea turtles (pleural scutes

orange).
.a. Dermochelys coriacea (Fig. 79 and 80 modified after a poster of S.A
ECKERT & P.C.H. PRITCHARD).

56
 Samia

Lizards (Sauria)

There are 236 species oflizards in 10 fami-

lies and 50 genera that occur in Central
America.

Key to families of lizards

1 a Third and fourth toe about same length
..... .. ....... ..... ..... .... ..... Helodermatidae suboculars
b Fourth toe distinctly longer than third toe a. Lateral view
..... ........ ..... .... .. .. .... .... .. .. ....................... 2
2 a Two pairs of scales between rostral plate interparietal plate
and first unpaired snout plate (Fig. 83a)
.. ... ........... ..... .......................... Anguidae
b Fewer than two pairs of scales between
rostral plate and first unpaired snout
plate (Fig. 83b) or snout scales small and
irregular ..... ..... ... .................................... 3
3 a Scales on dorsum and venter of body simi-
lar in shape and size ............ ................... 4
b Scales on dorsum and venter of body dif-
ferent in shape and size, ventral scales
usually larger than dorsal scales ............ 5 b. Dorsal view

4 a No eyelids present ........................... ........ . Fig. 82. Head scalation characters important
............... Gymnophthalmidae (in part) in the identification of lizards .
b Eyelids present ........... .. .... .... Scincidae
5 a Venter covered with large plates (Figs. 81
b-d) ............. ....... ... ........................... ...... 6
b Venter covered with small scales (Fig.
Sia) ... ..... .. ..... .... .... ..... .. ... .... ..... .... .... 8

a. Ce/estus b. Mabuya

Fig. 83. Scalation of dorsal surface of head.

a. Dactyloa latifrons b. Aspidoscelis deppii

Fig. 84.
Scalation of
dorsal surface
of head (rostral
brown; fronto-
c. Mesoscincus managuae d. Mabuya unimarginata nasal orange).

Fig. 81. Ventral scales in lizards. a. Ameiva b. Anadia

57
 Sauria

6 a No eyelids present ............. .. Xantusiidae

b Eyelids present ................ ..... ..... .. ....... 7
7 a Intemasals present, rostral separate from
frontonasal (Fig. 84a) ................... Teiidae
b Without intemasals, rostral in contact
with frontonasal (Fig. 84b) ........................
............... Gymnophthalmidae (in part)
8 a Dorsal surface of head covered with sub-
equal granular scales .... .... .............. ........ 9
b Dorsal surface of head covered with non-
granular scales of different sizes .......... 10
9 a No eyelids present .. .... ......... Gekkonidae
b Eyelids present ..... ...... ... Eublepharidae
lOa Dorsum of body with conspicuous beaded
scales ................................ Xenosauridae
b Dorsum of body without conspicuous
beaded scales ................... ....... lguanidae
Fig. 87. Abronia anzuetoi (southern slope of
Volcan de Agua, Escuintla, Guatemala).
Photo: J. A. Campbell

Anguidae
Anguids are cosmopolitan, with six genera
occurring in Central America. Most alli-
gator lizards have square scales that are
arranged in rings and strengthened by
bony dermal plates (osteoderms). While
representatives of the genera Mesapis and
Gerrhonotus are found more commonly,
the other species are known from only a
few specimens. This could be due to the
fact that the "rare" species live in habitats
(i.e., treetops) where, for the most part,
they evade our observation.
a. Diploglossus
Fig. 85. Lateral view of toe (scaly sheath
shaded).
Fig. 86. Lateral
head in Mesaspis
(preocular brown
and suboculars
orange).
Key to genera of Anguidae
1 a Body without distinct longitudinal fold .. 2
b Body with a distinct longitudinal fold .... 3
2 a Claws almost completely covered by scales
(Fig. 85a) ........ .. ............. ..... Diploglossus
b Claws not covered by scales (Fig. 85b)
...................................................... Celestus
3 a Longitudinal fold between ear opening
and shoulder reduced or absent; head
more or less flattened, often helmet-like
................. ......... ...... ...... .... .......... Abronia
b Longitudinal fold between ear opening
and shoulder conspicuously developed;
head not flattened, never helmet-like .... 4
4 a Suboculars differentiated from preoculars
and postoculars (Fig. 86), not in a single
continuous series; three pairs of large and
one pair of small chin shields .... ... .. ..
................ ........ ....... ........... .... ...... Mesaspis
b Suboculars, preoculars and postoculars in
a single continuous series of similar sca-
les; four or five pairs of large chin shields
················ ····················· ······················ ······· 5
5 a 10 longitudinal rows of ventral scales; 12
nuchals; five pairs of large chin shields;
usually two postrostrals ........ Coloptychon
b 12 or more longitudinal rows of ventral
scales; 10 nuchals; four pairs of large chin
shields; 0-1 postrostrals ........ Gerrhonotus

58
 Anguidae

Abronia

The tree-dwelling alligator lizards of the

genus Abronia are distributed in Mexico
and northern Central America. While
most species live in highlands between
1500 and 3000 m above sea level in pine,
oak, or cloud forest, the species of the
bogerti group (A bogerti, A. chizari, A.
ramirezi) are inhabitants of more tropical
pre-montane habitats between 360 and
1400 m above sea level (H. SMITH & SMITH
1981, CAMPBELL 1994). Abronia are diur-
nal and stay mostly up to 40 m above Fig. 88. Abronia aurita (El Quiche,
ground level in trees that are overgrown Guatemala). Photo: J. A. Campbell
with epiphytes, where they can climb
among the bromeliads, ferns, and orchids
(CAMPBELL & FROST 1993). The long pre-
hensile tail serves the alligator lizard as a
fifth leg and, thus, constiti,ites an impor-
tant climbing aid. Occasionally, however,
the animals will descend to the ground,
where they have been found beneath piec-
es of tree bark and tree trunks, and also
sitting openly on the leaf- or needle-cover-
ed ground (BOGERT & PORTER 1967'
CAMPBELL & FROST 1993, FRANZEN & HAFT
1999, MCCRANIE pers. comm. 1997). These
lizards have a calm nature. CAMPBELL &
FROST (1993) reported a gravid female A.
fimbriata that they observed in the quiet Fig. 89. Abronia bogerti (Cerro Baul, Oaxaca,
inlet of a small mountain stream. All spe- Mexico). Photo: T. Bille
cies of the genusAbronia, for which repro-
ductive data exist, are live-bearing, with
the number of young per litter varying
between one and twelve (WERLER 1951,
MARTIN 1955, H. SMITH & .ALVAREZ DEL
TORO 1962, ALVAREZ DEL TORO 1973,
CAMPBELL & FROST 1993). The coloration of
the juveniles varies from species to spe-
cies, but consists mostly of dark lateral
dorsal stripes on a pale background
(CAMPBELL & FROST 1993). Due to their
restricted geographical ranges and the
risk of habitat destruction, most species of
Abronia are considered amongst the most
endangered species of lizards anywhere.
One can assume that some species have
already become extinct before they could even Fig. 90. Abronia campbelli (Jalapa,
be discovered (CAMPBELL & FROST 1993). Guatemala). Photo: J. A. Campbell

59
 Anguidae

Abronia anzuetoi CAMPBELL & FROST 1993,

Bull. Amer. Mus. Nat. Hist. 216: 22; type
locality: Finca Rosario Vista Hermosa, 1219
m elevation, on S slope of Volcan de Agua,
Escuintla, Guatemala. SVL to 135 mm.
Southern slopes of Volcan de Agua in south-
ern central Guatemala, 1220-2290 m elevati-
on in cloud forest.
Abronia aurita (COPE 1869), Proc. Acad. Nat.
Sci. Philadelphia 20: 306; type locality: Forest
of Verapaz, Guatemala. SVL to 125 mm.
Central highlands of Guatemala, 2000-2660
m elevation in pine-oak forest. A female gave
birth to 12 juveniles between 26 and 28
January 1992 (CAMPBELL & FROST 1993). Fig. 93. Abronia fimbriata (Alta Verapaz,
Guatemala). Photo: J. A. Campbell

Abronia bogerti TIHEN 1954, Amer. Mus.

Novitates 1687: 3; type locality: N ofNiltepec,
between Cerro Atravesado and Sierra Madre,
Oaxaca, Mexico. SVL to 64 mm. Eastern
Oaxaca, Mexico, 760-1370 m elevation in rain
forest and lower cloud forest.
Abronia campbelli BRODIE & SAVAGE 1993,
Herpetologica 49: 421; type locality: Cerro
Tabl6n de las Minas near Pastoria, 14°46'N,
89°57'W, 1800-1900 m elevation, Jalapa,
Guatemala. SVL to 127 mm. Eastern
Guatemala, 1800-1900 m elevation in pine-
oak forest.
Abronia fimbriata (COPE 1885), Proc. Amer.
Phil. Soc. 22: 771; type locality: Pine forest of
Fig. 91. Abronia gaiophantasma (Baja Alta Verapaz, Guatemala. SVL to 130 mm.
Verapaz, Guatemala). Photo: J. A. Campbell Central-eastern Guatemala, 1400-2100 m
elevation in cloud forest. A female gave birth
to four juveniles (33 mm SVL) on 15 June
(CAMPBELL & FROST 1993).
Abronia frosti CAMPBELL, SASA, ACEVEDO &
MENDELSON 1998, Herpetologica 54: 222; type
locality: along road to Patalcal, 2835 m eleva-
tion, 15°51'19"N, 91°31'02"W, 5.9 km (by
road) NW intersection of Guatemala Road 9N
(near San Mateo Ixtatan), Sierra de Los
Cuchumatanes, Huehuetenango, Guatemala.
SVL to 101 mm. Northwestern Guatemala,
2835 m elevation in cloud forest.
Abronia gaiophantasma CAMPBELL & FROST
1993, Bull. Amer. Mus. Nat. Hist. 216: 19;
type locality: vicinity of La Union Barrios,
1600 m elevation, on W slope of Cerro Verde,
Baja Verapaz, Guatemala. SVL to 110 mm.
Central-eastern Guatemala, 1600-2350 m
elevation in pine and cloud forest.
Fig. 92. Abronia lythrochila (near San Abronia leurolepis CAMPBELL & FROST 1993,
Crist6bal de las Casas, Chiapas, Mexico). Bull. Amer. Mus. Nat. Hist. 216: 35; type lo-
Photo: G. Kohler
60
 Anguidae

cality: Santa Rosa, near Comitan, Chiapas,

Mexico. SVL to 105 mm. Only known from
the type locality, Santa Rosa, in the vicinity of
Comitan, in eastern Chiapas, Mexico, 1800-
2300 m elevation in cloud forest.
Abronia lythrochila SMITH & ALVAREZ DEL
TORO 1963, Herpetologica 19: 100; type lo-
cality: Nachij, between Tuxtla Gutierrez and
San Cristobal Las Casas, Chiapas, Mexico.
SVL to 106 mm. Central Chiapas, Mexico,
2000-3000 m elevation in pine-oak forest. A
copulation that took place on the ground and
lasted at least 45 minutes was observed on 2
August 1976 (CAMPBELL & FROST 1993). There
are 1-5 juveniles per litter, which are born
from June to August (SMITH & ALVAREZ DEL Fig. 95. Abronia matudai (Aguas Georginas, Quetz-
TORO 1962, ALVAREZ DEL TORO 1973, 1983). altenango, Guatemala). Photo: L. Melendez
Abronia matudai (HARTWEG & TIHEN 1946),
Occ. Pap. Mus. Zool., Univ. Michigan 497: 3;
type locality:Volcan Tacana, 2000 m eleva-
tion, Chiapas, Mexico. SVL to 96 mm.
Southwestern Guatemala and southeastern
Chiapas, Mexico, 1950-2630 m elevation in
cloud forest.
Abronia meledona CAMPBELL & BRODIE 1999,
Herpetologica 55: 163; type locality: near the
Torre de Guatel, near the aldea of Soledad
Grande, Jalapa, Guatemala. SVL to 120 mm.
Southeastern Guatemala, 2200-2660 m in
cloud forest.
Abronia montecristoi HIDALGO 1983, Occ. Pap.
Mus. Nat. Hist. Univ. Kansas 105: 6; type
locality: Hacienda Montecristo, 2250 m,
Cordillera de Alotepeque-Metapan, Santa
Ana, El Salvador. SVL to 90 mm. National
park Montecristo at the juncture of El
Salvador-Honduras-Guatemala and at Fig. 96. Abronia meledona (Jalapa, Guatemala).
Quebrada Grande in western Honduras, Photo: J.A Campbell
1370-2250 m elevation in cloud forest.

Fig. 94. Abronia lythrochila (Chiapas, Mexico). Fig. 97. Abronia montecristoi (Quebrada Grande,
Photo: A. Ramirez Copan, Honduras). Photo: J . R. McCranie
61
 Anguidae

Abronia ochoterenai (MARTIN DEL CAMPO

1939), Anal. Inst. Biol. Mexico 10: 357; type
locality: Santa Rosa, Comitan, Chiapas,
Mexico. SVL to 97 mm. Central Chiapas,
Mexico, 1800-2300 m elevation in cloud
forest.
Abronia ornelasi CAMPBELL 1984,
Herpetologica 40: 373; type locality: Cerro
Baul, Oaxaca, Mexico. SVL to 97 mm.
Eastern Oaxaca, Mexico, 1500-1600 m eleva-
tion in cloud forest.
Abronia ramirezi CAMPBELL 1994,
Herpetologica 50: 2; type locality: Rancho El
Recuerdo, 1350 m elevation, Cerro La Vela,
Sierra Madre de Chiapas, Municipio de
Jiquipilas, Chiapas, Mexico. SVL to 93 mm.
Cerro La Vela, Sierra Madre de Chiapas, Fig. 98. Abronia ornelasi (Oaxaca, Mexico).
western Chiapas, Mexico, 1350 m elevation in Photo: J . A. Campbell
cloud forest.
Abronia salvadorensis HIDALGO 1983, Occ.
Pap. Mus. Nat. Hist. Univ. Kansas 105: 1; type
locality: 10 km NE Perquin, Cordillera de
Nahuaterique, Honduras. SVL to 94 mm.
Sierra de Montecillos and Sierra de Opalaca
in Honduras, 1900-2250 m elevation in cloud
forest.
Abronia smithi CAMPBELL & FROST 1993,
Bull. Amer. Mus. Nat. Hist. 216: 30; type lo-
cality: cloud forest at 2020 m elevati,m on SE
slope of Cerro El Triunfo, Sierra Madre de
Chiapas, Chiapas, Mexico. SVL to 103 mm.
Cloud forest of the Sierra Madre de Chiapas
in southeastern Chiapas, Mexico, 1800-2800
m elevation. There are 3-4 juveniles per litter
(ALVAREZ DEL TORO 1973, CAMPBELL & FROST
1993).
Abronia vasconcelosii (BOCOURT 1871), Bull. Fig. 99. Abronia ramirezi (Cerro La Vela,
Nouv. Arch. Mus. Hist. Nat. Paris 7 (4): 107; Chiapas, Mexico). Photo: A. Ramirez V.
type locality: Arguetta [=Argueta, according
to CAMPBELL & FROST 1993], above 2000 m,
Guatemala. SVL to 123 mm. Guatemala
Plateau above 2200 m elevation in cloud
forest.

Abronia

"' A. anzuetoi
"'A. aurita
•A. bogerti
~'-r"----' • A. campbelli
6 A. fimbriata
\1 A. frosti
o A. gaiophantasma
O A. leurolepis
• A. vasconce/osii Fig. 100. Abronia salvadorensis (Zacate Blanco,
Intibuca, Honduras). Photo: J . R. McCranie
62

Key to Abronia
1 a 10 longitudinal ventral seal~ rows . . ·····:
.............................. ..... Abronia ramire:z:i
b 12 or more longitudinal ventral scale rows
·· ······················· ······· ·············· ············ ······ 2
2 a Conspicuous supra-auricular spines pre-
sent ........................................................... 3
b Without supra-auricular spines ..... ............
.............. ....... ............ ... ........... ... ............ 13
3 a Supranasals in contact with each other,
thereby separating anterior and posterior
pairs of internasals; frontonasal scale
absent ...................... Abronia fimbriata
b Supranasals not in contact with each
other, therefore anterior and posterior
pairs of internasals in contact; frontonasal
scale present or absent .. ... .................. ..... 4
4 a 12 longitudinal ventral scale rows .......... 5
b 14-16 longitudinal ventral scale rows ... . 7
5 a Frontonasal absent; dorsal body scales with-
out keels; posteriormost infralabial conspic-
uously elongate ...... ... Abronia leurolepis
b Frontonasal present; dorsal body scales
keeled; posteriormost infralabial not con-
spicuously elongate ........... ..................... . 6
6 a Posterior subocular in contact with lower
anterior temporal; preauriculars granular
and in several rows; three primary temp-
orals; no bright yellow ring around eye
present .. ............ .. Abronia ochoterenai
b Posterior subocular not in contact with
lower anterior temporal; preauriculars
usually subimbricate to imbricate and in
two rows; usually 4-5 primary temporals;
a bright yellow ring around eye present,
this color strikingly different from that of
rest of head .................... . Abronia smithi
7 a Circumorbital region not colored different-
ly from rest of head ............... ................... 8
Abronia
•A. matudai
"" A. meledona
• A montecristoi
• A.ochoterenai
!:::. A.
ornelasi
'V A.ramirezi
• A. salvadorensis
O A. smithi
• A. lythrochila
Anguidae
Fig. 101. Abronia smithi (El Triunfo, Chiapas,
Mexico). Photo: A. Ramirez V.
b A bright ring around eye present, this
color strikingly different from that of rest
of head ........... ........... .. .......................... ... 9
8 a 14-15 longitudim'\l dorsal scale rows; dor-
sal color variable ... Abronia lythrochila
b 12 longitudinal dorsal scale rows; dorsum
brownish ....... Abronia gaiophantasma
9 a 32-35 transverse ventral scale rows;
ground color gray to brown; circumorbital
region pale brown to cream color ...............
................................... Abronia campbelli
b 35-38 transverse ventral scale rows;
ground color green; circumorbital region
yellow to orange ................... ............... ... 10
lOa Dorsal body scales only sparsely pigmen-
ted with black .. .......... Abronia anzuetoi
b Dorsal body scales heavily pigmented with
black, anterior portion of scales almost
entirely dark ..... ... ........... ... .. ... ........... 11
Ila 9-11 supra-auricular spines ......... ..... ... .
. . . . . . . . . . . . . . . . . . . . . ........ .. Abronia meledona
b 4-7 supra-auricular spines .......... ... ...... 12
12a Dorsal body scales green in life with black
pigment ... .................... .. .. Abronia aurita
b Dorsal body scales yellowish green to tur-
quoise in life with black pigment
............... .. ............ Abronia vasconcelosii
13a Supranasals very large and in contact
with each other, thereby separating anter-
ior from posterior pairs of internasals;
posterior and dorsal edges of dorsal scales
distinctly pale colored Abronia ornelasi
b Supranasals, if present, small and not in
contact with each other, therefore anterior
63
 Anguidae

and posterior pairs of internasals in contact;

coloration of dorsal scales variable ... ....... 14
14a Parietal in contact with supraoculars ... 15
b Parietal scale not in contact with supra-
oculars ....................... ...................... ...... 17
15a Dorsal body pattern consists of distinct
alternating white or yellow transverse
bars; 12 longitudinal dorsal scale rows; 28-
32 transverse dorsal scale rows; 33-35
transverse ventral scale rows ... ... .. .... .. .
. . . . . . . . ............ ... .... . . . . . . . . . . .. Abronia frosti
b Dorsal body pattern different from that
indicated above; 14-16 longitudinal dorsal
scale rows; 33-41 transverse dorsal scale
rows; 36-55 transverse ventral scale rows Fig. 103. Abronia uasconcelosii (Cerro Alux,
............................. .. .................................. 16
Guatemala). Photo: G. Kohler
16a Supra-auricular scales slightly projecting
but not spine-like; prefrontal not in
contact with superciliaries; 4 secondary
and 4 tertiary temporals; 6 rows of nuchal
scales ........................... Abronia matudai
b Supra-auricular scales not differentiated;
prefrontal in contact with superciliaries; 3
secondary and 3 tertiary temporals; 8
rows of nuchal scales ..... ....... .. ..... ..... .
. . .. .... ... . . . . . . .............. .. ..... Abronia bogerti
17a 5 occipitals; 3 primary temporals in contact
with postoculars ... Abronia montecristoi
b 1-3 occipitals; 2 primary temporals in contact
with postoculars .... Abronia salvadorensis
Fig. 104. Abronia uasconcelosii (hatchling).
Photo: L . Melendez
secundary temporals primary temporals
Further Reading
TIHEN 1954, CAMPBELL & FROST 1993, KOHLER
supranasal 1996c, CAMPBELL et al. 1998, CHIPPINDALE et al.
supralabials 1998, MCCRANIE & WILSON 1999

prefrontal frontonasal

anterior internasal
rostral

occipitals

Fig. 102. Scalation on lateral and dorsal surface of Fig. 105. Celestus montanus (Cusuco,
head in Abronia montecristoi. Honduras). Photo: J. Kolby
64
 Anguidae

Celestus Celestus atitlanensis SMITH & Taylor 1950,

Bull. U.S. Natl. Mus. 199: 195; type locality:
Atitlan, Guatemala. SVL to 113 mm. Pacific
13 species of Mexican and Central versant of Guatemala to western El Salvador,
American anguids (11 of them in Central 700-1650 m elevation.
America) share the common trait of a Celestus bivittatus (BOULENGER 1894b), Proc.
cylindrical body without a longitudinal Zool. Soc. London 1894: 732; type locality:
fold. These species are placed into two Hacienda Rosa de Jericho [Santa Rosa de
genera (Celestus and Diploglossus) by Jerico], 3250 ft, Matagalpa, Nicaragua. SVL
to 103 mm. Eastern Guatemala to central
some authors (PETERS & DONOSO-BARROS Nicaragua, 990-1980 m elevation in pine-oak
1970, SAVAGE & LIPS 1994, MCCRANIE & forest.
WILSON 1996), while others (CAMPBELL &
Celestus cyanochloris COPE 1894, Proc. Acad.
CAMARILLO 1994) combine all these species Nat. Sci. Philadelphia 46: 200; type locality:
in the genus Diploglossus. Until more is Volcan Irazu, Costa Rica. SVL to 99 mm.
known about the evolutionary relation- Cordilleras de Tilaran and Central, Costa
ships of these species, I believe that it is Rica, 1200-1710 m elevation.
best to retain the two genera concept. Of Celestus hylaius SAVAGE & LIPS 1994, Rev.
the ten described species of the genus Biol. Trop. 41: 827; type locality: La Selva
Celestus, eight are found in Central Biological Station, 40 m elevation, Canton de
Sarapiquf, Heredia, Costa Rica. SVL to 107
America, whereas two (C. legnotus and C. mm. Caribbean versant of Costa Rica in low-
enneagrammus) are found further north- land rain forest, 40-430 m elevation.
west in Mexico. While most Celestus spe-
cies are highlands inhabitants that live
more than 1000 m above sea level in the
pine-oak forest, C. cyanochloris, C. hylaius,
and C. rozellae are found in lowland rain
forest. Some species are obvious tree-pwel-
lers (i.e., C. atitlansis, C. rozellae, and C.
scansorius), which spend their time at a
considerable height above ground.
However, individuals of these same spe-
cies have been observed basking on tree
trunks only 50-250 cm above the ground
(LEE 1996, MCCRANIE & WILSON 1996).
Other species (e.g., C. bivittatus) have been
found predominantly under fallen logs
(WILSON et al. 1986). Celestus rozellae is a
livebearer, producing litters of 3-5 juveni-
les during the months from May to July
(ALVAREZ DEL TORO 1983). A female
Celestus bivittatus from Honduras gave
birth to ten juveniles (SVL 28.2-30. 7 mm)
with bright red-orange tails on the 1st of
March 1980 (WILSON et al. 1986).

Fig. 106 (right top). Celestus rozellae (Quintana

Roo, Mexico). Photo: H. Bahena B.
Fig. 107 (right middle). Celestus hylaius (Coma-
dre, Costa Rica). Photo: R. W. Van Devender
Fig. 108 (right bottom). Celestus scansorius (La
Fortuna, Yoro, Honduras). Photo: J. R. McCranie
65
 Anguidae
Fig. 109. Celestus biuittatus (Honduras).
Photo: J . R. McCranie
Celestus montanus SCHMIDT 1933b, Field
Mus. Nat. Hist. Puhl. Zool. Ser. 20: 21; type
locality: Sierra de Merendon, west of San
Pedro Sula, Honduras. SVL to 93 mm.
....
~
N
Northwestern Honduras and adjacent
Guatemala, 915-1370 m elevation.
a.
fll
Celestus orobius SAVAGE & LIPS 1994, Rev.
Biol. Trop. 41: 823; type locality: area near
Hortensia , Palma and Fortuna on the
Carretera Interamericana, 1500-2000 m ele-
vation, Canton Perez-Zeledon, Cordillera de
Talamanca, San Jose, Costa Rica. SVL to 83
mm. Cordillera de Talamanca, Costa Rica,
1500-2000 m elevation.
Celestus rozellae SMITH 1942b, Proc. U.S.
Natl. Mus. 92: 372; type locality: Palenque,
Chiapas, Mexico. SVL to 102 mm. Southern
Mexico through northern Guatemala to
Belize and the Sierra de Espirito Santo near
the Guatemalan-Honduran border, sea level
to 1350 m.
Celestus scansorius MCCRANIE & WILSON
1996, Rev. Biol. Trop. 44: 260; type locality:
2.5 airline km NNE La Fortuna, 15°25'N,
87°19'W, 1550 m elevation, Cordillera
Nombre de Dios, Yoro, Honduras. SVL to 111
mm. Northwestern Honduras, 1550-1590 m.
Ce/estus
• C. atitlanensis
• C. bivittatus
Y C. montanus
C. raze/Jae
..a.. C. scansorius
66
Key to Celestus
1 a Suboculars and postoculars in a single
continuous series Celestus eyanochloris
b Subocular and postocular series juxta-
posed ................ ....... .. ... ......................... .. 2
2 a 14-18 lamellae under 4th toe .................. 3
b 20-27 lamellae under 4th toe .................. 4
3 a 17-18 lamellae under 4th toe; SVL to 110
mm .......... ... ........ Celestus atitlanensis
b 14-16, rarely 17, lamellae under 4th toe;
SVL to 90 mm .......... Celestus biuittatus
4 a Three almost subequal prefrontal plates,
median prefrontal separated from supra-
oculars; flank pattern consists of occelated
blotches .............. ..... Celestus montanus
b One single large prefrontal plate, in
contact with supraoculars; flank pattern
not as above ................... ... .. .... ............... 5
5 a Fewer than 70 transverse rows of dorsal
scales ..................... ...... Celestus orobius
b 73-81 transverse rows of dorsal scales .. 6
6 a No pale dorsolateral stripes; pale vertical
bars on neck present ... Celestus rozellae
b Pale dorsolateral stripes present; pattern
on neck not as above ............... .......... ...... 7
7 a 84-92 transverse rows of ventral scales;
76-81 transverse rows of dorsal scales
............ .. ...... ..... .... ..... Celestus hylaius
b 76-78 transverse rows of ventral scales; 74
transverse rows of ventral scales ............
.... .. ............. ... ....... Celestus scansorius
Further Reading
VILLA & WILSON 1988, CAMPBELL & CAMARILLO
1994, SAVAGE & LIPS 1994, MCCRANIE & WILSON
1996, TOWNSEND et al. 2005
Celestus
0 C. cyanoch/oris
• C. hylaius
..a.. C. orobius

Coloptychon
The genus Coloptychon consists of a single
species, C. rhombifer (PETERS 1877). The
holotype of C. rhombifer was collected in
the province of Chiriqui in Panama. Since
the species' original description more than
120 years ago, only two additional speci-
mens of C. rhombifer have been discover-
ed, both of them in the rainforests of
southeastern Costa Rica (GOOD 1988). The
natural history of this species is virtually
unknown. Coloptychon rhombifer has an
elongate slender body and a conspicuously
long tail (reaching two and half times the
SVL). A photograph of a preserved speci-
men of this species was published by GOOD
(1988). The dorsal body pattern consists of
eight pale subtriangular crossbars on a
dark background. The tail is also banded,
whereas the legs show only an indistinct
mottling.
Coloptychon rhombifer (PETERS 1877),
Monats. Akad. Wiss. Berlin 1877: 298; type
locality: Chiriqui, Panama. SVL to 120 mm.
Western Panama and southern Costa Rica,
50-500 m elevation.
Further Reading
PETERS 1877, GOOD 1988, SAVAGE 2002
.& Co/optychon rhombifer
Anguidae
Fig. 110. Coloptychon rhombifer. Drawing after
a photo in GOOD (1988). Above: Lateral view of
head (from DUMERIL et al. 1870-1909).
Diploglossus
The species of the genus Diploglossus are
stout-bodied lizards, the species D.
monotropis being the most colorful and
also the largest anguid lizard of Central
America, reaching a maximum SVL of215
mm. These lizards are mostly ground
dwellers, but can also be observed clirltb-
ing on trees with abundant vine growth. A
female of D. bilobatus was discovered
beneath the trunk of a tree where it
guarded a clutch of six eggs (TAYLOR 1956).
Diploglossus bilobatus (O'SHAUGHNESSY
1874), Ann. Mag. Nat. Hist. (4) 14: 257; type
locality: Costa Rica. SVL to 96 mm. Costa
Rica and western Panama, sea level to 1360
m elevation in rain forest.
Diploglossus monotropis (KUHL 1820),
Beitrage zur Zoologie und vergleichenden
Anatomie: 128; type locality: unknown. SVL
to 215 mm. Northern Nicaragua to western
Ecuador, sea level to 1000 m elevation in rain
forest.
Diploglossus montisilvestris MYERS 1973,
Amer. Mus. Novit. 2523: 3; type locality: SE
slope of Cerro Pirre, 1440 m elevation,
Serrania de Pirre, Darien, Panama. SVL to
100 mm. Eastern Panama, near border with
Colombia, 1440 m elevation in cloud forest .
67
 Anguidae

Key to Diploglossus
1 a A pair of prefrontals and one median
frontonasal (Fig. llla); dorsal scales dis-
tinctly keeled; SVL to 215 mm ............ ...... .
..... ................... Diploglossus monotropis
b A single large prefrontal (Fig. lllb); dor-
sal scales not or only faintly keeled; SVL
to93mm ................................................... 2
2 a First supralabial separates nasal from
rostral; one postnasal ............................. .
...... ........... Diploglossus montisilvestris
b Nasal in contact with rostral; two postna-
sals .................... Dip logloss us bilobatus

Fig. 113. Diploglossus bilobatus (Cabaceras,

Costa Rica). Photo: R. W. Van Devender

a. D. monotropis b. 0 . bilobatus
Fig. 111. Scalation of dorsal surface of head
(frontonasal brown; prefrontals orange).

Fig. 114. Juvenile Diploglossus bilobatus (Isla

Popa, Panama). Photo: G. Kohler

Further Reading
TAYLOR 1956, MYERS 1973
Fig. 112. Diploglossus monotropis (Rio San
Juan, Nicaragua). Photo: G. Kohler

• Dip/og/ossus bilobatus • Diplog/ossus monotropis

• Diploglossus montisilvestris

68

Abb. 115. Gerrhonotus liocephalus (Chiapas, Mexico).
Gerrhonotus
Four species of Gerrhonotus are currently
recognized, only one of which, G. lioceph-
alus, occurs in Central America (Goon
1994). The other species are distributed in
Mexico and the southern USA.
Gerrhonotus liocephalus is a very elongate
lizard with a very long and breakable tail.
Because of its extremely slender appear-
ance, this lizard is called "lagarto culebra"
(="snake-lizard") in Chiapas (ALVAREZ DEL
TORO 1960). On an olive-brown ground
• Gerrhonotus /iocephalus
Anguidae
color, this species exhibits irregular dark
brown cross bars on the dorsal surfaces of
the body and tail. The venter is dirty white
to pale yellow. The short legs are used only
for slow movement, whereas for more
rapid movement, when, for example, it is
trying to escape, it will dart away with
undulating body movements, with the legs
adpressed to the body. Gerrhonotus lioce-
phalus can be found both on the ground
and on low vegetation (bushes and small
trees). When climbing, it utilizes its pre-
hensile tail as a fifth limb to hold onto the
vegetation. It feeds mostly on insects,
which are slowly approached and then
grasped with its strong jaws in a fast for-
ward motion of the body. The females pro-
duce clutches with 10 to 15 eggs that are
about 13x8 mm in size (ALVAREZ DEL TORO
1960).
Gerrhonotus liocephalus WIEGMANN 1828, Isis
von Oken 21: 381; type locality: Mexico. SVL
to 190 mm. Widely across Mexico to eastern
Chiapas; 200-2000 m elevation.
Further Reading
.ALVAREZ DEL TORO 1983, Goon 1994
69
 Anguidae

Mesaspis

The genus Mesaspis is composed of six

species, two of which occur in Central
America: Mesaspis monticola inhabits the
mountain regions of Costa Rica and
western Panama, M. moreletii is found in
the highlands of northern Central
America (GOOD 1988). Both species are
ground dwellers that prefer open pine Fig. 116. Mesaspis monticola (Cerro de la
forest, paramo and clearings in the cloud Muerte, Costa Rica). Photo: M. Franzen
forest at elevations up to 3250 m (e.g., on
the Cerro de la Muerte, Costa Rica). Like
most Central American anguids, these
agile lizards prey mostly on insects and
other arthropods. Both species are vivi-
parous. Once a year, after a gestation peri-
...
t"I od of about twelve months, the female M .
monticola will give birth to a litter of 2 to
a
!!ii

10 young. The newborn M. monticola

fll measure 50-57 mm TL with a SVL of23-26
mm (FITCH 1970, 1973a, VIAL & STEWARD
1985, WICKNICK 1993). Fig. 117. Mesaspis moreletii (El Pital, Honduras).
Photo: G. Kohler

Mesaspis monticola (COPE 1877), Proc. Amer.

Phil. Soc. 17: 97; type locality: Pico Blanco,
Costa Rica. SVL to 85 mm; Costa Rica to
Chiriqui Province, Panama, 1800-3800 m el-
evation.
Mesaspis moreletii (BOCOURT 1871), Bull.
Nouv. Arch. Mus. Hist. Nat. Paris 7: 102; type
locality: Peten and pine forests of Alta Vera-
paz, Guatemala. SVL to 95 mm; southern
Mexico to northern Nicaragua, 1500-3000+ m
elevation.
Fig. 118. Mesaspis moreletii (male, Cerro
Kilambe, Nicaragua). Photo: J. Sunyer
Key to Mesaspis
1 a 14-16 longitudinal rows of dorsal scales;
prefrontal scale usually present ................ .
................ ..... ... ....... .. Mesaspis monticola
b 18-20 longitudinal rows of dorsal scales;
prefrontal scale usually absent ................. .
........ ........... .. .. ..... ....... Mesaspis moreletii

Further Reading
TAYLOR 1956, FITCH 1970, 1973a, VIAL &
STEWARD 1985, GOOD 1988, 1989, WICKNICK
1993, VESELY & KOHLER 2001, SUNYER &
KOHLER 2007
Mesaspis moreletii
• Mesaspis montico/a

70
 Eublepharidae

Eublepharidae
Currently, the geckos with moveable
eyelids are separated as a distinct family
(Eublepharidae) from the other geckos
(family Gekkonidae) (KLUGE 1987, GRIS-
MER 1988). The family Eublepharidae is
widely distributed in the Old World but in
Central America represented only by the
genus Coleonyx. Banded geckos are char-
acterized by moveable eyelids and a strik-
ing banded dorsal pattern. Their subdigi-
tal lamellae are neither broadened nor
adhesive.

Fig. 120. Coleonyx mitratus (San Salvador, El

Coleonyx Salvador). Photo: G. Kohler
Two species of the genus Coleonyx occur in
Central America. Coleonyx elegans and C.
mitratus are closely related. Both are ter-
restrial nocturnal lizards that hide
beneath rocks and fallen tree trunks and
in hollow tree trunks during the daytime.
Their diets consist mostly of insects and
spiders. The movements of these attrac-
tively patterned geckos are very graceful,
almost cat-like. When alarmed, they
expand their throat and assume a stiff
posture with the body held high, well off
the ground. Both species are oviparous
and produce clutches consisting of one or
two eggs each. At a temperature of 28-
300C, the eggs hatch after 62-68 days (C.
elegans) and 46-65 days (C. mitratus), Fig. 121. Coleonyx elegans (near Chetumal,
respectively (KALBERLAH 1996, KOHLER Quintana Roo, Mexico). Photo: H. Bahena B.
1997).

Coleonyx elegans GRAY 1845, Ann. Mag. Nat.

Hist. (1) 16: 163; type locality: Belize. SVL to
97 mm. On the Pacific versant from southern
Nayarit, Mexico, to western El Salvador, and
on the Caribbean versant from Veracruz,
Mexico, through the Yucatan Peninsula to
northern Guatemala and Belize, sea level to
1000 m elevation.
Coleonyx mitratus (PETERS 1863), Monatsber.
Kong. Akad. Wiss. Berlin 1863: 42; type lo-
cality: Panama. SVL to 91 mm. Guatemala to
Costa Rica on the Pacific versant and
northern Honduras on the Caribbean ver-
Fig. 119. Coleonyx mitratus (Isla de Utila, sant, sea level to 700 m elevation.
Honduras). Photo: G. Kohler
71
 Eublepharidae

Gekkonidae
Geckos of the family Gekkonidae occur
throughout much of the tropical and sub-
tropical regions of both the Old and New
Worlds. In Central America, this family is
represented by nine genera with 24 spe-
cies. Most gecko species are characterized
by broadened adhesive lamellae beneath
fingers and toes (allowing these animals to
Fig. 122. Coleonyx elegans (Mexico). climb on smooth, vertical surfaces), as well
Photo: R. D. Bartlett as by the absence of moveable eyelids
(their eyes are covered by a transparent
spectacle instead). Some gecko species are
crepuscular or nocturnal (e.g., Hemi-
dactylus and Phyllodactylus), whereas
Key to Coleonyx
others such as species of the genera
1 a Claws almost completely covered by a Gonatodes and Sphaerodactylus are active
scaly sheath (Fig. 123a); first sublabial tri-
angular; an elongate field of small scales during the day. All Central American
behind internasals .... .. Coleonyx elegans gecko species are oviparous, usually pro-
b Claws not covered by a scaly sheath (Fig. ducing one or two eggs per clutch.
123b); first sublabial squarish; no field of
small scales behind internasals .... .. ..........
......... ....................... ... Coleonyx mitratus Key to genera of Gekkonidae
la Digits with widened subdigital lamellae, at
least partially (Figs. 124a-f) ............... ...... 2
b Digits without widened subdigital lamellae,
lateral slender throughout their length (Figs.
view 124h-j), or with a single extremely enlarged
terminal subdigital scale (Fig. 124g) ......... 7
ventral 2a Dilation of subdigital lamellae restricted to
view
two symmetrically enlarged terminal plates
(Fig. 124£) ............. ............. Phyllodactylus
a. Coleonyx elegans b. Coleonyx mitratus
b Dilation of subdigital lamellae extends
Fig. 123. Toes (scaly sheath shaded). Drawings throughout most of digit (Figs. 124a-e) ..... 3
by M. Vesely. 3a Claw in contact with or only slightly beyond
widened subdigital lamellae (Fig. 124a)
........... .................... .. ..... ...... Thecadactylus
Further Reading b Claw much beyond distal part of widened
Kl.AUBER 1945. subdigital lamellae (Figs. 124b-e) ..... ........ 4
4a All subdigital lamellae undivided (Fig.
124c) ...................... .................. Aristelliger
b At least the distal subdigital lamellae divi-
ded or arranged in pairs (Figs. 124b,d,e) .. 5
5a Median subcaudals not enlarged, not
aligned in a median series ...... ....... .
...... ...... ................... ..... ....... Lepidodactylus
b Median subcaudals enlarged and
aligned in a median series ........ ................ 6
• Coleonyx elegans 6a Dorsal scales heterogeneous with large
tubercles .............................. Hemidactylus
• Coleonyx mitratus
A Aristelliger georgeensis b Dorsal scales homogeneous, no large
tubercles ...... .. ..... .... ..................... .. Gehyra
72
 Gekkonidae

7a A single extremely enlarged spherical ter-

minal subdigital scale present (Fig. 124g)
..... ........ ... .. .. ................. Sphaerodactylus
b Terminal subdigital scale not larger than
other subdigital lamellae .......................... 8
Sa Two supralabials to a point below anterior
margin of eye (Fig. 125a); head of adult
males colored more or less similar to body
........ ............... ................ .. Lepidoblepharis
b Three supralabials to a point below anteri-
or margin of eye (Fig. 125b); head of adult
males orange colored, contrasting with
grayish body ................... ........ ... Gonatodes

Fig. 126. Aristelliger georgeensis (Isla Cozumel,

Mexico). Photo: J. C. Lee

a b. ~ d. Aristelliger
Thecadactylus Lepidodactylus Aristel/iger Hemidactylus
The genus Aristelliger, with six species,
has a distribution centered mostly in the
Antilles. Only a single species occurs in
Central America. Aristelliger georgeensis
is mostly found in the vicinity of human
settlements. At night, these large geckos
can be observed on house walls and the
trunks of palm trees, where they hunt for
f. g.
insects. They prey upon anything they can
Gehyra Phyllodactylus Sphaerodactylus
subdue, even feeding on smaller lizards
(DUNN & SAXE 1950). Like many geckos, A.
georgeensis is also capable of producing a
vocalization that is described as a screech
or a low chirp (DUELLMAN 1965, SCHWARZ
Fig. 124. & HENDERSON 1991). The species is ovipa-
Under- rous (LEE 1996).
sides of
h. i. j. toes in
Lepidoblepharis Gonatodes geckos. Aristelliger georgeensis (BocOURT 1873),
Coleonyx
Miss. Sci. Mex., Rept. 1873: 41; type locality:
St. George Island, near Belize City, Belize.
SVL to 115 mm. Eastern coast of Yucatan
Peninsula including offshore islands, also on
Islas Providencia and San Andres, sea level
to 50 m elevation.

a. Lepidoblepharis b. Gonatodes Further Reading

K. SCHMIDT 1941, BAUER & RUSSELL 1993a, b,
Fig. 125. Lateral view of head (labials orange). LEE 1996
73
 Gek.konidae

Gehyra

The species of the genus Gehyra are wide-

ly distributed in Madagascar, Southeast
Asia, Australia, and Oceania. Like many
gecko species, Gehyra mutilata has been
introduced to numerous places in the tropics
worldwide, including Central America,
where the species is known from Chiapas,
Mexico (ALVAREZ DEL TORO 1983). These
crepuscular and nocturnal lizards prey
mostly on insects and spiders, but also eat
sweet fruit. Gehyra mutilata is oviparous
and produces clutches with 1 or 2 eggs.
Gehyra mutilata (WIEGMANN 1835), Acta
Acad. Caes. Leop. Carol. Nat. Cur. 17: 238;
"Manila" [Luzon, Philippines] . SVL to 64 mm.
Southeast Asia, Oceania and India; dispersed
in the Tropics worldwide.
fll
Further Reading
ALVAREZ DEL TORO 1983, JOHNSON 1990
Fig. 127. Juvenile ofGehyra mutilata
(Tapachula, Mexico). Photo: T. Bille
.a. Gehyra mutilata
Gonatodes albogularis
Fig. 128. Adult male of Gonatodes albogularis
(Bartola, Rio San Juan, Nicaragua).
Photo: G. Kohler
Gonatodes
The species of the genus Gonatodes are
diurnal, attractive colored geckos that
have their main distribution in tropical
South America. Of the more or less 20 spe-
cies, only one (G. albogularis) occurs in
Central America. Claims that G. annularis
reaches southern Central America (VILLA
et al. 1988, AUTH 1994) are erroneous and
are probably based on PETERS & DoNoso-
BARROS (1970). Gonatodes annularis is
restricted to the northern Amazon Basin
(AVILA-PIRES 1995, HOOGMOED pers. comm.
1999). Unlike most other gecko species,
the representatives of the genus
Gonatodes have round pupils and do not
have broadened adhesive lamellae
beneath fingers and toes. In Gonatodes,
there is a strong sexual dichromatism,
with the male exhibiting a strikingly
colorful pattern, whereas the more or less
grayish brown females are less con-
spicuous. Gonatodes albogularis can be
found in many different habitats, including
human settlements. These graceful geckos
are highly thermophilic and, particularly
in the morning hours, can be observed
basking in the sun on tree trunks, walls,
and rocks. The females lay single eggs

74
 Gekkonidae

Fig. 129. Hemidactylus garnotii (Rincon de Osa,

Costa Rica). Photo: G. Kohler

several times per year. The juveniles hatch

after an incubation period of two to three
months (KOHLER 1997).

Gonatodes albogularis (DUMERIL & BIBRON

1836), Erp. Gen. 3: 415; type locality:
Martinique. SVL to 40 mm. Central America
and northern South America, sea level to
1000 m elevation. The subspecies G. a. fuscus
(HALLOWELL 1855) occurs in Central America.

Further Reading
TAYLOR 1956; FITCH 1973a, b

Hemidactylus

Five species of Old World house geckos

(genus Hemidactylus) have been intro-
duced to numerous localities in Central
America (KOHLER 200ld, FRANKLIN 2000,
MCCRANIE & WILSON 2000, GREENBAUM
2002). These geckos are usually associated
with human habitation. At night, they can
be observed in houses, where they prey
upon arthropods. They are capable of
vocalization and their call sounds like
"gee-gee-gee." The clutches consist of two
eggs each, with the young hatching after
an incubation period of about two months.
Hemidactylus
• H. frenatus
..,. H. mabouia
0 H. turcicus
Fig. 130. Hemidactylus mabouia.
Photo: R. D. Bartlett
Hemidactylus brookii GRAY 1845, Cat.
Lizards Brit. Mus.: 153; type locality: Borneo
and Australia. SVL to 67 .5 mm. Introduced to
Panama and Honduras, sea level to 1000 m
elevation.
Hemidactylus frenatus DUMERIL & BIBRON
1836, Erp. Gen. 3: 366; type locality: Java and
Timor. SVL to 60 mm. Introduced to several
localities in Central America, sea level to
1230 m elevation.
Hemidactylus garnotii DUMERIL & BIBRON
1836, Erp. Gen. 3: 368; type locality: Tahiti,
French Polynesia. SVL to 65 mm. Introduced
to Costa Rica, sea level to 1160 m elevation.
Hemidactylus mabouia (MOREAU DE JONNES
1818), Bull. Soc. Philom. Paris 1818: 138; type
locality: Antilles. SVL to 75 mm. Introduced
to Panama and Honduras, sea level to 1000 m
elevation.
Hemidactylus turcicus (LINNAEUS 1758),
Systema Naturae, ed. 10: 202; type locality:
"Oriente". SVL to 59 mm. Introduced to
Panama and Mexico (Yucatan Peninsula) ,
sea level to 100 m elevation.

75
 Gekkonidae

Key to Hemidactylus
1 a Tail with denticulate fringe; two or three
pairs of enlaited chin shields, posterior
pair(s) not in contact with infralabials
(Fig. 133c) .......... Hemidactylus garnotii
b Tail without denticulate fringe, although
widely spaced ventrolateral spines can be
present; two pairs of enlarged chin shields,
both pairs in contact with infralabials
(Fig. 133a-b, d-e) ...................................... 2
2 a Enlarged subdigital lamellae of 4th toe do a. Hemidactylus mabouia b. Hemidactylus frenatus
not reach base of toe (Fig. 132a) ......... .
. . . . . . . . . ........ ....... Hemidactylus mabouia Fig. 132. Underside of 4th toe (lamellae shaded) .
b Enlarged subdigital lamellae of 4th toe
reach base of toe (Fig. 132b) ................. .. 3
3 a No tubercles on upper surface of hind
limbs and above ear opening ........ .......
... . . . . . . . . . ...... ....... Hemidactylus frenatus
b Distinct tubercles on upper surface of hind
limbs and above ear opening ............ ... .... 4
4 a Each tubercle on dorsum surrounded by
10-12 small scales; males with 21-32 pre-
anofemoral pores Hemidactylus brookii
b Each tubercle on dorsum surrounded by
15-20 small scales; males with 3-10 pre-
anofemoral pores .... ..................... ............. .
..... .............. ....... . Hemidactylus turcicus

Further Reading Fig. 133. Chin region

McCOY 1970, MARCELLINI 1971, SALVADOR 1981, in Hemidactylus.
POWELL & MAxEY 1990, AUTH 1994, LEE 1996,
POWELL et al. 1998 Anterior chin shields
orange, posterior ones
brown.

Fig. 131. Hemidactylus frenatus (David, Fig. 134. Hemidactylus brookii (Cordoba,
Chiriqui, Panama). Photo: M. Lundberg Colombia). Photo: M. Lundberg
76
 Gekkonidae

Fig. 135. Hemidactylus turcicus. Fig. 136. Lepidoblepharis xanthostigma

Photo: A. Nollert (Nusagandi, Panama). Photo: G. Kohler

Lepidoblepharis

The species of the genus Lepidoblepharis

are small ground dwellers that live in the
leaflitter on the rain forest floor, mostly in
close vicinity to rotten tree trunks, large
roots or decaying plant material that pro-
vide sufficient humidity. The females pro-
duce clutches consisting of single eggs,
which they lay in organic debris under
roots or loose bark several times per year.
Both Central American species of Fig. 137. Lepidoblepharis sanctaemartae
Lepidoblepharis appear to have an exten- (Panama). Photo: M. A. Guerra
sive reproductive season.

Further Reading
Lepidoblepharis sanctaemartae (RUTHVEN BARBOUR 1923, DUNN 1940a, TAYLOR 1956, RIVAS
1916), Occ. Pap. Mus. Zool., Univ. Michigan FUENMAYOR et al. 2002
21: 2; type locality: Fundaci6n, Colombia.
SVL to 35 mm. Central Panama to northern
Colombia and northwestern Venezuela, sea
level to 1200 m elevation.
Lepidoblepharis xanthostigma (NOBLE 1916),
Proc. Biol. Soc. Wash. 29: 87; type locality:
Zent, near Puerto Limon, Costa Rica. SVL to ~ Lepidoblepharis sanctaemartae
38 mm. Southern Nicaragua to Colombia, sea
level to 1360 m elevation. • Lepidoblepharis xanthostigma
.a. Lepidodactylus lugubris

Key to Lepidoblepharis
1 a Dorsal scales distinctly imbricate .....
. . . ....... Lepidoblepharis sanctaemartae
b Dorsal scales juxtaposed ..... ... ... .... .... .
.. .. . . . . . . . Lepidoblepharis xanthostigma
77
 Gekkonidae
Fig. 138. Lepidodactylus lugubris (EI Valle de
Anton, Panama). Photo: G. Kohler
...
t-4
a
N 1983). During the course of the year, sever-
Lepidodactylus
Cll
Lepidodactylus lugubris is an animal that
has spread with the advance of civilization
and been introduced worldwide. The first
published record of this species being
established in Central America appeared
in the early 1960s (H. SMITH & GRANT
1961). These authors collected a subadult
female in Panama. Meanwhile, L. lugubris
is known from additional localities in
Central America, including places in Costa
Rica, as well as the Corn Islands off the
Caribbean coast of Nicaragua (VILLA et al.
1988, KOHLER 1999c). The species repro-
duces parthenogenetically, with the fe-
males producing clutches consisting of one
or two eggs that hatch after 60 to 100
days, depending on incubation tempera-
ture (KOHLER 1997).
Lepidodactylus lugubris (DUMERIL & BIBRON
1836), ERP. G:EN. 3: 304; type locality: "Otaiti"
[= Tahiti, according to PETERS & DONOSO-
BARROS 1970) SVL to 42 mm. Southeast Asian
and lndoaustralian region; dispersed world-
wide, sea level to 700 m elevation.
Further Reading
H. SMITH & GRANT 1961, HENDERSON et al. 1976,
ROLL 2002
78
Phyllodactylus
Leaf-toed geckos are distinguished from
all other Central American geckonid gen-
era by a pair of enlarged leaf-like terminal
lamellae on the fingers and toes, whereas
the more proximal subdigital lamellae are
not widened. Leaf-toed geckos are crepus-
cular and nocturnal inhabitants of old
trees with loose bark and can also be
found on house walls. During the daytime
in its natural environment, these geckos
like to hide in tree crevices and beneath
loose bark, while in human habitations
they will make use of any dark refuge that
provides some kind of protection. The call
of this species can be described as a series
of kiss-like sounds (ALVAREZ DEL TORO
al clutches consisting of two eggs each are
glued to the surface in crevices or other
protected sites. In cases where several
females use the same egg laying site, mass
accumulations of eggs and eggshells can
be found (KOHLER unpubl. observ. on Utila,
Honduras).
Phyllodactylus insularis DIXON 1960,
Herpetologica 16: 9; type locality: Half Moon
Cay, Belize. SVL to 58 mm. Endemic on Half
Moon Key, Belize, sea level to 50 m elevation.
Phyllodactylus palmeus DIXON 1968, Proc.
Biol. Soc. Washington 81: 419; type locality:
0.5 km N Roatan, Isla de Roatan, Islas de la
Bahia, Honduras. SVL to 76 mm. Endemic to
the Islas de la Bahia and Cayos Cochinos,
Honduras, sea level to 200 m elevation.
Phyllodactylus tuberculosus WIEGMANN 1835,
Nova Acta Acad. Leop.-Carol. 17 (1): 241; type
locality: "California". SVL to 72 mm. Pacific
versant of southern Mexico to Costa Rica, sea
level to 1160 m elevation. The subspecies
Phyllodactylus t. ingeri DIXON 1960 (Belize),
Phyllodactylus t. magnus TAYLOR 1942
(northwestern Guatemala northward), and
Phyllodactylus t. tuberculosus (central
Guatemala southward) occur in Central
America.
 Gekkonidae

Fig. 139. Phyllodactylus insularis (Half Moon Fig. 143. Phyllodactylus tuberculosus (Isla
Cay, Belize). Photo: J. C. Lee Ometepe, Nicaragua). Photo: G. Kohler

Fig. 140. Key to Phyllodactylus

Dorsal scales in 1 a Dorsum with 14 or fewer longitudinal rows
Phyllodactylus of enlarged tubercle-like scales ...... .... .
tuberculosus .. ...... .. ...... Phyllodactylus tuberculosus
(tubercles shaded). b Dorsum with 16 or more longitudinal rows
of enlarged tubercle-like scales ................ 2
2 a 48-60 ventral scales from throat to vent;
39-47 tubercle-like scales in a paraverte-
bral row from occiput to base of tail .. .....
.. .. .. . .. ... ............. Phyllodactylus palmeus
b 60-66 ventral scales from throat to vent;
46-52 tubercle-like scales in a paraverte-
bral row from occiput to base of tail ........... .
...... .. .. .. .. ..... .... Phyllodactylus insularis

Further Reading
DIXON 1960, 1964

Fig. 141. Eggs and egg shells of Phyllodactylus

Phy/lodactylus
palmeus (Isla de Utila, Honduras).

• P. insularis
""" P. palmeus
P. tubercu/osus

Fig. 142. Phyllodactylus palmeus (Isla de

Guanaja, Honduras). Photo: G. Kohler
79
 Gekkonidae

Sphaerodactylus

With some 150 recognized species, the

least geckos (Sphaerodactylus), with their
center of diversity and distribution in the
Caribbean (about 70 species), form the
largest of all geckonid genera. Nine spe-
cies of least geckos occur in Central
America, where they are restricted to low-
land regions (sea level to about 1000 m
elevation). These are very small lizards (in
fact the smallest amniote is a Sphaero-
dactylus; HEDGES & THOMAS 2001). Most
Central American Sphaerodactylus have a
maximum SVL of 25 to 30 mm, except for
the island endemics S. rosaurae and S. Fig. 144. Sphaerodactylus rosaurae (Isla de
pacificus, which reach 40 mm. Least Roatan, Honduras). Photo: G. Kohler
geckos are typical leaf litter inhabitants;
they dart through the leaves, skillfully
hunting small arthropods, such as spring-
tails, cricket larvae and little spiders. The
best way to spot these small geckos is
when shuffiing slowly through the leaf
litter carefully watching the ground. These
geckos especially like to be in the proximity
of the roots of large trees. The population
density can be very high at certain sites,
reaching two or three individuals per
square meter. On the Islas de la Bahia,
Honduras, I found S . millepunctatus and
S. rosaurae syntopically, whereas the
former species occurs together with S.
dunni at Pico Bonito on the Caribbean
coast of Honduras. Least geckos produce
clutches consisting of single eggs that Fig. 145. Sphaerodactylus millepunctatus
hatch after 60 to 90 days . (Matamoros, Honduras). Photo: G. Kohler

.6. Sphaerodactylus argus

• Sphaerodactylus lineolatus
• Sphaerodactylus graptolaemus
• Sphaerodactylus homolepis

Fig. 146. Sphaerodactylus dunni (Pico Bonito,

Atlantida, Honduras). Photo: G. Kohler
80

Fig. 147. Sphaerodactylus lineolatus
(Nusagandi, Panama). Photo: G. Kohler
Fig. 148. Sphaerodactylus homolepis
(Tortuguero, Costa Rica). Photo: G. Kohler
Fig. 149. Sphaerodactylus glaucus (Bacalar La-
goon, Quintana Roo, Mexico). Photo: H. Bahena B.
Gekkonidae
Sphaerodactylus argus GOSSE 1850, Ann.
Mag. Nat. Hist. (2) 6 (35): 347; type locality:
Jamaica. SVL to 33 mm. Jamaica, Cuba,
Bahamas, Key West (Florida); scattered
populations along and off the Caribbean coast
of Central America, including Isla San
Andres, Isla de Maiz Grande, Isla Cozumel,
Provincias Bocas del Toro and San Blas
(Panama), sea level to 100 m elevation.
Sphaerodactylus dunni SCHMIDT 1936a, Proc.
Biol. Soc. Washington 49: 46; type locality: Rio
Naco, near Cofradia, Honduras. SVL to 28
mm. Northern Honduras, sea level to 700 m
elevation.
Sphaerodactylus glaucus COPE 1865, Proc.
Acad. Nat. Sci. Philadelphia 17: 192; type
locality: near Merida, Yucatan, Mexiko. SVL
to 29 mm. Veracruz and Oaxaca, Mexico,
through the Yucatan Peninsula and northern
Guatemala to western Honduras, 200-1000 m
elevation.
Sphaerodactylus graptolaemus HARRIS &
KLUGE 1984, Occ. Pap. Mus. Zool. Univ.
Michigan 706: 4; type locality: Isla del Cano,
Puntarenas, Costa Rica. SVL to 32 mm.
Southern Pacific versant of Costa Rica and
adjacent Panama, sea level to 700 m eleva-
tion.
Sphaerodactylus homolepis COPE 1886a,
Proc. Amer. Philos. Soc. 23: 77; type locality:
Nicaragua. SVL to 33 mm. Caribbean coast of
southern Nicaragua to central Panama, sea
level to 600 m elevation.
Sphaerodactylus lineolatus LICHTENSTEIN &
VON MARTENS 1856, Nomenclator Reptilium
et Amphibiorum Musei Zoologici
Berolinensis: 6; type locality: Veragua,
Panama. SVL to 32 mm. Western Panama to
northwestern Colombia, sea level to 800 m
elevation.
_ Sphaerodactylus
- (/
\l S. rosaurae
.A. S. argus
e S. dunni
§ S. g/aucus
• S. millepunctatus
81
 Gekkonidae

Sphaerodactylus millepunctatus HALLOWELL

1861, Proc. Acad. Nat. Sci. Philadelphia 12:
480; type locality: Nicaragua. SVL to 31 mm.
Isthmus of Tehuantepec, Mexico, to northern
Costa Rica, sea level to 750 m elevation; not
on the Yucatan Peninsula.
Sphaerodactylus pacificus STEJNEGER 1903,
Proc. Biol. Soc. Washington 16: 3; type lo-
cality: Isla de Coco, Costa Rica. SVL to 4 7
mm. Endemic on Isla de Coco, Costa Rica.
Sphaerodactylus rosaurae PARKER 1940, Ann. Fig. 150. Sphaerodactylus argus (Mandeville,
Mag. Nat. Hist. (17) 5: 264; type locality: Manchester, Jamaica). Photo: J . C. Lee
Helene Island, Islas de la Bahia, Honduras;
SVL to 39 mm. Endemic to Islas de la Bahia
and Cayos Cochinos, Honduras, sea level to
100 m elevation.

Key to Sphaerodactylus
1 a Dorsum with a median band of small
granular scales between large imbricate
dorsal scales (Fig. 154a) ..... ............. ... .. ... . Fig. 151. Sphaerodactylus graptolaemus (P.N. La
.. .... ......... .... Sphaerodactylus rosaurae
Gangreja, Costa Rica). Photo: D. M. Dehling
b All dorsal scales about the same size,
without a median band of small granular
scales (Fig. 154b) ............................ ..... .. 2
2 a Dorsum of adults with distinct large dark
blotches, spots, or mottling; SVL to 4 7 mm;
endemic on Isla de Coco, Costa Rica ......... .
... ........ ......... Sphaerodactylus pacificus
b Dorsal pattern not as described above;
SVL to 33 mm ..... ............... ............... ...... 3
3 a Enlarged spine-like superciliary situated a. b.
posterior to level of mideye (Fig. 152a);
snout conspicuously convex in lateral view Fig. 152. Scalation of the snout region (supra-
..... ...... ...... ....... Sphaerodactylus dunni nasals shaded); note also the different position
b Enlarged spine-like superciliary situated of the spine-like superciliary (arrow).
anterior to level of mideye (Fig. 152b);
snout flat, in lateral view running in a
straight line from eye to tip of snout ...... 4
4 a One supranasal scale (Fig. 152a) .. .. ....... 5
Fig. 153. Scalation of ven-
b Two supranasal scales (Fig. 152b) ....... .... 7 tral tail (enlarged subcau-
5 a Median subcaudals enlarged and aligned dals shaded).
in a median series (Fig. 153); dorsum with-
out dark cross-bands ........................... ...... .
........ ... . Sphaerodactylus graptolaemus
b Median subcaudals not enlarged, not •-

• ----
aligned in a median series; juveniles with
dark cross-bands on dorsum .... ....... ..... .... 6
6 a Dorsal and parietal scales strongly keeled;
subcaudals arranged alternately; 48-67 a. S . rosaurae
dorsals between levels of axilla and groin
. . . . . . . . . . ...... Sphaerodactylus homolepis Abb. 154. Dorsal scales in Sphaerodactylus.

82
 Gekkonidae

b Dorsal scales weekly keeled, parietal

scales usually smooth; subcaudals
arranged in sequences of one small scale,
one large scale, and a pair of small scales;
62-81 dorsals between levels of axilla and
groin ...... ... Sphaerodactylus lineolatus
7 a Dorsal scales smooth .............................. ..
..... .......... ... ..... Sphaerodactylus glaucus
b Dorsal scales keeled .... ... ...... ...... ... ....... .. 8
8 a 38-50 dorsals between levels of axilla and
groin; dorsum with longitudinal rows of
ocellated blotches ............... .......... ...... ... ...
. .. ....... .. ......... ...... Sphaerodactylus argus
b 4 7-74 dorsals between levels of axilla and
groin; dorsal pattern variable, but without
longitudinal rows of ocellated blotches Fig. 156. Thecadactylus rapicauda (Bartola,
........ Sphaerodactylus millepunctatus Rio San Juan, Nicaragua). Photo: G. Kohler

Further Reading Cll

HARRIS & KLUGE 1984 (Sphaerodactylus of
Central America), KOHLER 2001c
Thecadactylus 1....
N
~
Thecadactylus rapicauda is a nocturnal,
arboreal inhabitant of the tropical rain
forest. At night, these large geckos can be
observed on the trunks of large trees some
3 to 6 m above the ground, while during
the daytime they retreat into bark crevices
and hollow branches. Other typical day-
time hiding places include piles of leaf
litter among the buttress roots of large
trees. Turnip-tailed geckos are also com-
monly encountered on house walls. They
feed on invertebrates (mostly crickets,
moths, grasshoppers, and spiders) and
Fig. 155. Thecadactylus rapicauda (Quintana small lizards. Like most geckos, T. rapi-
Roo, Mexico). Photo: H. Bahena B. cauda is capable of vocalization and its
call sounds like "chack-chack-chack"
(BEEBE 1944). During the reproductive
season, female T. rapicauda lay several
clutches each consisting of a single egg.

Thecadactylus rapicauda (HOUTI'UYN 1782),

Verh. Zeeuw. Genootsch. Wet. Vlissingen 9:
323; type locality: West Indies. SVL to 121
mm. Mexico through Central America to
tropical South America; also on the Lesser
Antilles, sea level to 1050 m elevation.

Further Reading
TAYLOR 1956, AVILA-PIRES 1995, RUSSEL &
BAUER 2002, BERGMANN & RUSSELL 2007
83
 Gymnophthalmidae
Gymnophthalmidae
PRESCH (1983) suggested separating the
so-called "microteiids" into a separate
family (Gymnophthalmidae) from the so-
called "macroteiids" (Teiidae). This view
has been accepted by the majority of her-
petologists working in Central America
(IBANEZ et al. 1996, SAVAGE 2002, WILSON
pers. comm., McCRANIE pers. comm.),
whereas others preferred not to recognize
the Gymnophthalmidae as a distinct family
(HARRIS 1985, MYERS & DONNELLY 1996).
The lizards of the family Gymnoph-
thalmidae, including about 30 genera with
approximately 150 species, are distributed
in temperate, subtropical, and tropical
areas of southern Mexico, Central
America, and South America. Limb reduc-
tion has occurred several times within
Gymnophthalmidae, and in many of these
it is accompanied by body elongation. All
gymnophthalmids are oviparous, usually
producing two eggs per clutch.
Key to genera of
Gymnophthalmidae
1 a Without frontoparietals, frontal, therefore,
in contact with parietals (Fig. 160a); no
outer ear opening present; extremities
greatly reduced ........ .. .... ......... .... Bachia
Fig. 157. Neusticurus apodemus (Palmar, Costa
Rica). Photo: R. W. Van Devender
84
b Frontoparietals present, frontal, therefore,
separate from parietals (Fig. 160b); outer
ear opening present; extremities normal 2
2 a Without eyelids .......... Gymnophthalmus
b Eyelids present ... ................ ... ........... .... .... 3
3 a Posterior edges of the parietal scales and
the interparietal form a straight line (Fig.
161a); upper surface of tongue covered
with plicae ... ..... ............... Ptychoglossus
b Middle parietal scale clearly protrudes
past the posterior edges of the parietal
scales (Fig. 161b); Upper surface of tongue
covered with papillae ... .. ..... ..... ..... ... ... 4
4 a Dorsal scalation heterogeneous (large
tubercular scales sprinkled in with the
finer ground scalation) ....... .................... 5
b Dorsal scalation homogeneous .. .. .. .. ....... 6
5 a Prefrontals and frontal fragmented into
small irregular scales (Fig. 162b); a clearly
enlarged scale between first finger and
wrist ........... .. ...... ..... ... ..... Neusticurus
b Prefrontals and frontal form large shields
(Fig. 162a); no clearly enlarged scale be-
tween first finger and wrist ... .......... .... ... .
... ... . . . . . . . . . . . . . . ....... .. .. ........... Echinosaura
6 a Dorsal scales smooth .................. Anadia
b Dorsal scales keeled ................................. 7
7 a A double longitudinal row of widened
gular scales; a vertebral pale stripe usu-
ally present ... ..... .. ......... Prionodactylus
b No double longitudinal row of widened
gular scales; no vertebral pale stripe ......
......... .......................................... Leposoma
Fig. 158. Leposoma southi (Nusagandi,
Panama). Photo: G. Kohler
 Gymnophthalmidae

frontoparietals

rostral

interparietal

supraoculars
parietals superciliaries

Fig. 159. Scalation of dorsal surface of head in

Prionodactylus.

Fig. 160.
Dorsumof
head (frontal
brown;
parietals
orange).

a. Bachia b. Gymnophthalmus

Fig. 161.
Dorsumof
head (interpa-
rietal brown; Fig. 163. Anadia ocellata (Rara Avis, Costa
parietals Rica). Photo: T. Leenders
orange).

a. Ptychoglossus b. Leposoma

Anadia

In Central America, two species of the

Fig. 162. genus Anadia, A. ocellata and A. vittata,
Dorsumof are found (OFTEDAL 1974). Both are very
head (prefron- slender diurnal tree dwellers with long
tals brown; prehensile tails. The habitat of these
frontals
lizards is the rain forest where they prefer
orange).
trees with a plentiful supply of bromel-
iads. Large bromeliads also serve these
b. Echinosaura b. Neusticurus
animals as hiding places. Their diet con-
sists primarily of insects and spiders.
85
 Gymnophthalmidae
Fig. 164. Anadia ocellata (Provincia Limon,
Costa Rica). Photo: R. W. Van Devender
Anadia ocellata GRAY 1845, Cat. Lizards Brit.
Mus. : 58; type locality: Tropical America. SVL
to 75 mm. Costa Rica and western Panama;
500-1310 m elevation in premontane rain
forest.
Anadia vittata BOULENGER 1913, Proc. Zool.
Soc. London 1913: 1033; type locality: Pena
Lisa, 300 ft, Condoto, Choc6, Colombia. SVL
to 67 mm. Eastern Panama to southern
Colombia; 0-100 m elevation in lowland rain
forest.
KeY: to Anadia
1 a Pale dorsolateral stripes maximum of two
scales wide; obvious ocelli on body sides;
27-32 scales at midbody; males with a total
of 8-11 femoral pores ..... Anadia ocellata
b Pale dorsolateral stripes at least three
scales wide; ocelli, if present, not obvious;
30-37 scales at midbody; males with a total
of 11-14 femoral pores ..... Anadia vittata
Further Reading
0FTEDAL 1974
86
Bachia
The small lizards of the genus Bachia are
slender ground dwellers. Two species of the
predominantly South American genus
reach the southern portion of Central
America (DIXON 1973, MCDIARMID &
DEWEESE 1977). Bachia blairi was consid-
ered by DIXON (1973) to be synonymous
with B. pallidiceps, a decision for which
there was no evidence. MCDIARMID &
DEWEESE (1977) were able to show that
Bachia blairi was a separate species,
which differs from B. pallidiceps in
coloration, build, and several scale
characters. Representatives of this genus
are characterized by the absence of an
external ear opening and by the tendency
to reduced limb size. On the front limbs, B.
blairi and B. pallidiceps have four toes,
whereas the rear limbs have only three.
They are highly secretive ground dwellers
in the rain forest. These lizards are usual-
ly found in loose dirt or under trees or logs
(DUNN 1940a).
Bachia blairi (DUNN 1940a): Proc. Acad. Nat.
Sci. Philadelphia 92: 115; type locality:
Puerto Armuelles, Chiriqui, Panama. SVL to
70 mm. Southeastern Costa Rica and adja-
cent areas in Panama, sea level to 40 m ele-
vation.
Bachia pallidiceps (COPE 1862c), Proc. Acad.
Nat. Sci. Philadelphia 14: 356, type locality:
Rio Truando region, Colombia. SVL to 73
mm . Eastern Panama to northwestern
Colombia, sea level to 100 m elevation.
..t. Bachia blairi
• Bachia pallidiceps

Key to Bachia
1 a Pale dorsolateral stripes present; 22-24
scales at midbody; 45-48 cross rows of dor-
sal scales; 30-33 cross rows of ventral
scales ...... ..... ............ Bachia pallidiceps
b Without pale dorsolateral stripes; 24-25
scales at midbody; 43-45 cross rows of dor-
sal scales; 29-31 cross rows of ventral
scales .......... ......... .. ...... .... Bachia blairi
Further Reading
DIXON 1973, MCDIARMID & DEWEESE 1977
Fig. 165. Bachia pallidiceps (southeastern
Costa Rica). Photo: R. W. Van Devender
Echinosaura
The lizards of the genus Echinosaura are
small terrestrial "microteiids" that are dis-
tributed from western Panama across
northwestern Colombia to central-western
Ecuador. In his revision of the genus,
UZZELL (1965a) recognized a single species
(E. horrida) divided into three subspecies.
Recently, FRITTS et al. (2002) described a
new species of Echinosaura and elevated
the former subspecies to species. Another
new species was described by KOHLER et
al. (2004). Thus, as currently understood,
the genus Echinosaura contains five spe-
cies, two of which (E. palmeri and E. pana-
mensis) occur m southern Central
America.
Gymnophthalmidae
Although usually found in the vicinity of
streams, these lizards live somewhat more
terrestrially than the semiaquatic Neusti-
curus (UZZELL 1965a, b). According to
DUNN (1944), BREDER (1946), and UZZELL
(1965a), active Echinosaura can be observed
during the day as well as at night; how-
ever, they appear to be predominantly noc-
turnal (UZZELL 1965a). The animals are
active among the leaves of the ground
cover or hide under rocks and in rotting
plant material. Several reports state that
Echinosaura will become absolutely rigid
when handled (UZZELL 1965a, LEVITON &
ANDERSON 1966). The animals will stiffen
their bodies and press their limbs against
their sides. If they are turned on their
backs they will quickly turn over and fl)
freeze again. As with most microteiids,
Echinosaura produces clutches, each with
two soft-shelled eggs (UZZELL 1965a).
1.. N
~
Echinosaura palmeri BOULENGER 1911, Ann.
Mag. Nat. Hist. (8) 7: 23; type locality:
Noananoa, Rio San Juan, Choco, Colombia.
SVL to 70 mm. Colombia and eastern
Panama, near sea level to 1400 m elevation.
E chinosaura panamensis BARBOUR 1924,
Proc. New England Zool. Club 9: 8; type lo-
cality: La Loma, Bocas del Toro Province,
Panama. SVL to 71 mm. Western Panama,
300-1500 m elevation.
Fig. 166. Echinosaura panamensis (near Santa
Fe, Panama). Photo: G. Kohler
87
 Gymnophthalmidae
Fig. 167. Echinosaura panamensis (El Cope,
Panama). Photo: R. W. Van Devender
Key to Echinosaura
1 a Frontal plate transversely divided (Fig.
168a) ............. Echinosaura panamensis
b Frontal plate single (Fig. 168b) ... .............. .
.......... .. ......... ..... Echinosaura palmeri
Further Reading
DUNN 1944, UZZELL 1965a, FRITTS et al. 2002,
KOHLER et al. 2004
Gymnophthalmus
Gymnophthalmus speciosus is the sole
representative of the microteiids to extend
as far north as Mexico. Whereas this spe-
cies has, at times, been divided into a num-
ber of subspecies (MERTENS 1952c, PETERS
a. E. panamensis
Fig. 168. Dorsal head scalation in Echinosaura
(frontal orange).
88
Fig. 169. Gymnophthalmus speciosus (near
Guanagasapa, Escuintla, Guatemala).
Photo: G. Kohler
& DONOSO-BARROS 1970), more recent in-
vestigations indicate that such a division is
not justified (KOHLER 1996c). The usually
intensely red tail is particularly conspic-
uous, and gives this small lizard the local
name "Cola Roja" ("red tail"). These are
typical undergrowth dwellers that spend
their days searching for prey (mostly inver-
tebrates) under the wilted leaves. The
population density is apparently seasonal,
with an increase during the dry season
(TELFORD 1971). According to the investiga-
tions of TELFORD (1971) the female of G.
speciosus in Panama produces clutches of
1-2, sometimes as many as 3 or 4 eggs. The
egg-laying season, during which the fe-
males produce up to three clutches, lasts
from October to March, with hatchlings
• Echinosaura panamensis
• Echinosaura palmeri

Fig. 170. Gymnophthalmus speciosus (Isla de
Roatan, Honduras). Photo: G. Kohler
being observed in the months from
February to May (TELFORD 1971, ALVAREZ
DEL TORO 1983). The young measure 17-19
mm at hatching.
Gymnophthalmus speciosus (HALLOWELL
1861), Proc. Acad. Nat. Sci. Philadelphia 12:
484; type locality: Nicaragua. SVL to 40 mm.
Isthmus of Tehuantepec on the Pacific side
and Guatemala on the Caribbean side south-
wards into northern South America, as well
as in the Lesser Antilles, sea level to 1220 m
elevation. No subspecies are recognized
(KOHLER 1996a).
Further Reading
COLE et al. 1990 (evolution and systematics of
the genus); TELFORD 1971 (ecology, reproduc-
tion)
Gymnophthalmidae
Leposoma
Like most of the microteiids, the genus
Leposoma is also most heavily distributed
in South America. Only Leposoma rugiceps
and L. southi occur in southern Central
America. The males of L. southi generally
display an intensive orange coloration on
the underside of the body. Similar to spe-
cies of the genera Ptychoglossus and Gym-
nophthalmus, the Leposoma species are
also predominantly undergrowth dwellers
that are found between the wilting leaves
on the rain forest floor. The following
details on the reproduction of Leposoma
rugiceps are based on the investigations of
TELFORD (1971) in Panama. During the
egg-laying season, which lasts for several
months (May to November) during the
rainy season, the females produce clutches
of two eggs each. The young hatch with a
SVL of 16-19 mm. They are sexually ma-
ture within six to eight months.
Leposoma rugiceps (COPE 1869), Proc. Acad.
Nat. Sci. Philadelphia 20: 305; type locality:
Rio Magdalena, Colombia. SVL to 44 mm.
Eastern Panama to Colombia, sea level to 500
m elevation.
Leposoma southi RUTHVEN & GAIGE 1924,
Pap. Mus. Zool., Univ. Michigan 147: 1; type
locality: Progreso, Chiriqui, Panama. SVL to
55 mm. Southern Costa Rica to Colombia, sea
level to 700 m elevation.
Fig. 171. Leposoma rugiceps (Panama).
Photo: M. A. Guerra
89
 Gymnophthalmidae

Fig. 172. Leposoma southi (Isla Popa, Panama). Fig. 174. Leposoma southi (southwestern Costa
Photo: G. Kohler Rica). Photo: R. W. Van Devender

Key to Leposoma
1 a Frontonasal divided ........ Leposoma southi
b Frontonasal undivided Leposoma rugi,ceps

Further Reading
RUIBAL 1952, UZZELL & BARRY 1971 (taxonomy),
TELFORD 1971 (ecology)

Abb. 173. Leposoma rugiceps (Pearl Islands,

Panama). Foto: C. W. Myers
a. L. southii b. L. rugiceps

Fig. 175. Dorsal head scalation in Leposoma

(frontonasals orange).

• Neusticurus apodemus
• Prionodactylus vertebra/is

90
 Gymnophthalmidae

Neusticurus apodemus UZZELL 1965b, Bull.

Amer. Mus. Nat. Hist. 132: 298; type locality:
15 km SW San Isidro del General, 865 m ele-
vation, San Jose, Costa Rica. SVL to 47 mm.
Southern portion of the Puntarenas Province,
Costa Rica, near sea level to 900 m elevation.

Further Reading
UZZELL 1965b

Prionodactylus

Only one species of this predominantly

South American genus Prionodactylus
Fig. 176. Neusticurus apodemus (Palmar, Costa reaches the border area between Panama
Rica). Photo: R. W. Van Devender and Colombia in the extreme southern
portion of Central America. These agile
lizards are ground dwellers in the rain
forest, where they search for prey (mostly
insects and spiders) during the day. To
Neusticurus reproduce, the female P. uertebralis pro-
duces clutches of two eggs each, which are
Water tegus of the genus Neusticurus have only superficially buried (UZZELL 1973).
their primary distribution in South
America, with only one species,
Neusticurus apodemus, reaching southern Prionodactylus vertebralis (O'SHAUGHNESSY
1879), Ann. Mag. Nat. Hist. (5) 4: 298; type
Central America in Costa Rica. As with all locality: Intac, Ecuador. SVL to 68 mm.
species of this genus, N. apodemus is Extreme eastern Panama (bordering
semiaquatic, found along fast flowing Colombia) to northwestern Peru, 700-2500 m
streams in the rain forest. Usually water elevation.
tegus are found on rocks or roots by the
edge of the stream, and they flee into the Further Reading
UZZELL 1973
water at the slightest disturbance. Some
active N. apodemus can also be found in
leaf litter on the forest floor (UZZELL
1965b). Although representatives of the
genus Neusticurus are active primarily
during the day, N. apodemus has been
observed swimming in the water at night,
as well as during the day (UZZELL 1965b).
While swimming, Neusticurus will fold in
their legs alongside the body and hold only
the head above the water's surface while
making snake-like movements with the
body and tail. Nothing is known about
their reproduction, but most likely N.
apodemus, as with other Neusticurus spe-
cies, produces two eggs per clutch, which
are deposited by the females under rotting Fig. 177. Prionodactylus vertebralis (Cerro
logs (UZZELL 1965b). Mali, Darien, Panama). Photo: C. W. Myers
91
 Gymnophthalmidae
Fig. 178. Ptychoglossus festae (Nusagandi,
Panama). Photo: G. Kohler
Ptychoglossus
The primary distribution of the genus
Ptychoglossus is in northwestern South
America, with only a few species reaching
Central America. Whereas P. plicatus
reaches at least as far as northern Costa
Rica, P. festae and P. myersi are restricted
in their distribution to eastern Panama.
Ptychoglossus festae is most easily distin-
guished externally by the presence of small
lateral scales between the dorsal and ven-
tral scales. However, the only definitive dif-
ference between the other two species can
only be seen in the hemipenial morphology
(HARRIS 1994). The species of the genus
Ptychoglossus are small diurnal under-
growth dwellers. Thus, they occupy an eco-
logical niche similar to that of members of
the genera Leposoma and Gymnophthalmus.
• Ptychoglossus plicatus
92
Fig. 179. Ptychoglossus plicatus (Fortuna,
Costa Rica). Photo: G. Kohler
Ptychoglossus festae (PERACCA 1896), Boll.
Mus. Zool. Comp. Anat. Univ. Torino 11: 2;
type locality: Rio Cianati, Darien, Panama.
SVL to 55 mm. Eastern Panama and north-
ern Colombia; sea level to 800 m elevation in
rain forest.
Ptychoglossus myersi HARRIS 1994, Herpetol.
Monogr. 8: 250; type locality: cloud forest on
the SE slope of Cerro Pirre, 1000 m, 7°50'N,
77°43'W, Serrania de Pirre, Darien, Panama.
SVL to 52 mm. Border area of Panama and
Colombia; 800-1300 m elevation in cloud
forest.
Ptychoglossus plicatus (TAYLOR 1949b), Univ.
Kansas Sci. Bull. 33: 272; type locality:
Morehouse Finca, 5 miles SW Turrialba,
Cartago, Costa Rica. SVL to 66 mm. Costa
Rica to northern Colombia; 400-1890 m ele-
vation in rain forest and cloud forest.
• Ptychoglossus festae
"" Ptychoglossus myersi

Fig. 180. Ptychoglossus plicatus (Fortuna,
Costa Rica). Photo: G. Kohler
Fig. 181. Ptychoglossus myersi (between Rio
Jaque and Rio Imamad6, Darien, Panama).
Photo: C. W. Myers
Key to Ptychoglossus
1 a A longitudinal fold separates dorsal scales
from ventral scales (Fig. 182a) ................
.. .. .... ... ...... ...... ..... Ptychoglossus festae
b Without a longitudinal fold between dorsal
scales and ventral scales (Fig. 182b) ........ 2
2 a Hemipenis with 23-28 ring folds ..........
....... ...... ............ Ptychoglossus plicatus
b Hemipenis with 9-12 ring folds ............... .
. .. .. .. .. . . . .. .... .......... Ptychoglossus myersi
a. Ptychoglossus festae b. Pfychoglossus p/icatus
Abb. 182. Lateral body scales.
Further Reading
RUIBAL 1952, HARRIS 1994
Helodermatidae
Helodermatidae
Of the two surviving species of the genus
Heloderma, only one (H. horridum) is
found in Central America. These are robust
lizards that can reach a total length of over
70 cm, with massive heads and short,
powerful limbs. They have fat tails, which
are not capable of autotomy or regenera-
tion. The back is covered with large ossified
knobby scales. The tongue is fleshy, but
forked at the tip. The Heloderma species
are the only surviving venomous lizards.
Elongated, multi-lobed poison glands are
located along both sides of the lower jaw. A
separate opening from each lobe of the
gland leads to the base of the grooved
teeth, which are up to 6 mm in length.
When the teeth clamp down, the venom is
squeezed out of the glands by the pressure
of the jaw muscles and flows through the
grooved teeth into the wound primarily by
capillary action. Because Gila monsters
are shy and live very secretively, they pose
no threat to humans. Biting incidents
occur only when the animals are provoked
or captured. These otherwise calm lizards
can launch themselves with unexpected
speed and bite very suddenly. A Gila mon-
ster will not readily release anything that
comes between its powerful jaws, and it
will bite down forcibly. Gila monster bites
should be taken very seriously and can
lead to localized as well as systemic symp-
toms of envenomation, which absolutely
require medical attention (MEBS 1992).
In Mexico and Central America, three sub-
species of Heloderma horridum are recog-
nized: H. h. horridum (WIEGMANN 1829)
along the Pacific side of Mexico from south-
ern Sonora to Chiapas, H. h. alvarezi
BOGERT & MARTIN DEL CAMPO 1956 in the
Rio Grijalva Basin, Chiapas, and H . h.
charlesbogerti CAMPBELL & vANNIN! 1988 in
the Rio Motagua Basin, Guatemala. Gila
monsters are crepuscular and nocturnal
ground-dwellers in dry forest that become
active just after dusk. On cloudy days, they
can sometimes be observed during the day
outside of their hiding places (burrows,
93
 Helodermatidae

Fig. 183. Heloderma horridum charlesbogerti Fig. 184. Heloderma horridum horridum
(El Arenal, Zacapa, Guatemala). (Mixtequilla, Oaxaca, Mexico). Photo: G. Kohler
Photo: G. Kohler

which they have either excavated them-

selves or were pre-existing). They hunt
down their prey, which consists of nestling
rodents and birds, as well as bird and rep-
tile eggs, with constant tongue flicking.
Gila monsters can survive for several
months without food by living off the fat
stored in their tails. During the rainy sea-
son, these animals are more active than
during the drier months of the year.
Females will produce one clutch per year
(during the months from October to
December), consisting of 4-15 elongated,
thick-shelled eggs, which hatch after four Fig. 185. Heloderma horridum alvarezi (Tuxtla
to five months (ALVAREZ DEL TORO 1960, Gutierrez, Chiapas, Mexico). Photo: G. Kohler
FITCH 1970). Gila monsters are long-lived
animals, reaching an age of over 20 years.

Heloderma horridum (WIEGMANN 1829), Isis

von Oken 22: 421; type locality: Mexico. SVL
to 325 mm. Southern Sonora to Chiapas,
Mexico, also in Rio Motagua valley,
Guatemala, sea level to 1000 m elevation. For
subspecies see text above.

Further Reading
BOGERT & MARTiN DEL CAMPO 1956 (compre- Heloderma horridum
hensive work on the biology of this genus)

94
 Iguanidae

Iguanidae
The phylogenetic relationships and taxon-
omy of the pleurodont iguanians has been
subject to considerable controversy in
recent years. ETHERIDGE & DE QUEIROZ
(1988) demonstrated the existence of eight
monophyletic groups within the family
lguanidae that were reclassified as eight
families by FROST & ETHERIDGE (1989). The
latter workers were unable to find any evi-
dence for the monophyly of all pleurodont
iguanians (the former family lguanidae).
Initially, this new classification was
strongly criticized (BOHME 1990, LAzELL
1992) and later rejected on the basis of new Fig. 186. Adult male of Iguana i. rhinolopha
molecular data (MACEY et al. 1997, SCHULTE (Belize). Photo: G. Kohler
et al. 1998).

Most recently, FROST et al. (2001) have pres-
ented a modified classification in which
they recognized the pleurodont iguanians
as a monophyletic group, based on the evi-
dence provided by MACEY et al. (1997).
FROST et al. (2001) proposed to maintain
the subgroups as families and combine
these in a higher level taxon (Pleurodonta).
However, forthcoming work on this group
of lizards is likely to result in additional
changes (e.g., the former Tropiduridae and
Polychrotidae were shown to be paraphyl-
etic; see FROST et al. 2001), and there is a
high probability that some of these groups
will still be subject to change pending fur-
ther studies. Because the current status of
iguanian phylogenetics is still contentious,
it appears best to combine all pleurodont
iguanians in a single family (lguanidae)
and assign the status of subfamilies to the
monophyletic subgroups.
b Lamellae beneath fingers and toes smooth
or keeled but not widened (Fig. 187a-c);
extensile dewlap present or not ............... 3
3 a Head noticeably extended posteriorly, ei-
ther as vertical fin or horizontal shelf (hel-
met, crown or fin; Fig. 189), most conspic-
uous in adult males .................................. 4
b Head not extended or drawn out posterior-
ly to form helmet, crown or fin ............... 6
4 a Each subdigital lamella with a knob (Fig.
187a); horizontal fin (crown) on head ......... .
......................................... Laemanctus
knob
fringe
widened lamellae

Key to lguanidae genera

1 a Head with several horn-like projections; a
lateral abdominal fringe with soft spines
present (Fig. 409, p. 149) ...... Phrynosoma
b Head without horn-like projections, or, if
head with horn-like projections, then no
lateral abdominal fringe with soft spines 2
2 a Fingers and toes with widened smooth a. Laemanctus b. Basiliscus c. Sceloporus d. Ano/is
subdigital lamellae (Fig. 187d); males with
an extensile dewlap ....... anoles (genera
Anolis, Ctenonotus, Dactyloa, Norops) Fig. 187. Ventral surface of toes.
95
 Iguanidae

b Subdigital lamellae without knobs (Fig.

187b); vertical fin or helmet on head ....... 5
5 a Toes of hind foot with fringe of flat scales
forming serrate margin (Fig. 187b); lateral
body scales homogeneous ........ Basiliscus
Basiliscus Corytophanes Laemanctus
b Toes of hind foot without fringe of flat plumifrons cristatus serratus
scales forming serrate margin; lateral body
scales heterogeneous ......... Corytophanes Fig. 189. Lateral view of head.
6 a Tail with whorls of enlarged spinous scales
(Figs. 190a, 192) ........ ............................... 7
b Tail without spines ................................... 8
7 a A large, elongate subocular scale present
(Fig. 190b) ........ ... ................... Ctenosaura
b Orbit bordered below by an arc of several
subequal scales ................ Morunasaurus
8 a Large pendulous dewlap present; one or b.
two conspicuously enlarged rounded scales
below tympanum; SVL to 50 cm ... Iguana
b No large pendulous dewlap; no conspic- Fig. 190. a. Scalation of tail in Ctenosaura
uously enlarged scales below tympanum; quinquecarinata;
small to medium-sized lizards .......... ....... 9 b. Lateral head scales in Ctenosaura similis
(enlarged subocular scale orange).
9 a Very large interparietal plate present (Fig.
191a) ......................... ... .... .... ................. 10
b Interparietal plate, if present, small (Fig.
191b) ........................................................ 11
lOa A dorsolateral longitudinal fold alongside
body present, usually with some enlarged
scales; a complete transverse gular fold,
differentiated from adjacent gular scales
by very fine scales ...... .. .......... Urosaurus
b No dorsolateral longitudinal fold alongside
body; no transverse gular fold .. .. ..... .
. . . .. . . .. . . . . . ..... .... . . . . . . . . . . . . . . ....... Sceloporus
lla Tail about three times SVL; no vertebral
row of enlarged scales .............. Polychrus
b Tail less than twice SVL, a vertebral row of a. Sceloporus (inter-
parietal scale shaded)
enlarged scales present .... ..... Enyalioides
Fig. 191. Dorsal head scalation (interparietal
orange).

Fig. 188. Urosaurus bicarinatus (ventral view). Fig. 192. Tail of Morunasaurus annularis.
Arrow points to gular fold. Photo: G. Kohler Photo: G. Kohler

96
 Iguanidae

relationships of the anoles is still too

Anoles (genera Anolis, Ctenonotus,
limited to accurately define the natural
Dactyloa, Norops) groupings.

The anoles (Anolis sensu lato), with more
than 300 valid species, represent the most
diverse lizard genus, with more species
being described almost every year. They
are a conspicuous component of the fauna
wherever they occur, and throughout their
range it is common for several species of
anoles to occur together. Despite the fact
that anoles have been the focus of interest
by herpetologists for more than a century,
the phylogenetic relationships and taxon-
omy of this group of lizards remains sub-
ject to serious controversy in the recent
literature. Indeed, our understanding of
the systematics of these lizards is still
poor. In the present work, I follow the clas-
sification proposed by GUYER & SAVAGE
(1987, 1992) and SAVAGE & GUYER (1989) in
using the generic names Anolis, Cteno-
notus, Dactyloa, and Norops for the anole
species of Central America. This decision
reflects the strong support for the assertion
that Norops represents a monophyletic
lineage. Norops is the beta section of anoles
of ETHERIDGE (1959), whose species are
characterized by autotomic caudal verte-
brae with transverse processes that are
directed anterolaterally; see figure 3A in
ETHERIDGE (1967). However, the appro-
priateness of GUYER and SAVAGE's (1987,
1992) other proposed genera (the alpha
group of ETHERIDGE 1959) appear more
uncertain. The anole classification of
GUYER & SAVAGE (1987) was heavily
criticized by CANNATELLA & DE QUEIROZ
(1989) and by E. WILLIAMS (1989).
Most species are bush or tree-dwellers, al-
though some are only found amongst the
leaves on the ground. Even some semi-
aquatic species that occur only in the vici-
nity of water, such as Norops barkeri, N
lionontus, N poecilopus, and N aquaticus,
are found in Central America. Typical leaf-
dwellers in rain forest are N humilis and
N uniformis, which in some areas reach a
high population density (KOHLER 1998c).
The true giants, however, amongst the
Central American anoles are found in the
genus Dactyloa, such as D. insignis (SVL
to 160 mm). These are tree-dwellers that I'll
are difficult to spot in the rain forest. With
luck, they can be observed on the trunks of
large trees where the animals usually sit
1...
N
i-:1
with their heads pointed down. According
to SCOTT et al. (1976), these animals usual-
ly remain on tree trunks at heights be-
tween 40 and 200 cm above the ground,
although specimens sitting up higher
would be difficult to observe (SAVAGE &
TALBOT 1978). All anole species reproduce

For most "species groups" of anoles, as pro-

posed by various researchers in the litera-
ture, evidence that these actually repre-
sent monophyletic groups is meager. Some
of them probably are natural groups of spe-
cies, while others represent artificial
groupings of species that are superficially
similar and are placed together because of
particular character combinations. At this Fig. 193. Norops sericeus (copulating,
time, our knowledge of the phylogenetic Mazatenango, Guatemala). Photo: G. Kohler

97
 Iguanidae
by laying eggs, and, during the breeding
season, will produce several clutches, each
consisting of a single egg, at intervals of
one to two weeks. The ovary and oviduct on
each side of the body are alternately active.
As soon as one egg is laid from the right
oviduct, for example, a new follicle is pro-
duced by the left ovary and passes through
the left oviduct, where it is fertilized and
shelled within a few days, before being
laid. In the meantime, the right ovary has
produced a new follicle, which is ready to
ovulate. As long as there is enough food
available and the weather conditions are
suitable, this cycle will continue and eggs
will be laid at regular intervals (FITCH
1982).
The study of hemipenis morphology has
shown to be a valuable tool for defining
species bounderies in mainland anoles. In
the species studied up to date, hemipenis
morphology was shown to exhibit phenoty-
pic stability across the species' ranges,
covering large geographic areas in some
taxa (KOHLER & SUNYER 2008). Based on
difference in hemipenis morpology, several
taxonomically cryptic species of anoles
have been detected indicating a substanti-
al cryptic diversity among Central
American anoles (KOHLER & KREUTZ 1999;
KOHLER et al. 2003, KOHLER et al. 2007'
KOHLER & SUNYER 2008). Obviously, there
are some very effective barriers in place
that prevent hybridization between these
very similar species. More research is nee-
ded to study the distribution and possible
interactions of these species in their
various contact zones.
In the following list, information for body
and dewlap coloration are for live speci-
mens; except where otherwise indicated,
dewlap coloration pertains only to adult
males. Norops binotatus (PETERS 1863b) is
excluded because there appears to be no
valid record for this species in Central
America. BARBOUR (1923) erroneously
reported this taxon for Panama, based on
misidentified N vittigerus (KOHLER unpubl.
data).
98
Anolis allisoni BARBOUR 1928, Proc. New
England Zool. Club 10: 58; type locality:
Coxen Hole, Ruatan, Bay Islands, Honduras.
SVL to 87 mm. Body green, dewlap pink.
Islas Guanaja and Roatan, Cayos Cochinos,
Honduras, as well as in Belize and on Cuba,
sea level to 300 m elevation.
Ctenonotus cristatellus (DUMERIL & BIBRON
1837), Erp. Gen. 4: 143; type locality:
Martinique [in error]. SVL to 70 mm. Body
brown, dewlap greenish yellow with orange
margin. Puerto Rico and Virgin Islands;
introduced in Costa Rica, sea level to 750 m
elevation; occurrence on Isla Cozumel doubt-
ful (LEE 1996).
Dactyloa casildae (AROSEMENA, IBANEZ & DE
SOUSA 1992, Rev. Biol. Trop. 39: 255-262; type
locality: Margenes de Quebrada Frank, 1100
m, 8°44'N, 82°13'W, Reserva Foresta! de
Fortuna, Chiriqui, Panama. SVL to 108 mm.
Body green with 4-6 oblique brown bands
(not formed of adjacent ocelli) interspersed by
blue-outlined yellow patches; some females
have a broad, dark brown mid-dorsal stripe
extending from nape to base of tail; dewlap
dirty cream with broad yellow scale rows
irregularly interspersed with smaller eme-
rald green scales. Fortuna region in the
Provinces Chiriqui and Bocas del Toro,
Panama, 1050-1400 m elevation.
Dactyloa chloris (BOULENGER 1898), Proc.
Zool. Soc. London 1898: 110; type locality:
Paramba, Ecuador. SVL to 60 mm. Body
green, dewlap yellowish white, pale blue
anteriorly. Eastern Panama to northwestern
Ecuador.
Dactyloa chocorum (WILLIAMS & DUELLMAN
1967), Breviora, Mus. Comp. Zool. 256: 2; type
locality: Rio Tuira at Quebrada La Plata, 100
m, Darien, Panama. SVL to 79 mm in males,
to 73 mm in females. Body green with or
Fig. 194. Dactyloa chloris (Rio Jaque, Darien,
Panama). Photo: C. W. Myers
Fig. 195. Anolis allisoni (Isla de Roatan, Fig. 198. Ctenonotus cristatellus (Cahuita,
Honduras). Photo: G. Kohler Costa Rica) Photo: D. M. Dehling

Fig. 196. Dactyloa chocorum (Reserva Foresta} Fig. 199. Dactyloa chocorum (Cerro Narices,
Fortuna, Panama, 400 masl). Photo: B. Akeret Panama). Photo: S. Lotzkat

Fig. 197. Dactyloa casildae (female, Reserva Fig. 200. Dactyloa casildae (male, La Nevera,
Foresta} Fortuna, Panama). Photo: S. Lotzkat Panama). Photo: S. Lotzkat
99
 lguanidae

without darker green oblique bands; dewlap

pale orange with green lines. Southern Costa
Rica to Colombia, 100-1100 m elevation.
Dactyloa frenata (COPE 1899), Sci. Bull.
Philadelphia Mus. 1: 6; type locality: unknown.
SVL to 143 mm in males, to 118 mm in fe-
males. Body green with oblique rows of dark
olive green ocelli; dewlap uniformely cream-
colored (occasionally with black shading).
Costa Rica to northern Colombia, sea level to
820 m elevation in lowland and premontane
rain forests.
Dactyloa kunayalae (HULEBAK, POE, IBANEZ &
WILLIAMS 2007), Phyllomedusa 6 (1): 6; type
locality: "Parque Nacional General de
Division Omar Torrijos Herrera, 5 km N El
Cope, Cocle Province, Panama (8°40.315'N,
80°35.518'W)." SVL to 109.3 mm. Body green
with oblique rows of dark olive green ocelli,
males with blue coloration on neck and anteri-
or body; basal portion of male dewlap orange
with large white basal portion and multiple
rows of yellow-green or blue scales; female
dewlap pale bluish-green with orange mar-
gin. Central Panama, 370-800 m elevation in
primary and secondary rain forest.
Dactyloa insignis (COPE 1871), Proc. Acad.
Nat. Sci. Philadelphia 24: 213; type locality:
San Jose, Costa Rica. SVL to 160 mm in
males, to 135 mm in females. Body yellowish
gray to tan or greenish with a distinct large,
pale-centered, dark-outlined blotch above
shoulder; dewlap orange red, with or without
several horizontal green or white bars and
dark spotting and with a white to greenish
free margin. Costa Rica and Panama, sea
level to 1600 m elevation.
Dactyloa microtus (COPE 1871), Proc. Acad. Fig. 202. Dactyloa frenata (Costa Rica).
Nat. Sci. Philadelphia 24: 214; type locality: Photo: A. Flaschendrager
San Jose, Costa Rica. SVL to 111 mm in
males; to 104 mm in females. Body yellowish
gray to tan with several pale brown-centered
dark bands that may change to green on
flanks and a distinct postorbital dark-bor-
dered pale stripe; dewlap salmon with white
scales. Costa Rica and Panama, 425-1500 m
elevation.
Dactyloa latifrons (BERTHOLD 1845), Nachr.
Georg-Augusts Univ. und K. Ges. Wiss.
Gi:ittingen 3: 39; type locality: western
Colombia (see MYERS & BoHME 1996). SVL to
124 mm in males, to 97 mm in females. Body
green with a distinct large, pale-centered,
dark-outlined blotch above shoulder; dewlap
pale greenish gray with pale yellow scales.
Eastern Panama to northwestern Ecuador.
Fig. 203. Dactyloa latifrons (near Noanama,
Darien, Panama). Photo: C. W. Myers
100
Fig. 204. Female Dactyloa kunayalae Fig. 207. Male Dactyloa kunayalae (Panama).
(Panama). Photo: S. Poe Photo: S. Poe

Fig. 205. Subadult female of Dactyloa microtus Fig. 206. Dactyloa microtus (adult male, La
(La Nevera, Panama). Photo: G. Kohler Nevera, Panama). Photo: S. Lotzkat

Fig. 208. Adult female of Dactyloa insignis Fig. 209. Dactyloa insignis (Las Cruces, Punta-
(near Santa Fe, Panama). Photo: G. Kohler renas, Costa Rica). Photo: K.-H. Jungfer
101

Fig. 210. Norops altae (San Rafael de
Varablanca, Costa Rica).
Fig. 211. Norops alvarezdeltoroi (Chiapas,
Mexico).
Fig. 212. Norops amplisquamosus (Cusuco
National Park, Honduras). Photo: G. Kohler
Fig. 213. Norops anisolepis (San Cristobal de
las Casas, Chiapas, Mexico). Photo: G. Kohler
102
Norops altae (DUNN 1930b), Proc. New
England Zool. Club 12: 22; type locality: Finca
Acosta, 7000 ft, Volcan Barba, Costa Rica.
SVL to 52 mm. Body grayish brown, dewlap
rust orange grading to yellow on free margin.
Highlands of Costa Rica, 1220-2000 m eleva-
tion.
Photo: G. Kohler Norops alvarezdeltoroi (NIETO MONTES DE
OcA 1996), J . Herpetol. 30: 20; type locality:
19,5 km N, 8,1 km W Ocozocoautla, 940 m,
l6°56'N, 97°27'W, Chiapas, Mexico. SVL to 66
mm. Body grayish brown, dewlap dark red
with a suffusion of black pigment. Region of
El Ocoto, Chiapas, Mexico, 910-940 m eleva-
tion.
Norops amplisquamosus MCCRANIE, WILSON
& WILLIAMS 1992, Carib. J. Sci. 28: 209; type
locality: Finca El Cusuco, 1550 m, 15 3l'N, 88
12'W, 5,6 km WSW Buenos Aires, Sierra de
Omoa, Cortes, Honduras. SVL to 46 mm.
Body grayish brown, dewlap yellow-orange.
Only known from type locality, 1530-1720 m .
Norops anisolepis (SMITH, BURLEY & FRITTS
1968), J. Herpetol. 2: 147; type locality: 10 mi
Photo: G. Kohler SE San Christ6bal Las Casas, Chiapas,
Mexico. SVL to 47 mm. Body grayish brown,
dewlap pinkish red . Meseta Central de
Chiapas, Mexico, 1500-2500 m elevation.
Norops aquaticus (TAYLOR 1956), Univ.
Kansas Sci. Bull. 38: 141; type locality:
Palmar, Puntarenas, Costa Rica. SVL to 71
mm. Body grayish yellowish green with broad
transverse brown bands and a pale greenish
yellow lateral longitudinal stripe; dewlap
red-orange with yellow stripes. Costa Rica
and Panama, 30-1170 m elevation.
Norops apletophallus (KOHLER & SUNYER
2008), Herpetologica 64 (1): 98; type locality:
"Panama City, Metropolitan National Park
(8°58'60"N, 79°32'46"W), 45 m, Panama
Province, Panama." SVL to 4 7 mm. Body
grayish brown; dewlap almost uniformly
orange. Central and eastern Panama, sea
level to 470 m elevation.
Norops auratus (DAUDIN 1802), Hist. Nat.
Rept.: 89; type locality: unknown. SVL to 52
mm. Body grayish brown, dewlap blue.
Central Panama and northern South
America, sea level to 2000 m elevation.
Norops barkeri (SCHMIDT 1939), Field Mus.
Nat. Hist. Puhl. Zool. Ser. 24: 7; type locality:
Cascajal, upper Uzpanapa River, Veracruz,
Mexico. SVL to 101 mm. Body grayish brown,
dewlap dark orange with yellowish orange
margin. Caribbean versant of Isthmus of
Tehuantepec (Veracruz, Oaxaca,Tabasco, and
Chiapas, Mexico), 200-1100 m elevation.
Fig. 214. Norops apletophallus (Mahe, Fig. 216. Norops auratus (Los Algarrobos,
Panama). Photo: A. Batista Chiriqui, Panama). Photo: G. Kohler

Fig. 215. Norops aquaticus (Las Cruces, Punta- Fig. 217. Norops barkeri (Chiapas, Mexico).
renas, Costa Rica). Photo: G. Kohler Photo: G. Kohler

Norops bicaorum KOHLER 1996b, Sencken-

bergiana biol. 76: 21; type locality: trail to
Rock Harbour, about 3 km north of the town
of Utila (16° 6,34'N; 86° 53,94'W), Isla de
Utila, Islas de la Bahia, Honduras. SVL to 75
mm. Body grayish brown, dewlap orange with
suffusion of black pigment centrally and with
white scales. Endemic on Isla de Utila, Islas
de la Bahia, Honduras, sea level to 100 m ele-
vation.
Norops biporcatus (WIEGMANN 1834), Herp.
Mex. 47; type locality: Santa Rosa de Pansos
[=Panzos], Alta Verapaz, Guatemala. SVL to
102 mm. Body green (capable of rapid color Fig. 218. Norops bicaorum (Isla de Utila,
change to brown), dewlap blue with dark Honduras). Photo: G. Kohler
orange margin. Southern Mexico through
Central America to northern South America,
sea level to 2000 m elevation. The nominate
subspecies occurs in Central America.
KOHLER & BAUER (2001) demonstrated that
the holotype of Norops biporcatus (WIEGMANN
1834) is conspecific with the syntypes of N
petersii (BocoURT 1873). In order to stabilize
the current and long established usage of the
names N biporcatus and N petersii, the
International Commission on Zoological
Nomenclature (ICZN) set aside the status of
the original holotype of N biporcatus and
designated MNHM 2426 (holotype of Anolis
copei BOCOURT 1873) as the neotype of N
biporcatus (ICZN 2002). Fig. 219. Norops biporcatus (Bartola,
Nicaragua). Photo: G. Kohler
103
 Iguanidae
Naraps capita (PETERS 1863b), Monats. Akad.
Wiss. Berlin 1863: 142; type locality: Costa
Rica. SVL to 96 mm. Body grayish brown
with green elements, dewlap yellowish green.
Caribbean versant of southern Mexico
through Central America to eastern Panama
and probably adjacent regions in Colombia,
sea level to 1300 m elevation.
Naraps campbelli KOHLER & SMITH 2008
Herpetologica 64 (2): 207; type locality: "traii
to L~~un~ Yolnabaj, Aldea Yalambojoch,
Mumc1pahdad Nenton, Departamento de
Huehuetenango, Guatemala, 1540 m elevati-
on, l6°00'17"N, 91°34'05"W.". SVL to 51.3
mm. Body grayish brown, dewlap color
unknown. Known only from two localities in
the Sierra de los Cuchumatanes, western
Guatemala, at 1540 and 1660 m elevation.
Naraps carpenteri (ECHELLE, ECHELLE &
FITCH 1971), Herpetologica 27: 355; type lo-
cality: Rio Reventazon, 7 km ESE Turrialba,
Cartago, Costa Rica. SVL to 45 mm. Body
greenish, dewlap orange. Nicaragua, Costa
Rica, and Panama, sea level to 1100 m eleva-
tion. Anolis pracellaris MYERS 1971a is
Fig. 220. Naraps capita (female, Reserva
Forestal Fortuna, Panama). Photo: G. Kohler
Fig. 221. Naraps campbelli (female, Huehue-
tenango, Guatemala). Photo: J. A. Campbell
104
regarded to be a synonym of N. carpenteri
according to FITCH et al. (1976).
Naraps cabanensis (STUART 1942), Occ. Pap.
Mus. Zool. Michigan 464: 6; type locality: 3
km S Finca Samac (6 airline km of Cohan),
1350 m, Alta Verapaz, Guatemala. SVL to 50
mm. Body grayish brown, dewlap purple.
Highlands on the Caribbean versant of
Guatemala, 1300-1410 m elevation.
Naraps campressicauda (SMITH & KERSTER
1955), Herpetologica 11: 193; type locality:
Near La Gloria, Oaxaca, Mexico. SVL to 55
mm. Body grayish brown, dewlap uniform
rose to purple. Disjunct populations in
east~rn Oaxaca and western Chiapas,
Mexico, 500-1200 m elevation.
Noraps crassulus (COPE 1864), Proc. Acad.
Nat. Sci. Philadelphia 16: 173; type locality:
Cohan, Alta Verapaz, Guatemala. SVL to 59
mm. Body grayish brown, dewlap red.
Highlands of northern Central America
(southern Mexico, Guatemala, Honduras, and
El Salvador), 1300-3000 m elevation.
Naraps cristifer (SMITH 1968a), Trans. Kansas
Acad. Sci. 71: 195; type locality: near
Acacoyagua, Chiapas, Mexico. SVL to 88 mm.
Body grayish brown. Pacific versant of
Chiapas, Mexico, and western Guatemala.
Naraps cryptalimifrans (KOHLER & SUNYER
2008), Herpetologica 64 (1): 101; type locality:
"Cerro Brujo (9°11'16.4"N, 82°11'25.4"W), 10
m, Bocas del Toro Province, Panama." SVL to
45 mm. Body grayish brown; dewlap dirty
white with a small basal orange blotch.
Caribbean slope of eastern Panama and adja-
cent Costa Rica, sea level to 250 m elevation.
Naraps cupreus (HALLOWELL 1861), Proc.
Acad. Nat. Sci. Philadelphia 12: 481; type
locality: Nicaragua. SVL to 57 mm. Body
grayish brown, dewlap brown with orange
margin. Honduras to Costa Rica, sea level to
1250 m elevation.
Naraps cuprinus (SMITH 1964), Herpetologica
20: 31; type locality: Vicinity of Zanatepec,
Oaxaca, Mexico. SVL to 69 mm. Body grayish
brown, dewlap red. Eastern Oaxaca and
western Chiapas, Mexico, 200-500 m eleva-
tion.
Naraps cusuca MCCRANIE, KOHLER & WILSON
2000? Senckenbergiana biol. 80: 214; type
locahty: Parque Nacional El Cusuco Centro
de Visitantes, 15°29.92'N, 88°12.88'W, 1550 m
elevation, Cortes, Honduras. SVL to 46 mm.
Body grayish brown, dewlap unicolor dirty
white. Known only from vicinity of type lo-
cality, 1550-1935 m.
Fig. 222. Norops carpenteri (Bartola, Rio San Fig. 226. Norops cristifer (Depto. Retalhuleu,
Juan, Nicaragua). Photo: G. Kohler Guatemala). Photo: L. Melendez

Fig. 223. Norops cobanensis (Pasado El Fig. 227. Norops cryptolimifrons (Cerro Brujo,
Quetzal, Baja Verapaz, Guatemala). Bocas de Toro, Panama). Photo: G. Kohler
Photo: G. Kohler

Fig. 224. Norops compressicauda (El Ocote Fig. 228. Norops cupreus (Selva Negra,
region, Chiapas, Mexico). Photo: G. Kohler Matagalpa, Nicaragua). Photo: G. Kohler

Fig. 225. Norops crassulus (near Santa Rosa Fig. 229. Norops cusuco (Cerro Cusuco, Cortes,
Pass, Quiche, Guatemala). Photo: G. Kohler Honduras). Photo: G. Kohler
105
 Iguanidae
Norops datzorum (KOHLER, PONCE, SUNYER &
BATISTA 2007), Herpetologica 63 (3): 385; type
locality: "La Nevera, 8°29'45"N, 81°46'35"W,
1600 m elevation, Serrania de Tabasara,
Comarca Ngobe Bugle, Distrito de Nole
Duima, Corregimiento de Jadeberi, Panama".
SVL to 49 mm. Body greenish, dewlap color
unknown. Serrania de Tabasara, Panama.
Norops dollfusianus (BocoURT 1873), Miss.
Sci. Mex., Rept. 1873: 84; type locality: Volcan
Atitlan, 1200 m, Guatemala. SVL to 44 mm.
Body grayish brown, dewlap yellow. Pacific
versant of eastern Chiapas, Mexico, and
western Guatemala, 200-1200 m elevation.
Norops fortunensis (AROSEMA & IBANEZ D.
1993), Rev. Biol. Trop. 41: 267; type locality:
Reserva Foresta! de Fortuna, along the Rio
Chiriqu:i, in front of the estacion hidromete-
reol6gica del Instituto de Recursos
Hidraulicos y Electrificaci6n (IRHE) in Bijau,
1050-1075 m elevation, Chiriqu:i Province,
Panama. SVL to 48 mm. Body grayish brown,
dewlap orange red with red margin and yel-
low scales. Fortuna region, Chiriqu:i and
Bocas del Toro Provinces, Panama, 1000-1200
m elevation. Anolis exsul AROSEMENA &
IBANEZ D. 1994 is regarded to be a synonym of
N fortunensis according to PONCE & KOHLER
(2008).
Norops fungosus (MYERS 1971b), Amer. Mus.
Novitat. 2471: 13; type locality: north slope
Cerro Pando, 1450 m, Cordillera de
Talamanca, Bocas del Toro, Panama. SVL to
4 7 mm. Body grayish brown, dewlap red.
Cordillera de Talamanca in Panama and
Costa Rica, 1200-1600 m elevation.
Norops fuscoauratus (D'ORBIGNY 1837), in
DUMERIL & BIBRON: Erp. Gen. 4: 110; type
locality: Chile [in error). SVL to 50 mm. Body
grayish brown, dewlap pink to pinkish-
brown. Eastern Panama; widely distributed
in the Amazon Basin, sea level to 1000 m.
Fig. 230. Norops datzorum (female holotype).
Photo: G. Kohler
106
Norops gruuo (KOHLER, PONCE, SUNYER &
BATISTA 2007), Herpetologica 63 (3): 376; type
locality: "near the headwaters of Rio San
Felix, ca. 2 km N Escopeta Camp, ca. 8°32'N,
81°50'W, Serrania de Tabasara, 900 m eleva-
tion, Comarca Ngobe Bugle, Distrito de Nole
Duima, Corregimiento de Jadeberi, Panama''.
SVL to 47 mm. Body grayish brown, dewlap
orange. Only known from type locality.
Norops haguei (STUART 1942b), Occ. Pap.
Mus. Zool. Univ. Michigan 464: 3; type local-
ity: 2 km south of Finca Chichen, 1750 m,
Alta Verapaz, Guatemala. SVL to 53 mm.
Body grayish brown, dewlap red. Only known
from type locality.
Norops heteropholidotus (MERTENS 1952a),
Zool. Anz. 148: 89; type locality: Hacienda Los
Planes, 2000 m, Miramundo, Santa Ana, El
Salvador. SVL to 59 mm. Body grayish brown,
dewlap red with pale to dark brown scales.
Highlands of northern El Salvador and south-
western Honduras, 1870-2200 m elevation.
Norops hobartsmithi (NIETO-MONTES DE 0CA
1995), Puhl. Espec. Mus. Zool. Facultad de
Ciencias, UNAM, 10: 160; type locality:
Parador Selva Negra on Mexico Highway 195
(approx. 0.5 km W Pinabeto), 1992 m,
l 7°12'58"N, 92°57'4 7"W, Municipality of
Rayon, at the border with Municipality of
Pueblo Nuevo Solistahuacan, Chiapas,
Mexico. SVL to 49 mm. Body grayish brown,
dewlap rose. Highlands of northwestern
Chiapas, Mexico, 1525-2000 m elevation.
Norops humilis (PETERS 1863b), Monats.
Akad. Wiss. Berlin 1863: 138; type locality:
Veragua, Panama. SVL to 44 mm. Body gray-
ish brown, dewlap red-orange with yellow
margin or dewlap uniform red orange (popu-
lations in eastern Panama). Northern Costa
Rica along both versants to central Panama
east of the Canal Zone, sea level to 1600 m
elevation.
Norops johnmeyeri (WILSON & MCCRANIE
1982), Trans. Kansas Acad. Sci. 85: 133; type
locality: 5.6 km WSW Buenos Aires, 1580 m,
El Cusuco, Cortes, Honduras. SVL to 70 mm.
Body grayish brown, dewlap orange red with
large central blue blotch. Females with well
developed dewlap (yellow with large central
blue blotch). Sierra de Omoa (Dptos. Copan
and Cortes), Honduras, 1410-1995 m.
Norops kemptoni (DUNN 1940a), Proc. Acad.
Nat. Sci. Philadelphia 92: 111; type locality:
Finca Lerida, Chiriqu:i, Panama. SVL to 55
mm. Body grayish brown, dewlap red with
orange blotch anteriorly, and with white
scales. Highlands of Chiriqu:i, Panama, 1000-
Fig. 231. Norops dollfusianus (Finca Irlanda, Fig. 236. Norops gruuo (male, type locality).
Chiapas, Mexico). Photo: G. Kohler Photo: G. Kohler

Fig. 232. Norops fortunensis (Reserva Forestal Fig. 237. Norops heteropholidotus (El Pital,
Fortuna, Panama). Photo: M. Ponce Honduras). Photo: G. Kohler

Fig. 234. Norops fungosus (Reserva Forestal Fig. 238. Norops humilis (Monteverde, Costa
Fortuna, Panama). Photo: A. Batista Rica). Photo: G. Kohler

Fig. 235. Norops fuscoauratus (Guaquira, Fig. 239. Noropsjohnmeyeri (Cusuco National
Venezuela). Photo: J. Sunyer Park, Cortes, Honduras). Photo: G. Kohler
107
 Iguanidae
1800 m elevation. Anolis pandoensis SAVAGE
& GUYER 1998 is regarded to be a synonym of
N kemptoni according to HULEBAK & PoE
(2006) and PONCE & KOHLER (2008).
Norops kreutzi MCCRANIE, KOHLER & WILSON
2000, Senckenbergiana biol. 80: 218; type
locality: 2.5 airline km NNE La Fortuna,
l5°26'N, 87°18'W, 1670-1690 m elevation,
Yoro, Honduras. SVL to 51 mm. Body grayish
brown, dewlap pale yellow with purple scales.
Known only from the type locality.
Norops laeviventris (WIEGMANN 1834),
Herpetologica Mexicana: 47; type locality:
Mexico. SVL to 52 mm. Body grayish brown,
dewlap dirty white. Southern Mexico to
western Panama, 1160-2000 m elevation.
Based on the studies of BOTTCHER (1999) and
KOHLER (2001e) I consider Anolis intermedius
PETERS 1863b, Anolis nannodes COPE 1864,
and Anolis bourgeaei BOCOURT 1873 to be
synonyms of N laeviventris .
Norops lemurinus (COPE 1861d), J. Acad. Nat.
Sci. Philadelphia 12: 213; type locality:
Veragua, Panama. SVL to 75 mm. Body gray-
ish brown, dewlap red (often with black
scales). Central Veracruz, Mexico, along
Caribbean versant of Central America to cen-
tral Panama; also on Pacific versant in Costa
Rica and Panama, sea level to 1080 m eleva-
tion.
Norops limifrons (COPE 1862b), Proc. Acad.
Nat. Sci. Philadelphia 13: 178; type locality:
Veragua, Panama. SVL to 48 mm. Body gray-
ish brown, dewlap dirty white with a basal
orange-yellow spot or (populations in central
and eastern Panama) dewlap almost com-
pletely orange-yellow. Eastern Honduras to
Panama, sea level to 1350 m elevation. Anolis
biscutiger TAYLOR 1956 and Anolis godmani
BOULENGER 1885 are both regarded to be
synonyms of N limifrons according to SAVAGE
(2002).
Fig. 240. Norops kemptoni (male, Alto
Chiquero, Chiriqui, Panama). Photo: G. Kohler
108
Norops lionotus (COPE 1861d), J . Acad. Nat.
Sci. Philadelphia 12: 210; type locality:
Cucuyas de Veragua, Panama. SVL to 78 mm.
Body grayish brown, dewlap yellow to yellow-
orange. Eastern Honduras to central
Panama, sea level to 1200 m elevation. Based
on the studies of OBERMEIER (1998), I consider
Anolis oxylophus COPE 1876 to be a synonym
of N lionotus.
Norops loveridgei (SCHMIDT 1936a), Proc.
Biol. Soc. Washington 49: 47; type locality:
Portillo Grande, 4100 ft, Yoro, Honduras. SVL
to 118 mm. Body grayish brown, dewlap or-
ange with pink streaks. Mountainous regions
in the Dptos. de Yoro and Atlantida,
Honduras, 550-1100 m elevation.
Norops macrophallus (WERNER 1917), Mitt.
Zool. Mus. Hamburg 34: 31; type locality: San
Jose, Guatemala. SVL to 45 mm. Body gray-
ish brown, dewlap rose with a orange yellow
basal blotch. Pacific versant of northern
Central America (Guatemala and El
Salvador), 40-1320 m elevation.
Norops magnaphallus (POE & IBANEZ 2007),
Journal of Herpetology 41 (2): 264; type loca-
lity: "eastern entrance to Sendero Quetzales,
8 km north of Boquete, approximately
08°49.0'N, 82°26.6'W, Chiriqui Province,
Panama". SVL to 55 mm. Body grayish
brown, dewlap brick red with white scales.
Known only from the region around Boquete
and Cerro Horqueta, western Panama.
Norops matudai (SMITH 1956), Herpetologica
12: 1; type locality: region of Soconusco,
Chiapas, Mexico. SVL to 42 mm. Body gray-
ish brown, dewlap purple. Sierra Madre de
Chiapas, Chiapas, Mexico, and adjacent high-
lands in Guatemala, 200-1000 m elevation.
Norops muralla KOHLER, MCCRANIE &
WILSON 1999, Amphibia-Reptilia 20: 285;
type locality: along trail to Cerro de Enmedio,
1500 m, Parque Nacional La Muralla,
Olancho, Honduras. SVL to 56 mm. Body
grayish brown, dewlap red. Parque Nacional
La Muralla, Honduras, 1440-1740 m elevation.
Norops ocelloscapularis KOHLER, MCCRANIE
& WILSON 2001a, Herpetologica 57: 248; type
locality: near Quebrada Grande off a trail to
Laguna del Cerro, l5°04.82'N, 88°55.45'W,
1200 m elevation, Copan, Honduras. SVL to
4 7 mm. Body grayish brown, dewlap orange.
Known only from vicinity of type locality,
1150-1370 m elevation.
Norops pachypus (COPE 1876), J . Acad. Nat.
Sci. Philadelphia (2) 8: 122; type locality:
Slopes of Pico Blanco, Costa Rica. SVL to 50
mm. Body grayish brown, dewlap red-orange.
Fig. 241. Norops laeviventris (male, near Fig. 245. Norops loveridgei (south-slope of
Zapote, Costa Rica). Photo: G. Kohler Cerro Bufalo, Atlantida, Honduras).
Photo: J . R. Mccranie

Fig. 242. Norops lemurinus (male, Isla Fig. 246. Norops macrophallus (near Guanaga-
Bastimentos, Panama). Photo: G. Kohler sapa, Escuintla, Guatemala). Photo: G. Kohler

Fig. 243. Norops limifrons (Los Algarrobos, Fig. 247. Norops magnaphallus (male).
Chiriqui, Panama). Photo: G. Kohler Photo: S. Poe

Fig. 244. Norops lionotus (near Pueblo Wiso, Fig. 248. Norops matudai (male, Chiapas,
Jinotega, Nicaragua). Photo: G. Kohler Mexico). Photo: A. Ramirez V.
109
 lguanidae
Highlands of Costa Rica and Volcan Chiriqui,
Panama, 1370-1560 m elevation.
Norops parvicirculatus (ALVAREZ DEL TORO &
SMITH 1956), Herpetologica 12: 5; type local-
ity: El Suspiro, 1200 m, Chiapas, Mexico. SVL
to 50 mm. Body grayish brown, dewlap
yellowish orange with a red central blotch.
Central-eastern Chiapas, Mexico, 500-1200 m
elevation.
Norops pentaprion (COPE 1862b), Proc. Acad.
Nat. Sci. Philadelphia 14: 178; type locality:
Truando River, Colombia. SVL to 79 mm.
Body grayish brown, dewlap red. Along the
Caribbean versant of Central America from
Tamaulipas, Mexico, to Panama, including
Yucatan Peninsula; also on Pacific versant in
Costa Rica and Panama, sea level to 900 m
elevation (to 1780 m in Veracruz, Mexico,
according to VOGT et al. 1997). Two subspe-
cies are recognized: Norops p. pentaprion
(Caribbean versant of Central America from
eastern Honduras to Panama) and Norops p.
beckeri (BOULENGER 1881) (Caribbean ver-
sant from Tamaulipas, Mexico, to Honduras;
including Yucatan Peninsula).
Fig. 249. Norops ocelloscapularis (holotype).
Photo: G. Kohler
Fig. 250. Norops pachypus (Cerro Picacho,
Chiriqui, Panama). Photo: G. Kohler
110
Norops petersii (BocOURT 1873), Miss. Sci.
Mex., Rept.: 79; type locality: Alta Verapaz,
Guatemala. SVL to 102 mm. Body greenish,
dewlap purple with pale yellow margin and
pale yellow gorgetal scales. San Luis Potosi,
Mexico, to Honduras, 200-2130 m elevation.
Norops pijolensis MCCRANIE, WILSON &
WILLIAMS 1993b, J. Herpetol. 27: 393; type
locality: East slope of Pico Pijol, 2050 m,
15°10'N, 87°33'W, Montana de Pijol north-
west of Tegucigalpa, Yoro, Honduras, 1860-
2050 m elevation. SVL to 60 mm. Body gray-
ish brown, dewlap rose with pink scales. Only
known from the type locality. Note that the
species name has been emended from the
original spelling ''pijolense" to "pijolensis" to
correspond to the masculine gender of the
genus name Norops (following H. SMITH &
LARsEN 1974a).
Norops poecilopus (COPE 1862b), Proc. Acad.
Nat. Sci. Philadelphia 14: 179; type locality:
near Cartagena and on Rio Truando,
Colombia. SVL to 68 mm. Body grayish
brown, dewlap yellow to yellowish orange.
Panama east of the Canal Zone and
Colombia, sea level to 1000 m elevation.
Norops polylepis (PETERS 1873), Monats.
Akad. Wiss. Berlin 1873: 738; type locality:
Chiriqui, Panama. SVL to 57 mm. Body gray-
ish brown, dewlap orange. Southwestern
Costa Rica and Panama, sea level to 1500 m
elevation.
Norops pseudokemptoni (KOHLER, PONCE,
SUNYER & BATISTA 2007), Herpetologica 63
(3): 380; type locality: "La Nevera, 8°29'45"N,
81°46'35"W, 1600 m elevation, Serrania de
Tabasara, Comarca Ngobe Bugle, Distrito de
Nole Dilima, Corregimiento de Jadeberi,
Panama". SVL to 54 mm. Body grayish
brown, dewlap brick red grading into orange
anteriorly with white scales, anterodorsal
corner of dewlap cream color. Only known
from type locality.
Norops pseudopachypus (KOHLER, PONCE,
SUNYER & BATISTA 2007), Herpetologica 63
(3): 383; type locality: "La Nevera, 8°29'45"N,
81°46'35"W, 1600 m elevation, Serrania de
Tabasara, Comarca Ngobe Bugle, Distrito de
Nole Duima, Corregimiento de Jadeberi,
Panama". SVL to 48 mm. Body grayish
brown, dewlap uniform orange yellow.
Serrania de Tabasara, Panama.
Norops purpurgularis MCCRANIE, CRUZ &
HOLM 1993a, J. Herpetol. 27: 386; type local-
ity: 2.5 km airline NNE La Fortuna, 1690 m,
15 26'N, 87 18'W, Cordillera Nombre de Dios,
Yoro, Honduras. SVL to 60 mm. Body grayish
Fig. 251. Norops pentaprion (Bartola, Rio San Fig. 255. Norops polylepis (Rio Sereno,
Juan, Nicaragua). Photo: G. Kohler Chiriqui, Panama). Photo: G. Kohler

Fig. 252. Norops petersii (Chiapas, Mexico). Fig. 256. Norops pseudokemptoni (La Nevera,
Photo: G. Kohler Panama). Photo: G. Kohler

Fig. 253. Norops pijolensis (east-slope of Pico Fig. 257. Norops pseudopachypus (La Nevera,
Pijol, Yoro, Honduras). Photo: J. R. McCranie Panama). Photo: G. Kohler

Fig. 254. Norops poecilopus (Nusagandi, Comar-

ca de San Blas, Panama). Photo: G. Kohler
111
 lguanidae
brown, dewlap pink. Mountainous regions in
the Dptos. de Yoro and Atlantida, Honduras,
1690-2040 m elevation.
Norops pygmaeus (ALVAREZ DEL TORO & SMITH
1956), Herpetologica 12: 7; type locality: El
Ocote, Chiapas, Mexico, 600 m . SVL to 35
mm. Body grayish brown, dewlap pale red.
Region of El Ocote, Chiapas, and south-
eastern Oaxaca, Mexico.
Norops quaggulus (COPE 1885), Proc. Amer.
Phil. Soc. 22: 391; type locality: Rio San Juan,
Nicaragua. Body grayish brown, dewlap red-
orange with yellow margin. On the Caribbean
versant from eastern Honduras through
Nicaragua to extreme northern Costa Rica,
sea level to 1350 m elevation.
Norops roatanensis KOHLER & MCCRANIE
2001, Senckenbergiana biol. 81: 240; type
locality: between West End Point and Flowers
Bay, l6°17.98'N, 86°34.82'W, 30 m elevation,
Isla de Roatan, Islas de la Bahia, Honduras.
SVL to 63 mm. Body grayish brown, dewlap
pink red with suffusion of black pigment cen-
trally and with white scales. Endemic to Isla
de Roatan, Islas de la Bahia, Honduras.
Norops rodriguezii (BOCOURT 1873), Miss. Sci.
Mex., Rept.: 62; type locality: Panzos,
Guatemala. SVL to 49 mm. Body grayish
brown, dewlap orange with darker. basal
area. Isthmus of Tehuantepec, Mexico, to
western Honduras, sea level to 2000 m eleva-
tion.
Norops rubribarbaris KOHLER, MCCRANIE &
WILSON 1999, Amphibia-Reptilia 20: 280;
type locality: 4 km S of San Luis de los
Planes, 1700 m elevation, N slope of Montana
de Santa Barbara, Santa Barbara, Honduras.
SVL to 48 mm. Body grayish brown, dewlap
red. Only known from the type locality.
Fig. 259. Norops quaggulus (Cerro El Toro,
Atlantico Norte, Nicaragua). Photo: G. Kohler
112
Norops sagrei (DUMERIL & BIBRON 1837), Erp.
Gen. 4: 149; type locality: Cuba. SVL to 57
mm. Body grayish brown, dewlap yellow-
orange to red-orange with yellow margin.
Southeastern Mexico to Honduras, sea level
to 200 m elevation. In Central America, the
subspecies Norops s. mayensis SMITH &
BURGER 1949 occurs.
Norops salvini (BOCOURT 1873), Miss. Sci.
Mex., Rept.: 75; type locality: ""Escuintla
(Republique du Guatemala)", type locality
erroneous according to KOHLER (2007). SVL to
64 mm. Body grayish brown, dewlap red with
white scales. Costa Rica and Panama, 1400-
1830 m elevation. Anolis vociferans MYERS
1971b is regarded to be a synonym of N. sal-
vini according to KOHLER (2007).
Norops sericeus (HALLOWELL 1856), Proc.
Acad. Nat. Sci. Philadelphia 8: 227; type local-
ity: Jalapa [= Xalapa), Veracruz, Mexico. SVL
to 52 mm. Body grayish brown, dewlap yel-
low-orange with central blue blotch. Isthmus
of Tehuantepec on the Pacific versant and
Tamaulipas, Mexico, on the Caribbean ver-
sant along both versants to Costa Rica, sea
level to 1500 m elevation.
Norops serranoi KOHLER 1999a, Salamandra
35: 39; type locality: forest in the vicinity of
butterfly farm of Dr. Francisco Serrano
(l3°49,46'N, 89°59,98'W), 225 m elevation,
Departamento Ahuachapan, El Salvador.
SVL to 85 mm. Body grayish brown, dewlap
brick red with suffusion of black pigment cen-
trally and with white scales. Pacific versant
of northern Central America from Chiapas,
Mexico, to eastern El Salvador, sea level to
950 m elevation.
Norops sminthus (DUNN & EMLEN 1932),
Proc. Acad. Nat. Sci. Philadelphia 84: 26; type
locality: San Juancito, 6900 ft, Honduras.
SVL to 55 mm. Body grayish brown, dewlap
red with pale to dark brown scales. South-
central Honduras, 1840-2100 m elevation.
Norops townsendi (STEJNEGER 1900), Bull.
Mus. Comp. Zool. 36: 163; Isla del Coco, Costa
Rica. SVL to 49 mm. Body grayish brown
with a lateral stripe bordered above and
below by a black stripe; dewlap dark brown.
Isla del Coco, Costa Rica, sea level to 630 m
elevation.
Norops tropidogaster (HALLOWELL 1856),
Proc. Acad. Nat. Sci. Philadelphia 8: 224; type
locality: Colombia. SVL to 55 mm. ~ody gr~y­
ish brown, dewlap red-orange with white
scales. Eastern Panama and northern South
America, sea level to 700 m elevation.
Fig. 260. Norops roatanensis (Isla de Roatan, Fig. 264. Norops sericeus (Rancho Alegre,
Honduras). Photo: G. Kohler Chiapas, Mexico). Photo: G. Kohler

Fig. 261. Norops rodriguezii (Quirigua, Fig. 265. Norops serranoi (Los Tarrales,
Guatemala). Photo: G. Kohler Suchitepequez, Guatemala). Photo: G. Kohler

Fig. 262. Norops sagrei (male, Hopkins, Fig. 266. Norops sminthus (La Tigra, Francisco
Belize). Photo: G. Kohler Morazan, Honduras). Photo: M. Lundberg

.!.lo.:
Fig. 263. Norops salvini (Jurutungo, Chiriqui, Fig. 267 Norops tropidogaster (Metropolitan
Panama). Photo: G. Kohler National Park, Panama). Photo: G. Kohler

113
 Iguanidae
Norops tropidolepis (BOULENGER 1885), Cat.
Lizards Brit. Mus. 2nd ed. 2: 53; type locality:
Volcan Irazu, Costa Rica. SVL to 59 mm.
Body grayish brown, dewlap purplish red to
pink. Volcan Irazu and Monteverde, Costa
Rica, 1220-2500 m elevation.
Norops tropidonotus (PETERS 1863b), Monats.
Akad. Wiss. Berlin 1863: 135; type locality:
Huanusco, Veracruz, Mexico. SVL to 60 mm.
Body grayish brown, dewlap dark orange
with yellow to yellowish orange margin and a
central dark streak. Southern Mexico to
northern Nicaragua, 90-1740 m elevation.
Norops uniformis (COPE 1885), Proc. Amer.
Phil. Soc. 22: 392; type locality: Yucatan,
Mexico. SVL to 40 mm. Body grayish brown,
dewlap rose with purple spots. Southern
Mexico to western Honduras, sea level to
1300 m elevation.
Norops utilensis KOHLER 1996a, Sencken-
bergiana biol. 75: 24; type locality: 2 km NNE
of the town of Utila, Isla de Utila, Islas de la
Bahia, Honduras. SVL to 59 mm. Body gray-
Fig. 268. Norops tropidolepis (male, Monte-
verde, Costa Rica). Photo: G. Kohler
Fig. 269. Norops tropidonotus (male; Rancho
Alegre, Chiapas, Mexico). Photo: G. Kohler
114
ish brown, dewlap red with white scales. Isla
de Utila, Honduras.
Norops villai (FITCH & HENDERSON 1976),
Milwaukee Pub. Mus. Contrib. Biol. Geol. 9: 1;
type locality: Great Corn Island, Nicaragua.
SVL to 55 mm. Body grayish brown, dewlap
chocolate brown. Great Corn Island,
Nicaragua.
Norops vittigerus (COPE 1862b), Proc. Acad.
Nat. Sci. Philadelphia 14: 179; type locality:
Truando region, Colombia. SVL to 72 mm.
Body grayish brown, dewlap red-orange with
a dark blotch. Panama and northern South
America, sea level to 900 m elevation.
Norops wampuensis MCCRANIE & KOHLER
2001, Senckenbergiana biol., Frankfurt a. M.,
81 (1/2): 228; type locality: confluence of Rios
Aner and Wampu, 15°03'N, 85°07'W, 110 m
elevation, Olancho, Honduras. SVL to 51 mm.
Body grayish brown, dewlap orange-red with
yellow margin. Known only from the type
locality, 95-110 m elevation.
Norops wermuthi KOHLER & OBERMEIER 1998,
Senckenbergiana biol. 77: 129; type locality:
road from Matagalpa to Jinotega at km 146
(13°01.995'N, 85°55.848'W), 1400 m elevati-
on, Departamento Jinotega, Nicaragua. SVL
to 54 mm. Body grayish brown, dewlap red
with pale to dark brown scales. Northern
Nicaragua, 1230-1500 m elevation.
Norops woodi (DUNN 1940a), Proc. Acad. Nat.
Sci. Philadelphia 92: 110; type locality: El
Volcan, Chiriqui, Panama. SVL to 100 mm.
Body grayish brown, dewlap dark-orange,
brownish olive or black (capable of color
change). Puntarenas Province, Costa Rica, to
Chiriqui, Panama, 1150-2500 m elevation.
Anolis attenuatus TAYLOR 1956 is regarded to
be a synonym of N. woodi according to SAVAGE
(2002).
Norops yoroensis MCCRANIE, NICHOLSON &
KOHLER 2002, Amphibia-Reptilia 22: 466;
type locality: 2.5 airline km NNE La Fortuna,
1600 m elevation, 15°26'N, 87°l8'W,
Cordillera Nombre de Dios, Yoro, Honduras.
SVL to 47 mm. Body grayish brown, dewlap
unicolor orange. Northwestern Honduras
(Yoro), 1180-1600 m elevation.
Norops zeus KOHLER & MCCRANIE 2001,
Senckenbergiana biol. 81: 236; type locality:
Liberia, Parque Nacional Pico Bonito, 90 m
elevation, Atlantida, Honduras. SVL to 44
mm. Body grayish brown, dewlap unicolor
dirty white. Caribbean versant of the
Cordillera Nombre de Dios in northern
Honduras, 90-900 m elevation.
Fig. 270. Norops tropidonotus (female; Rancho Fig. 274. Norops uittigerus (Cordoba,
Alegre, Chiapas, Mexico). Photo: G. Kohler Colombia). Photo: M. Lundberg

Fig. 271. Norops uniformis (Cerro San Gil, Fig. 275. Norops wermuthi (Cerro El Toro,
Guatemala). Photo: G. Kohler Atlantico Norte, Nicaragua). Photo: G. Kohler

Fig. 272. Norops utilensis (Isla de Utila, Fig. 276. Norops woodi (male, Rio Chevo,
Honduras). Photo: G. Kohler Chiriqui, Panama). Photo: J. Sunyer

Fig. 273. Norops woodi (juvenile, 11 km N San Fig. 277. Norops woodi (female, Rio Chevo,
Isidro del General, Costa Rica). Photo: G. Kohler Chiriqui, Panama). Photo: J. Sunyer
115
 lguanidae

• Norops barkeri

J
Norops capita
• Ano/is al/isoni
• Ctenonotus cristatellus Norops biporcatus

Dactyloa insignis Dactyloa frenata

• Dactyloa microtus O Norops gruuo
O Dactyloa latifrons • Norops fortunensis
D Dactyloa chloris o Norops kemptoni
• Dactyloa chocorum • Norops pseudokemptoni
,.. Dactyloa casildae ,.. Norops altae
• Dactyloa kunaya/ae • Norops auratus

Norops Norops

• N. a/varezdeltoroi
o N. campbelli • N. crassu/us
• N. hobartsmithi • N. haguei
"''...-- ~-, ,.. N. cobanensis '"""'...r'--.l ,.. N. anisolepis
• N. johnmeyeri • N. amplisquamosus
D. N. pijolensis D. N. rubribarbaris
\/ N. purpurgularis D N. parvicirculatus
D N. loveridgei ON. matudai
N. sagrei N. rodriguezii

Norops
o N. yoroensis
• N. cuprinus
,.. N. vil/ai
• N. do/lfusianus

N. lemurinus
• N. serranoi
• N. vittigerus

116
 lguanidae

Norops Norops

O N. cryptolimifrons
• N. apletophallus
 Iguanidae

Anolis allisoni Ctenonotus cristatellus Dactyloa casildae (male)

Dactyloa casildae (female) Dactyloa chocorum (male) Dactyloa insignis (female)

Dactyloa kunayalae (male) Dactyloa microtus (male) Norops altae

Norops aluarezdeltoroi Norops amplisquamosus Norops anisolepis

Fig. 278 - 349. Alloles with extended dewlaps. Because in most species females have no or very
small dewlaps (an exception in this respect is Norops johnmeyeri as well as most Dactyloa spe-
cies) only males are shown.
118
 Iguanidae

Norops apletophallus Norops aquaticus Norops auratus

Norops barkeri Norops bicaorum Norops biporcatus

1
....
~

Norops capita Norops carpenteri Norops cobanensis

Norops compressicauda Norops crassulus Norops cryptolimifrons

119
 lguanidae

Norops cupreus Norops datzorum Norops dollfusianus

Norops fortunensis Norops fungosus Norops fuscoauratus

Norops gruuo Norops heteropholidotus Norops humilis (central

Panama)

Norops johnmeyeri (male) Norops johnmeyeri (female) Norops kemptoni

120
 lguanidae

Norops laeviventris Norops lemurinus Norops lemurinus

(Isla Bastimentos, Panama) (Peten, Guatemala)

Norops limifrons Norops limifrons Norops lionotus

(Monteverde, Costa Rica) (Los Algarrobos, Panama)

Norops macrophallus Norops magnaphallus Norops pachypus

Norops pentaprion Norops pentaprion Norops petersii

(David, Panama) (Bartola, Nicaragua)

121
 Iguanidae

Norops poecilopus Norops polylepis (near Piedras Norops polylepis (near

Blancas, Costa Rica) Palmar Sur, Costa Rica)

Norops polylepis (near Balzar, Norops polylepis (Manuel Norops pseudokemptoni

Costa Rica) Antonio, Costa Rica)

Norops pseudopachypus Norops quaggulus Norops rodriguezii

(Guatemala)

Norops rodriguezii Norops sagrei Norops saluini

(Guatemala)

122
 Iguanidae

Norops sericeus Norops serranoi Norops tropidogaster

Norops tropidolepis Norops tropidonotus Norops uniformis (Belize)

Norops uniformis Norops utilensis Norops vittigerus (female)

(Guatemala)

Norops wermuthi Norops woodi Norops zeus

123
 lguanidae

Key to anoles (genera Anolis, Dacty-

loa, Ctenonotus, Norops)
Elaborating an identification key to all Central
American anoles is a difficult task, partly
because some species are known from a single
specimen or a few individuals. In these cases,
the variation of supposely diagnostic character- a. Norops tropidonotus
istics is insufficiently documented. Some species
are extremely similar in their external morpho-
logy and coloration, but can be distinguished
readily by their very different hemipenial Fig. 351.
morphology, which, in some cases, is interpreted
as evidence for only distant relationships of Scalation anteri-
these cryptic species (e.g., Norops cupreus and or and posterior
N macrophallus; KOHLER & KREUTZ 1999). The of tympanum.
following information (including SVL) refers to b. Norops humilis
adults only. Juveniles of many anole species are
extremely difficult to identify. Also, there is con-
siderable intraspecific (particularly sexual)
variation in some species. Therefore, a few spe-
cies appear more than once in the following key.
...
t"t
N
Statements regarding coloration (of body and
dewlap) refer to specimens in life; dewlap color
refers to adult males, if not otherwise specified a. Norops biporcatus b. Norops petersii
(females of most anole species have no or only a
rudimentary dewlap). Subdigital lamellae
counts are given for phalanges ii to iv of the
fourth toe (not including distal phalanx), if not
otherwise specified.

1 a Mostly green in life (capable of rapid color c. Norops limifrons d. Norops utilensis
change to olive brown); in adult males head
conspicuously large and elongate; dewlap
pink; 2-3 scales between nasal and rostral;
SVL 65-87 mm; short-legged (4th toe of
adpressed hindlimb reaches usually to ear
opening, in some individuals to point be-
tween ear opening and eye); dewlap pink f. Dactyloa /atifrons
............ .............................. Anolis allisoni
b Combination of characters different from Fig. 352. Ventral scales in anoles:
that above ............................................. 2 a. imbricate, strongly keeled, mucronate
b. imbricate, weakly keeled, non-mucronate
c. smooth, non-imbricate
d. granular, not keeled
e. imbricate, smooth
f. slightly imbricate, smooth

Fig. 353. Norops wermuthi.

a (left). Mid-dorsum; note several rows of enlarged
Fig. 350. Norops tropidonotus; note tube-like dorsal scales.
axillary pocket (arrow). Photo: G. Kohler b (right). Heterogeneous lateral body scales.
124

2 a A deep tube-like axillary pocket present
(Fig. 350) .................................................... 3
b No deep tube-like axillary pocket present,
at most, a shallow axillary depression .... 8
3 a Scales anterior to ear opening flat and kee-
led and about twice as large as scales
posterior to ear opening (the latter are
usually granular; Fig. 351a) ................. .... 4
b Scales anterior to ear opening about same
size as scales posterior to ear opening (all
granular; Fig. 351b) ................................ .. 6
4 a Dewlap uniform rose to purple; tail in
males conspicuously compressed ......... .
.......................... Norops compressicauda
b Dewlap orange red with yellow margin
(with or without a central dark streak); tail
not conspicuously compressed ... ............... 5
5 a Adult males 50-60 mm SVL, adult females
45-55 mm SVL; dewlap orange red with a
yellow margin and a distinct central dark
streak .......... ........ Norops tropidonotus
b Adult males 40-50 mm SVL, adult females
42-51 mm SVL; dewlap orange red with a
yellow margin, but without a central dark
streak ..................... Norops wampuensis
6 a Usually 22-32 dorsal scales between levels
of axilla and groin, rarely up to 36; flank
usually with 1-3 pale vertical lines (lines
can be broken); dewlap rose with purple
spots .. ... ...................... Norops uniformis
b Usually 30-39 dorsal scales between levels
of axilla and groin, rarely as few as 23;
flank without pale vertical lines; dewlap
red orange with or without yellow margin
······ ····················· ·· ························· ··· ·· ······ 7
7 a Usually two large elongate scales in the
anterior supraciliary region (Fig. 354a);
postaxillary pocket relatively wide and
shallow; hemipenis relatively large with
well-developed elongate lobes and with a
strongly calyculate surface on both the
truncus and the lobes; maximum SVL 46.2
mm in males and 50.0 mm in females
........... .... .. ..... Norops humilis (in part)
b Usually three large elongate scales in the
anterior supraciliary region (Fig. 354b);
postaxillary pocket usually narrow, tube-
like and deep; hemipenis relatively small
with short and stout lobes and without a
strongly calyculate surface on either the
truncus or the lobes; maximum SVL 37.0
mm in males and 41.0 mm in females
..... ..... ..... ............ ...... Norops quaggulus
8 a Tail distinctly compressed laterally and
with a crest of enlarged scales (especially
visible in males) ....... ............................... 9
Iguanidae
b Tail not distinctly compressed laterally,
although a slightly carinate crest can be
present .................................................... 12
9 a Ventrals smooth; tail with a crest of supra-
caudal scales of unequal size (sequences of
smaller scales that increase in size to the
following lateral whorl; Fig. 357a); dewlap
greenish yellow with orange margin ......... .
.... .. ....... .. ........ Ctenonotus cristatellus
b Ventrals faintly to distinctly keeled; supra-
caudal scales of equal size (Fig. 357b-d);
dewlap dark orange, orange-red or orange
with pink streaks ........ .. .. ...................... 10
lOa 2-3 scales between supraorbital semi-
circles; supraoculars smooth .... ............... .
........... ...... ......................... Norops barkeri
b Either 0-1 or 4-6 scales between supra-
orbital semicircles; supraoculars keeled 11
Ila 0-1 scales between supraorbital semi-
circles; males with enlarged postanal
scales ............... ........ ........ Norops sagrei
b 4-6 scales between supraorbital semi-
circles; males without enlarged postanal
scales .......... Norops loveridgei (in part)
12a Head conspicuously broad and stout, snout
rather blunt in lateral aspect; long-legged
(4tn toe of adpressed hindlimb reaches at
le'.'lst to anterior border of eye); usually
with a pale band across chin (most distinct
in juveniles; Fig. 356);dorsal scales smooth
juxtaposed, mostly pentagonal or hexa~
gonal; tip of tail enlarged, not tapering; no
enlarged postanal scales in males; SVL 80-
96 mm; dewlap uniform greenish yellow
....................................... Norops capito
Fig. 354. Supraciliary region of a. Norops
humilis and b. N. quaggulus.
125
 lguanidae

b Combination of characters different from

that above ........................ ..... .... ...... .... 13
13a Short-legged (4th toe of adpressed
hindlimb reaches usually to ear opening, in
some individuals to point between ear
opening and eye); large adult size (SVL to
100 mm); 9-11 supralabials to level below
center of eye; males without enlarged
Fig. 355. Enlarged post- Fig. 356. Norops postanal scales; dorsals small, not much
larger than flank scales ................... ... .. 14
anal scales in Norops capita: ventral surface
tropidogaster. of head. b Combination of characters different from
that above ..... .. ....... .... .. ..... . ... ... .......... 15
supracaudals 14a Dorsal head scales faintly keeled; ventrals
only slighty keeled with rounded posterior
margins (Fig. 352b), 62-78 ventrals be-
tween levels of axilla and groin; 120-140
scales around midbody; throat region with
dark streaks; dewlap purple with pale
yellow margin and pale yellow gorgetal
scales ................................. Norops petersi
a. Ctenonotus cristatellus b. Norops sagrei
b Dorsal head scales strongly keeled;
ventrals strongly keeled and mucronate
(Fig. 352a), 40-52 between levels of axilla
and groin; 100-112 scales around midbody;
no dark streaks in throat region; dewlap
blue with dark orange margin ..............
.......... .......................... Norops biporcatus
c. Norops capita d. Norops polylepis
15a Iris metallic blue in life; large adult size
Fig. 35 7. Lateral scalation of tail (scales of (SVL to 100 mm); body often with dark
whorls shaded). green or brown bands with numerous
small dots; 2-3 scales between supraorbital
semicircles; 1 scale between suboculars
and supralabials; males with enlarged
postanal scales; dorsal ground color dull
brick red or dark olive in life, pale yellow to
tan with numerous small punctations in
preservative; dewlap large, dark-orange,
brownish olive or black (capable of color
change) ....................... .. .... Norops woodi
b Combination of characters different from
that above ................ . .... .. ........ .. .. ... ... 16
16a Adult size more than 100 mm ................ 17
b Adult size less than 90 mm ................... 23
a. Norops utilensis b. Norops pentaprion 17a Ventrals distinctly keeled; 1-2 scales be-
Fig. 358. Ventral surfaces of foot. tween suboculars and supralabials; males
without enlarged postanal scales ... ........ .. .
... .................. Norops loveridgei (in part)
b Ventrals smooth; suboculars and suprala-
bials in contact or separated by 1 scale row;
males with enlarged postanal scales ..... 18
18a Short-legged (4th toe of adpressed hind-
limb reaches usually to ear opening, in
some individuals to point between ear open-
a. Dactyloa latifrons b. Dactyloa frenata ing and eye); brown and cream I white in
coloration, often with a greenish cast in life
Fig. 359. Eye region (enlarged superciliaries .......................... ....... ................................. 19
orange).

126
 Iguanidae

b Long-legged (4th toe of adpressed

hindlimb reaches at least to center of eye;
mostly green in life ...... ......... ..... .. ... ...... 20
19a Sublabials distinctly enlarged (Fig. 360a);
interparietal plate not distinct, no parietal
eye; 4th toe with fewer than 50 subdigital
lamellae (from base of digit to claw)
......... ... ........................ Dactyloa microtus
b Sublabials not differeciated (Fig. 360b); a
well developed interparietal plate present, a. Dactyloa microtus b. Dactyloa insignis
distinctly larger than liadjacent scales,
parietal eye present; 4t toe with more Fig. 360. Chin region (sublabial scales orange).
than 50 subdigital lamellae (from base of Drawings by L. Czupalla
digit to claw) .......... ...... Dactyloa insignis
20a All of upper margin of eye with enlarged
superciliaries forming a stiff ridge (Fig.
359a) ..... .. ... ... .......... Dactyloa latifrons
b Enlarged superciliaries restricted to anter-
ior half of eye, followed by small granular
scales posteriorly, not forming a stiff ridge
(Fig. 359b) ............................................... 21
2la Distal phalanx (including claw) longer
than phalanges ii and iii combined; basal
portion of male dewlap orange with large
white basal portion and multiple rows of
yellow-green or blue scales; female dewlap
pale bluish-green with orange margin ...... .
.... . . . . . .......... ... ......... Dactyloa kunayalae a. Norops yoroensis
b Distal phalanx (including claw) shorter
than phalanges ii and iii combined; male
and female dewlap coloration not as above
.............................................................. .... 22
22a Dorsum with 4-6 oblique brown bands (not
formed of adjacent ocelli) interspersed by
blue-outlined yellow patches; dewlap dirty
cream with broad yellow scale rows irregu-
larly interspersed with smaller emerald
green scales; nasal in contact with rostral
or separated by one scale ......... .. .............. .
... ... . .. . .. . ...................... Dactyloa casildae
b Dorsum with several dark green ocelli b. Norops aquaticus
(most outlined with yellow) that are close-
ly arranged and form dark oblique bands;
dewlap dull, cream-colored (occasionally Fig. 361. Dorsal view of head.
with black shading); nasal separated from
rostral by one or two scales .... ................. .
............ ......... .................. Dactyloa frenata
23a Interparietal scale relatively large, surro-
unded by minute scales, separated from
supraocular semicircles by 6-9 scales (Fig.
361b); body yellowish green, with broad
transverse brown bands and a pale gree-
nish yellow lateral longitudinal stripe;
dewlap red-orange with irregular yellow
stripes ............... .......... Norops aquaticus Norops polylepis Norops wermuthi
b Interparietal scale large or small, but sur- Fig. 362. Chin region (postmental scales
rounded by scales of moderate size, 0-9 sea- orange).

127
 Iguanidae

les between interparietal and supraorbital

semicircles (Fig. 361a); body pattern diffe-
rent than above; dewlap color different
than above ............................................... 24
24a Bright green in life ................................. 25
b Mostly gray or brown (at most with a olive
green cast) in life ........ .............. ............. 26
25a Veotrals distinctly keeled; short-legged
(4th toe of adpressed hindlimb does not
reach to eye); SVL to 60 mm .. ... ................ . a. Norops cristifer
. . . . . . . . . . . . . . . . . . . . . . . . . . . . ...... Dactyloa chloris
b Ventrals smooth; long-legged (4th toe of
adpressed hindlimb reaches at least to
anterior border of eye); SVL to 73 mm ...... .
.. .... . . . . . . . . .. ................ Dactyloa chocorum
26a Eyes blue in life; long-legged (4th toe of
adpressed hindlimb reaches beyond anteri-
or border of eye); distal phalanx of fingers
and toes conspicuously long, about the
same length as combined length of penulti-
mate and antepenultimate phalanges; dor-
sal ground color olive brown with a gree- b. Norops pentaprion
nish cast; lateral body with 3-5 oblique or
vertical pale lines ........................................ . Abb. 363. Mid-dorsal scales in Norops cristifer
... ............ .......... Norops alvarezdeltoroi and N. pentaprion. Photos: G. Kohler
b Combination of characters different than
that above, particularly, distal phalanx of
fingers and toes not conspicuously long,
much shorter than combined length of b Proximal subdigital scales of toes differen-
penultimate and antepenultimate phalan- tiated as slightly broadened lamellae (Fig
ges ......................... ............................ 27 358b); scales along dorsal surface of tail
carinate and distinctly enlarged; SVL to 79
27a Ventrals granular or smooth and subimbri- mm ....................................... ........... ........ 31
cate; flank scales homogeneous; very short- 31a Adult males with a well-developed serra-
legged (4th toe of adpressed hindlimb rea- ted mid-dorsal crest (Fig. 363a); SVL to 88
ches not much further than shoulder); 5 or mm ... ......... .......... ... ..... Norops cristifer
fewer horizontal rows of loreals; supraocu-
lars smooth; lining of throat black b A double-row of slightly enlarged mid-dor-
(pentaprion group) .... .......................... 28 sal scales, but no serrated crest present
(Fig. 363b); SVL to 79 mm ..................... .
b Combination of characters different from ................................... Norops pentaprion
that above .... ........................................... 32
32a Most dorsal head and body scales strongly
28a A pair of small bony parietal protuber- keeled; lateral body scales homogeneous;
ances posterior and lateral to interparietal ventral scales strongly keeled and
plate; scales along dorsal surface of tail mucronate; 7-12 distinctly enlarged dorsal
carinate, but not enlarged ......................... . scale rows; males without enlarged post-
....................... ........... .... . Norops fungosus anal scales; relatively long-legged (4tfi toe
b No bony parietal protuberances; scales of adpressed hindlimb reaches at least to
along dorsal surface of tail carinate (en- posterior border of eye); dewlap red orange
larged or not) ......................................... 29 with yellow margin or dewlap uniform red
orange without yellow margin (populations
29a Ventrals keeled; a dark interorbital bar in eastern Panama); maximum SVL 46.2
present; males with enlarged postanal mm in males and 50.0 mm in females
scales ........... ..................... Norops salvini ....... .... .... ....... Norops humilis (in part)
b Ventrals smooth; no dark interorbital bar; b Combination of characters different from
males without enlarged postanal scales 30 that above ........................................... . 33
30a Proximal subdigital scales of toes granular 33a Most dorsal head scales distinctly multica-
(Fig. 358a); scales along dorsal surface of rinate; lateral body scales homogeneous;
tail carinate, but not enlarged; SVL to 59 ventral scales strongly keeled; no distinct-
mm ... ............. ... ......... Norops utilensis ly enlarged dorsal scale rows; relatively
128

long-legged (4th toe of adpressed hindlimb
reaching to tympanum; maximum SVL 49
mm; body with a greenish cast in life ........ .
..................... ............... Norops datzorum
b Combination of characters different from
that above ............................ .. .............. 34
34a 2-9, 5 or more in most populations, scales
between supraorbital semicircles; first and
second canthals usually with more than
one keel; 1-2 scales between suboculars
and supralabials; long-legged (4Th toe of
adpressed hindlimb reaches to a point
beyond eye) (pachypus complex) ........ 35
b Combination of characters different from
that above ................. ............. ............. 38
35a Dewlap bicolored: red or purple with a yel-
lowish-orange central blotch or with a yel-
low margin ................ .. Norops pachypus
b Dewlap uniform yellow, purplish red, or
pink························································· 36
36a Usually 7 or more scales between interpa-
rietal scale and supraorbital semicircles;
usually 16 or more scales between second
canthals; dewlap uniform yellow ... .... .
....................... .. Norops pseudopachypus
b Usually 6 or fewer scales between interpa-
rietal scale and supraorbital semicircles;
usually 15 or fewer scales between second
canthals; dewlap uniform purplish red or
brick red .................................................. 37
37a 4-6 scales between supraorbital semi-
circles; dewlap uniform yellow
................................. Norops tropidolepis
b 2-4 scales between supraorbital semi-
circles; dewlap uniform brick red
............................. Norops magnaphallus
38a Usually a conspicuous lyriform marking in
occipital region, can be fragmented (pre-
sent in about 80% of individuals); ventrals
strongly keeled and mucronate; relatively
long-legged (4th toe of adpressed hindlimb
reaches 8:t least to posterior border of eye);
males without enlarged postanal scales;
dewlap red, red-orange or brown-red
(lemurinus group) ................................ 39
b Combination of characters different from
that above ........ ...... ............................ 43
39a 10-20 dorsal scale rows irregularly en-
larged; flank scales usually heterogeneous
(enlarged scales scattered among smaller
scales); SVL 50-60 mm (males) or 55-72
mm (females); dewlap red-orange with a
dark blotch ...... ... ........ Norops vittigerus
b Only the two medial dorsal scales rows
slightly enlarged; flank scales homoge-
neous; dewlap without a dark blotch, al-
though often with black gorgetal scales 40
Iguanidae
40a Dewlap red to red-orange without suffu-
sion of black pigment, often with black gor-
getal scales; males, on the average, smaller
than females .............. . Norops lemurinus
b Dewlap red to pink-red, with suffusion of
black pigment centrally; gorgetal scales
white; males, on the average, larger than
females, or both sexes about the same size
...... .......................................................... 41
41a 5-8 horizontal rows of loreals; hemipenis
unilobate; Pacific versant of northern
Central America ........... Norops serranoi
b 8-10 horizontal rows of loreals; hemipenis
bilobate; endemic on Islas de la Bahia,
Honduras ........................... ................. 42
42a Dewlap orange red, with suffusion of black
pigment centrally; males average 65 mm
SVL (maximum 75 mm), females 64 mm
SVL (maximum 67 mm); endemic on Isla
de Utila ................. .. .. Norops bicaorum
b Dewlap pink red, with suffusion of black
pigment centrally; males average 55 mm
SVL (maximum 62 mm), females 57 mm
SVL (maximum 63 mm); endemic on Isla
de Roatan ................ Norops roatanensis
43a Lateral body scales heterogeneous, solitary
enlarged keeled or elevated scales (often
whitish) scattered among smaller (mostly
granular) laterals (Fig. 353b); males with
enlarged postanal scales; suboculars and
supralabials mostly in contact (crassulus
group) ..... ................................... ... ........ 44
b Combination of characters different from
that above, particularly, lateral body scales
homogeneous .......................................... 55
44a Enlarged dorsal scales abruptly different
in size from much smaller lateral body
scales; dewlap orange-yellow ................ .
... .......... ......... Norops amplisquamosus
b Enlarged dorsal scales grade into smaller
lateral body scales (Fig. 353a); dewlap color
different from that above ...................... 45
45a Dewlap dirty white, pale yellow or pale
gray; ratio tail length I SVL 1.9 or less .. 46
b Dewlap color different; ratio tail length I
SVL greater than 1.8 ........... ....... ......... 48
46a Dewlap extending to about level of axillae
................ .. ... ......... Norops laeviventris
b Dewlap extending well onto chest posteri-
or to level of axillae ........... .............. .... 47
47a Dewlap pale yellow with purple gorgetal
scales; 6-7 scales between second canthals
.............. ....... ................ Norops kreutzi
b Dewlap dirty white or pale gray with gor-
getal scales of same color; 7-10 scales be-
tween second canthals ..... Norops cusuco
129
 lguanidae
48a Ventrals strongly keeled; 8 or more rows of
distinctly enlarged dorsal scales ........... 49
b Ventrals smooth or only faintly keeled; 6-8
rows of distinctly enlarged dorsal scales
.............................................. ............... ..... 52
49a Ratio tail length I SVL 2.5 or greater; 4
scales between second canthals; 8 rows of
distinctly enlarged dorsal scales; about 29
ventrals in one head length ...... ... .. .
..... .... .. ................ Norops rubribarbaris
b Ratio tail length I SVL smaller than 2.4; 5-
9 scales between second canthals; 10-23
(mostly 12-15) rows of distinctly enlarged
dorsal scales; 13-27 ventrals in one head
length ........................ ... ................ ........... 50
50a Usually fewer than 27 dorsals in one head
length (17-33, mean 24); no nuchal crest
with enlarged scales ... Norops crassulus
b Usually more than 27 dorsals in one head
length (21-43, mean 30); a nuchal crest
with enlarged scales usually present .... 51
51a 1-2 scales between supraorbital semi-
circles; 22-26 lamellae beneath 4. toe; SVL
to 4 7 mm .................. Norops anisolepis
b 0-1 scales between supraorbital semi-
circles; 23-30 lamellae beneath 4. toe; SVL
to 53 mm ......................... Norops haguei
52a Ventrals perfectly smooth ...................... 53
b Ventrals faintly keeled ........................... 54
53a Medial dorsal scales uniform in size, with-
out interspersed small scales .................. .
............ ........... Norops heteropholidotus
b Small scales irregularly interspersed
among the enlarged medial dorsal scales
.................................... Norops muralla
54a Hemipenis with one asulcate-side process;
dewlap of adult males of moderate size
with relatively numerous gorgetal scales
..................................... Norops sminthus
b Hemipenis with two asulcate-side proces-
ses; dewlap of adult males small, with rela-
tively few gorgetal scales ............. .
... ....... .. ................ ..... Norops wermuthi
55a Ventrals perfectly smooth, flat and
distinctly imbricate (Fig. 352e); 2 medial
dorsal scale rows enlarged; canthus rostra-
lis distinct; supraoculars distinctly keeled
(schiedei group) ... ................................ 56
b Combination of characters different from
that above ..... ...... .... ....... .... ........ ....... 64
56a Dewlap of males orange-red with a large
central blue blotch; dewlap of females well
developed, yellow with a large central blue
blotch; SVL to 70 mm (males) or 65 mm
(females) ............... .. . Noropsjohnmeyeri
130
b Dewlap color different; females without a
well developed dewlap; SVL to 60 mm .. 57
57a Parietal ridges usually well developed, a
distinct parietal depression present; males
usually with slightly enlarged postanal
scales .............. .. .. ... ..... ... ........................ 58
b Parietal ridges absent to only slighty
developed, parietal depression absent or
only slightly developed; males usually
without enlarged postanal scales ........... 59
58a 6-8 scales between second canthals; snout
scales multicarinate; 52-62 dorsal scales
between levels of axilla and groin; 7-9
(rarely 6) horizontal rows of loreals;
dewlap purple .... Norops purpurgularis
b 10-11 scales between second canthals;
snout scales unicarinate; 43-54 dorsal
scales between levels of axilla and groin; 6-
7 horizontal rows of loreals; dewlap rose
with purple spot ..... ..... Norops pijolensis
59a Scales of supraorbital semicircles distinct-
ly smaller than scales between semicircles;
dewlap yellowish orange with a red central
blotch ................ Norops parvicirculatus
b Scales of supraorbital semicircles about
equal in size to, or larger than scales be-
tween semicircles; dewlap color different
...................................... ............................ 60
60a Suboculars and supralabials usually in
contact (lorilabial row incomplete); anteri-
or nasal usually fused with circumnasal 61
b Usually one complete scale row separating
suboculars and supralabials; anterior
nasal usually distinct ....................... .. .... 62
61a Scales comprising supraorbital semicircles
usually distinctly larger than scales bet-
ween semicircles (Fig. 364a); smooth or
only some faintly keeled midventral scales;
some keels usually oblique or oriented
transversely immediately to prefrontal
depression; scales in prefrontal depression
mostly smooth; circumorbital row usually
complete; hemipenis small and unilobate
.................................... Norops cobanensis
b Scales comprising supraorbital semicircles
about the same size as scales between
semicircles (Fig. 364b); most midventral
scales weakly, yet definitely keeled; keels
oriented nearly longitudinally immediate-
ly to prefrontal depression; scales in pre-
frontal depression strongly keeled; circu-
morbital row rudimentary or almost
absent; hemipenis large and bilobate ....... .
... ......... ......... .......... ..... Norops campbelli
62a Nasal and postrostral scales fused; 6 or
fewer medial rows of dorsal scales slightly

enlarged (distinctly larger than flank
scales); dewlap rose ................................. .
................................ Norops hobartsmithi
b Nasal and postrostral scales distinct; 4-20
medial rows of dorsal scales slightly en-
larged (distinctly larger than flank scales);
dewlap color not as above ... ............. ..... 63
63a Ventrals usually weakly to moderately
keeled; extensive black, diffuse coloration
on flanks common in males; dewlap red
......... ....................... ....... Norops cuprinus
b Ventrals smooth or only faintly keeled; no
black flank coloration in males; dewlap
pinkish red with yellow margin ................. .
....................................... Norops matudai
64a Ventrals smooth, or, if faintly keeled, then
with rounded posterior margins; anterior
nasal usually single ............................ .
(fuscoauratus group) .......................... 65
b Ventrals distinctly keeled, imbricate and
usually mucronate; anterior nasal usually
divided .............. .. .............................. .... 81
65a 8-10 rows of dorsal scales distinctly en-
larged and abruptly larger than much
smaller laterals; males without enlarged
a. Norops cobanensis
b. Norops campbelli
Fig. 364. Dorsal view of head.
Drawings by D. Dick and L. Czupalla
Iguanidae
postanal scales; dewlap pale red ..... ......... .
..................................... Norops pygmaeus
b 0-4 rows of dorsal scales slightly enlarged,
usually grading into small lateral scales;
males with enlarged postanal scales or not;
dewlap color different ............................. 66
66a Long-legged (4th toe of adpressed hindlimb
reaches at least to center of eye, usually
beyond eye) ............. ................................. 67
b Short-legged (4th toe of adpressed
hindlimb does not reach to eye, usually not
beyond ear opening) .... ...... ...... .. .. ........ 75
67a Ventrals slightly keeled and slightly imbri-
cate ......... ....... ..................... ................... 68
b Ventrals usually smooth and non-imbricate
...................... .... ..... .. ......... ..... ... ...... .... 70
68a An ocellated shoulder spot present ... ..... .
......................... Norops ocelloscapularis
b No ocellated shoulder spot ...................... 69
69a Dewlap yellow; suboculars and supralabi-
als usually in contact ............................ . .
......... .... .. ................. Norops dollfusianus
b Dewlap orange; usually 1 scale between
suboculars and supralabials ......... ........... .
. . . . . . . . . .. . . . . . . ................ ... Norops yoroensis
70a 2 elongate, overlapping superciliaries; SVL
to 57 mm; dewlap orange with a darker
basal portion or dirty white with an orange
basal portion ................. Norops polylepis
b 1 elongate superciliary; SVL to 48 mm;
dewlap color vaiable ...... ........................ 71
71a 4th toe of adpressed hindlimb reaches to
center of eye; dewlap orange with a darker
basal portion ............. Norops rodriguezii
b 4th toe of adpressed hindlimb reaches
beyond eye; dewlap color different ....... 72
72a Dewlap dirty white with a basal orange-
yellow spot or dewlap almost completely
orange-yellow ................. ...... ................... 73
b Dewlap uniformly dirty white, no basal
orange-yellow spot ................ Norops zeus
73a Dewlap large, larger than 150 mm2, almost
uniformily orange-yellow ..... ....... .
...... ........... ............. Norops apletophallus
b Dewlap small, smaller than 100 mm2, dirty
white with a basal orange-yellow spot .... 74
74a Hemipenis unilobate .... Norops limifrons
b Hemipenis bilobate ................................... .
.............. ............. Norops cryptolimifrons
75a Body greenish in life (in preservative yel-
low, with numerous small dark punctua-
tions, resulting in a fine speckled appear-
131
 ance); venter usually with a lichenous re-
ticulum; dewlap uniformly orange ........... .
... .... ............. ....... ........ Norops carpenteri
b Body in life gray or brown (in preservative,
with no fine speckled appearance); venter
immaculate; dewlap variable ................. 76
76a Total number ofloreal scales 75-88; a small
orange dewlap present in females (Fig.
365b); hemipenis bilobate ..... ..................... .
.... ............... .... .. . Norops pseudokemptoni
b Total number ofloreal scales 33-68; female
dewlap, if present, dirty white (Figs. 365a,c);
hemipenis unilobate or bilobate ............. 77
77a Tail with conspicuous dark and pale verti-
cal banding, pale bands almost white; male
dewlap more or less uniform orange;
posterior insertion of male dewlap at level
of axilla .. ............................ Norops gruuo
b Tail without conspicuous dark and pale
vertical banding, if present, pale bands are
cream to pale brown; male dewlap colorati-
on variable; posterior insertion of male
dewlap about one head length posterior to
level of axilla ... .................. ...... ............. ... 78
78a Ventrals smooth and non-imbricate ... .. . 79
b Ventrals faintly to moderately keeled and
usually slightly imbricate ... ......... .......... 80
79a A small dirty white dewlap present in
females ; posterior portion of male dewlap
greenish beige, anterior portion reddish
orange; 4-6 granular scales between enlar-
ged supraoculars and posterior (smaller)
superciliary; anterior superciliary at least
three times the length of posterior superci-
liary .. ... ...... ............. Norops fortunensis
b No dewlap in females ; male dewlap more
or less uniform reddish orange with a pur-
ple cast; 2-4 granular scales between
enlarged supraoculars and posterior
superciliary; anterior superciliary not
more than two times the length of posteri- Fig. 365. Female dewlaps in a. Norops kemptoni;
or superciliary ....... Norops fuscoauratus b. N pseudokemptoni and c. N fortunensis.
80a Male dewlap more or less uniform reddish Photos: G. Kohler
orange; hemipenis bilobate; ratio head
length/head width 1.5-1.8 (mean 1.65);
SVL to 51 mm ..... .. .... ... ...... Norops altae
point between shoulder and tympanum,
b Posterior portion of male dewlap rose pink, rarely beyond tympanum); tympanum very
anterior portion orange, a cream colored small (less than 1/2 the size of interpa-
basal area in some populations; hemipenis rietal plate); no enlarged postanal scales;
unilobate; ratio head length/head width dewlap yellowish orange with large blue
1. 7-2.0 (mean 1.80); SVL to 57 mm ............ . blotch .. ............................ Norops sericeus
............................. .. ...... Norops kemptoni b Combination of characters different from
Sia 8-10 medial dorsal scale rows slightly that above ...... ....... ..... ... ...... .. ............ 82
enlarged; anterior superciliary conspic-
uoQsly large and elongate; short legged 82a Usually 4 (rarely only 2) medial dorsal
(4th toe of adpressed hindlimb reaches to a scale rows slightly enlarged; 0-2 scales be-

132
 tween supraorbital semicircles; supraocu-
lars distinctly keeled; ventrals faintly to
distinctly keeled, but not mucronate.; males
without enlarged postanal scales; 4th toe of
adpressed hindlimb reaches to a point be-
tween shoulder and eye; dewlap of males
large (cupreus complex) ......... ............ 83
b Combination of characters different from
that above ................. .... ....... ..... .......... 85
83a Ventrals faintly keeled; dewlap uniformly
chocolate brown; endemic on Great Corn
Island, Nicaragua ............... Norops villai
b Ventrals distinctly keeled; dewlap color
different; distributed on mainland Central
America .................. ................. .... .... ....... 84
84a Surface of hemipenis covered with numer-
ous small spines (Fig. 366a); dewlap rose
with a orange-yellow basal blotch ......... .
.... .. .. ...................... Norops macrophallus
b Surface of hemipenis calyculate (Fig.
366b); dewlap brown or purple with darker
brown or orange brown basal blotch ..... .. ..
. . . . . . . . . .. . . . . . . . . .......... ......... Norops cupreus
85a Short legged (4th toe of adpressed
1
....
~

hindlimb reaches to a point between shoul-

der and tympanum, rarely beyond tym-
panum); dewlap blue; 0-1 scales between
supraorbital semicircles; dorsals large with
their keels arranged in longitudinal rows;
males with enlarged postanal scales
.......... .... ....... ...... ............. Norops auratus
b Combination of characters different from
that above .......................... ......... ......... 86
86a A conspicuous pale lateral stripe from neck
to at least mid-trunk present; 18-24 rows of
medial dorsal scales distinctly enlarged,
dorsals flattened, penta- or hexagonal; 1
scale between suboculars and supralabials;
2-4 scales between supraorbital semi-
circles; dewlap uniformly yellow or orange
(lionotus group) ................................... 87
b Combination of characters different from
that above .................... .......... ............. 88
87a 6-17 scales between second canthals ....... ..
...... ............... ............... Norops lionotus
b 20-22 scales between second canthals
......... ... ........ .... ........ Norops poecilopus
88a Males with strongly enlarged postanal
scales; no lateral stripe bordered above and
below by a black stripe along each side of
the body ............... Norops tropidogaster
b Males without enlarged postanal scales; a
lateral stripe bordered above and below by
a black stripe along each side of the body
........ .. ......................... Norops townsendi
Fig. 366. Hemipenis (sulcate view). Scale bar=
1 mm. Photo: G. Kohler
Further Reading
DUNN 1930b, BARBOUR 1932, 1934, H. SMITH &
KERSTER 1955, STUART 1955, ETHERIDGE 1959,
MEYER & WILSON 1971, E. WILLIAMS 1976,
SAVAGE & TALBOT 1978, GUYER & SAVAGE 1987,
1992, SAVAGE & GUYER 1989, E. WILLIAMS 1989,
MCCRANIE et al. 1992, 1993a,b, AROSEMA &
IBANEZ 1994, KOHLER 1996a-c, 1999a, KOHLER &
MCCRANIE 1998, KOHLER & OBERMEIER 1998,
SAVAGE & GUYER 1998, KOHLER et al. 1999,
KOHLER & MCCRANIE 2001, KOHLER et al.
200la, MCCRANIE & KOHLER 2001, MCCRANIE et
al. 2000, 2001, NICHOLSON et al. 2001, KOHLER &
VESELY 2003, KOHLER & ACEVEDO 2004, KOHLER
et al. 2005, KOHLER et al. 2006, KOHLER 2007,
KOHLER et al. 2007, KOHLER & SUNYER 2008,
KOHLER & SMITH 2008, PONCE & KoHLER 2008
(morphology, taxonomy, systematics); SEXTON et
al. 1971, FITCH 1973a, b, FITCH et al. 1976,
FITCH 1982 (ecology, ethology); FLASCHEN-
DRAGER & WIJFFELS 1997 (captive husbandry
and breeding)

133
 lguanidae

Basiliscus
All four known species of basilisks are
found in Central America. Earlier claims
that the distribution of the Ecuador
basilisk (Basiliscus galeritus) reached as
far as Costa Rica (LANG 1989) proved to be
erroneous, probably due to misidentifica-
tion of a juvenile B. basiliscus (J.M. SAVAGE
pers. comm. 1996). However, this species
has been collected in the Darien region of
Panama (R. IBANEZ pers. comm. 2000).
Basilisks, with their head and dorsal
crests, which give them a truly dragon-like
appearance, are among the most impres-
sive and bizarre lizards of Central America.
Their ability to run on their hind legs even
across rapidly flowing water has earned
them the local name "!guano Jesucristo".
Two important adaptations that allow
these animals to walk on the water's sur-
face are broad fringes of skin on the toes,
which distribute their body weight over a
larger area and the relatively heavy tail,
which acts as a counter-balance to the
upraised body. These attractive lizards are Fig. 367. Adult male of Basiliscus vittatus (Isla
found in a number of habitats, wherever de Utila, Honduras). Photo: G. Kohler
water in the form of streams, ponds, or
swamps is present, and are quite numer-
ous in many areas. On the Islas de la
Bahia, Honduras, Basiliscus vittatus is vir- Female basilisks produce three to seven
tually ubiquitous. Basilisks feed on insects, clutches of eggs per season, each with 4-18
spiders, and small vertebrates, as well as eggs, which hatch in 2-3 months (KOHLER
the occasional bit of vegetable matter. 1993b).

_. . . .
• Basiliscus basiliscus
• Basiliscus plumifrons
"" Basi/iscus galeritus

Basiliscus vittatus

134

Fig. 368. Juvenile Basiliscus basiliscus
(Manuel Antonio, Costa Rica). Photo: G. Kohler
Fig. 369. Adult male of Basiliscus plumifrons
(near Pueblo Wiso, Jinotega, Nicaragua).
Photo: G. Kohler
Fig. 370. Adult male of Basiliscus galeritus
(Ecuador). Photo: G. Kohler
Iguanidae
Fig. 371. Adult male of Basiliscus basiliscus
(Manuel Antonio, Costa Rica). Photo: G. Kohler
Basiliscus basiliscus (LINNAEUS 1758),
Systema Naturae, ed. 10: 206; type locality:
1•
"America australi". SVL to 240 mm .
Southwestern Nicaragua to northwestern ;;!
South America, sea level to 1200 m elevation.
Basiliscus galeritus DuMERIL 1851, Cat.
Meth. Coll. Rept. Mus. Paris: 61; type locality:
"N.-Grenade" [=Colombia] . SVL to 240 mm.
Darien region of Panama to western Ecuador,
sea level to 1000 m elevation.
Basiliscus plumifrons COPE 1876, J. Acad.
Nat. Sci. Philadelphia (2) 8: 125; type locality:
Sipurio, Costa Rica. SVL to 240 mm. Eastern
Honduras to Panama, sea level to 780 m ele-
vation.
Basiliscus vittatus WIEGMANN 1828, Isis von
Oken 21: 373; type locality: Mexico. SVL to
170 mm. Jalisco and Tamaulipas, Mexico, to
Panama, sea level to 1500 m elevation.
Fig. 372. Subadult of Basiliscus vittatus
(Belize). Photo: G. Kohler
135
 Iguanidae

Key to Basiliscus
1 a Ventral scales keeled; 1-2 mental shields in
contact with infralabials (Fig. 374a); head
crest of adult males single, triangular in
outline ....................... Basiliscus vittatus
b Ventral scales smooth; 3-4 mental shields
on each side in contact with infralabials
(Fig. 374b); head crest of adult males dif-
ferent than above ......... ........... ............. 2 a. Basi/iscus vittatus b. Basiliscus basiliscus
2 a A series of enlarged, raised middorsal Fig. 374. Arrangement of chin shields (orange).
scales, regularly separated from each other
by 2-4 small scales (Fig. 375); head crest of
adult males single, fleshily thickened at
base; no dorsal fin .. Basiliscus galeritus
b Middorsum with slightly enlarged scales
showing no evidence of regular spacing;
head crest of adult males different; a large
dorsal fin present ............................... 3 Fig. 375. Mid-dorsal scales in Basiliscus galeritus.
3 a Ground color green; one or two longitudi-
nal rows of pale spots along sides of body;
....N
I:"'
head crest of adult males comprising two
lobes; chin region of juveniles uniform Corytophanes
......... ... ............... Basiliscus plumifrons
b Ground color brown; continuous pale longi- The helmeted iguanas are among the rela-
tudinal stripes along sides of body; head tives of the basilisks and their distribution
crest of adult males single, often extending
as a thin process posteriorly; chin region of is restricted to Mexico and Central
juveniles with distinct longitudinal dark America. All helmeted iguanas have a later-
streaks .................. Basiliscus basiliscus ally flattened body and delicate proporti-
ons. Even at single locality, the coloration
of the C. cristatus can vary from dark
Further Reading
MATURANA 1962, LANG 1989, KOHLER 1993e, brown to olive green and yellow with irreg-
1999b ular blackish brown spots or crossbands.
The ventral side is pale brown in color with
a few dark spots. Males are easily distin-
guished from the females by the hemipenial
bulge at the base of the tail, larger helmets
and somewhat larger heads.

~e
The helmeted iguana is a peaceful tree-
dweller that spends most of its time in par-
tial shade, lying in wait for its predomi-
nantly insect prey. Analysis of stomach
B. vlftatus B. galeritus contents reveals that C. cristatus in the
wild rarely, if ever, eat during the day
(ANDREWS 1979). Helmeted iguanas pro-

0~
duce 4-11 eggs per clutch, each weighing 2-
4 g and measuring 10-15 x 18-30 mm
(REAM 1965, BOCK 1987). Depending on
B. plumifrons B. basiliscus temperature, the eggs will hatch after an
incubation period of 60-155 days (KOHLER
1999b). Hatchling C. cristatus have a total
length of57-75 mm and a SVL of25-30 mm
Fig. 373. Shape of helmet in Basiliscus. (REAM 1965, KOHLER et al. 1994).

136
 Iguanidae

Fig. 376. Corytophanes percarinatus

(El lmposible National Park, Ahuachapan, El
Salvador). Photo: G. Kohler

Fig. 378. Corytophanes hernandesii (El Ocote

region, Chiapas, Mexico). Photo: G. Kohler

Fig. 377. Corytophanes cristatus (Saslaya Fig. 379. Corytophanes percarinatus (Chiapas,
National Park, Nicaragua). Photo: G. Kohler Mexico). Photo: A. Ramirez V.
137
 lguanidae

Corytophanes cristatus (MERREM 1822),

Tentamen Systematis Amphibiorum: 50; type
locality: "Ceylon" [in error]. SVL to 120 mm.
Veracruz, Mexico, through Central America
as far as northwestern Colombia, sea level to
1640 m elevation in tropical and subtropical
wet and rain forest.
Corytophanes hernandesii (WIEGMANN 1831),
in GRAY, Synopsis Species Class Reptilia, in
GRIFFITH, Cuviers Animal Kingdom 9: 45;
type locality: Mexico. SVL to 100 mm. From
Central Veracruz, Mexico, along the base of
the Yucatan Peninsula into northwestern
Honduras, sea level to 1100 m elevation in
rain forest and dry forest.
Corytophanes percarinatus DUMERIL 1856, Fig. 380. Corytophanes cristatus (Parque
Arch. Mus. Hist. Nat. Paris 8: 518; type loca- Nacional Pico Bonito, Atlantida, Honduras).
lity: "Ascuintla, dans ! 'Amer. centrale"
[= Escuintla, Guatemala] . SVL to 100 mm.
Photo: G. Kohler
Isthmus of Tehuantepec, Mexico, along the
Pacific coast as far as El Salvador and into
southwestern Honduras, 200-2500 m eleva-
tion.

Key to Corytophanes
1 a Crest of helmet continues without inter-
ruption into the dorsal crest .. ................ 2
b Crest of helmet not continuous with dorsal
crest .. ... ..... Corytophanes hernandesii
2 a Helmet restricted to bony skull process; a. C. cristatus b. C. percarinatus
scales on dorsal surface of head keeled or
at least clearly lined ............. ............ ..... .. .
. . . . . . . . . . . . ... . Corytophanes percarinatus
b Helmet extends past bony skull process; Fig. 381. Shape of
scales on dorsal surface of head smooth head (lateral view) in
........................ .. Corytophanes cristatus the species of
Corytophanes.
Further Reading c. C. hernandesii
LANG 1989, KOHLER et al. 1994, KOHLER 1999b.

Corytophanes hernandesii
Corytophanes cristatus
~~~~.....;:...::~--"'~~~ ..... • Corytophanes percarinatus

138

Ctenosaura
Depending upon species, spiny-tailed iguan-
as can reach a total length of anywhere
from 25 to over 100 cm. These striking ig-
uanas are found in the lowlands of Mexico
and Central America from the south-
eastern Baja California and the southern
Sonora in western Mexico, also more or
less above the Tropic of Cancer in eastern
Mexico as far as Panama, and on certain
islands in the eastern Pacific, the western
Caribbean and the Sea of Cortez.
Of the larger species (maximum total
length over 100 cm), C. acanthura and C.
pectinata reach only the north of Central
America in the area of the Isthmus of
Tehuantepec, Mexico, whereas C. similis is
distributed all across the Central
American land bridge. These three species
are predominantly ground dwellers, but
they will also climb in trees. The smaller
ctenosaur species, as a rule, are tree-dwel-
lers. When threatened, they will retreat
into knotholes, and block the entrance with
their spiny tails. The beautifully colored
Yucatan Spiny-tailed Iguana (Ctenosaura
defensor) is a ground dweller in the rocky
terrain of the northern Yucatan Peninsula.
Limestone crevices serve as hiding places.
All spiny-tailed iguanas lay eggs, with
clutch sizes ranging from 5-15 (small spe-
cies) and 20-88 (C. similis). Suitable
nesting sights are open, sun exposed
areas where up to five females will
use a common system of holes to
lay their eggs (FITCH & HACKFORTH-
J ONES 1983). Three nest tunnels
examined by FITCH & HACKFORTH-JONES
(1983) were 11-22 m long and reached 20-
36 cm under the surface of the
ground. The eggs require an incubation of
approximately three months (KOHLER
1993a). The hatchlings of most spiny-tailed
iguana species are predominantly green in
color with green markings. One exception
in this regard is the Utila Iguana (C. bake-
ri), whose hatchlings are grey-brown with
lguanidae
virtually no markings, possibly due to the
habitat (border zones of mangrove swamp)
in which the hatchlings of this species are
found . Spiny-tailed iguanas reach sexual
maturity at two to three years of age.
Based on research of the geographic varia-
bility of the spiny-tailed iguana formerly
designated as Ctenosaura palearis,
BUCKLEY & AxTELL (1997) have described
the Honduran populations as a separate
species, C. melanosterna. Thus C. palearis
is restricted in its distribution to the
Motagua Basin of Guatemala. Similarly,
the populations of what was formerly
known as C. quinquecarinata have been
split into two species. Because of the holo-
type's identity, the name quinquecarinata
has to apply to the Nicaraguan and Costa
Rican populations and the Mexican popu-
lations received a new name, C. oaxacana
(HASBUN & KOHLER 2001, KOHLER &
1
....
..:1
HAsBUN 2001). Recently, the distributional
range of the three large ctenosaur species
(C. acanthura, C. pectinata, and C. similis)
in the area of the Isthmus of Tehuantepec
and in the Central Depression of Chiapas
was studied (KOHLER & MENDOZA
manuscript). Contrary to earlier views that
considered the ctenosaur species in the
Central Depression of Chiapas to be C. pec-
tinata (ALVAREZ DEL TORO 1960, 1983;
JOHNSON 1989, 1990, STAFFORD 1990), our
field work provided evidence that it is
actually C. acanthura.
139
 Iguanidae
Ctenosaura acanthura (SHAW 1802), Gen.
Zool., London 3: 216, type locality: unknown.
SVL to 315 mm. Along the Caribbean versant
from central Tamaulipas to southern
Veracruz, Mexico; also in the Central
Depression of Chiapas, Mexico, sea level to
1000 m elevation.
Ctenosaura alfredschmidti KOHLER 1995a,
Salamandra 31: 5; type locality: 70 km E
Escarcega on road to Chetumal, Campache,
Mexico. SVL to 170 mm. Southern Campeche
and Quintana Roo, Mexico, and adjacent regi-
ons in Peten, Guatemala.
Ctenosaura bakeri STEJNEGER 1901, Proc. U.S.
Natl. Mus. 23: 467; type locality: Isla de Utila,
Islas de la Bahia, Honduras. SVL to 300 mm.
Endemic to Isla de Utila, Islas de la Bahia,
Honduras, sea level to 30 m elevation.
Ctenosaura defensor (COPE 1866), Proc. Acad.
Nat. Sci. Philadelphia 18: 124; type locality:
Yucatan, Mexico. SVL to 145 mm. Northern
portion of the Yucatan Peninsula, Mexico, sea
level to 100 m elevation.
Fig. 383. Ctenosaura acanthura (Monte Bonito,
Chiapas, Mexico). Photo: G. Kohler
Fig. 384. Ctenosaura alfredschmidti (southern
Campeche, Mexico). Photo: G. Kohler
140
Ctenosaura fiauidorsalis KOHLER & KLEMMER
1994, Salamandra 30: 197; type locality: 1 km
S La Paz, 750 m elevation, l4°16'N, 87°40'W,
La Paz, Honduras. SVL to 170 mm. Disjunct
populations from eastern Guatemala to
eastern El Salvador and southwestern
Honduras to the Comayagua Valley in south-
central Honduras; 35 to 1010 m elevation.
Ctenosaura melanosterna BUCKLEY & Ax.TELL
1997, Copeia 1997: 139; type locality: 2 km S
Coyoles Central, Yoro, Honduras. SVL to 310
mm. Valle de Rio Aguan and Cayos Cochinos,
Honduras, sea level to 250 m elevation.
Ctenosaura oaxacana KOHLER & HASBUN
2001, Senckenbergiana biologica 81: 260; type
locality: Tehuantepec, Oaxaca, Mexico. SVL
to 170 mm. Restricted to the Pacific versant
of the Isthmus of Tehuantepec, Oaxaca,
Mexico; sea level to 250 m elevation.
Ctenosaura oedirhina DE QUEIROZ 1987a,
Copeia 1987: 892; type locality: approx. 4.8
km W Roatan on path to Flowers Bay, Isla de
Roatan, Islas de la Bahia, Honduras. SVL to
270 mm. Isla de Roatan and its satellite
islands (Santa Elena, Barbareta, Big Pigeon
Cay), Islas de la Bahia, Honduras, sea level to
100 m elevation.
Ctenosaura palearis STEJNEGER 1899, Proc.
U.S. Natl. Mus. 21: 381, type locality: Gualan,
Guatemala. SVL to 300 mm. Valle de Rio
Motagua, Guatemala, 150-700 m elevation.
Ctenosaura pectinata (WIEGMANN 1834),
Herpetologica Mexicana: 42; type locality:
unknown. SVL to 353 mm. Southern Sinaloa,
Mexico, along the Pacific versant to western
Chiapas, sea level to 2000 m elevation.
Ctenosaura quinquecarinata (GRAY 1842),
Zool. Misc. 1842: 59; type locality:
"Demerara?" [based on morphological and
molecular genetic data, HAsBUN & KOHLER
2001 allocated the C. quinquecarinata holo-
type to Nicaraguan and Costa Rican popula-
tions) . SVL to 200 mm. Isolated populations
in Nicaragua and northwestern Costa Rica,
sea level to 250 m elevation.
Ctenosaura similis (GRAY 1831), Synopsis
Species Class Reptilia, in GRIFFITH, Cuviers
Animal Kingdom 9: 38; type locality: un-
known. SVL to 489 mm. Isthmus of
Tehuantepec (central Tabasco on Caribbean
versant and central Oaxaca on Pacific ver-
sant), Mexico, across all of Central America
(except highlands and very humid regions) to
Panama, sea level to 1320 m elevation.
Fig. 385. Ctenosaura bakeri (Isla de Utila, Fig. 389. Ctenosaura oaxacana (Oaxaca,
Honduras). Photo: W. Steiger Mexico). Photo: G. Kohler

Fig. 386. Ctenosaura defensor (near Telcha- Fig. 390. Ctenosaura oedirhina (Isla de
quillo, Yucatan, Mexico). Photo: G. Kohler Roatan, Honduras). Photo: G. Kohler

Fig. 387. Ctenosaura flavidorsalis (near La Fig. 391. Ctenosaura palearis (upper Motagua
Paz, Honduras). Photo: G. Kohler Valley, Guatemala). Photo: J. A. Campbell

Fig. 388. Juvenile of Ctenosaura melanosterna. Fig. 392. Ctenosaura pectinata (Oaxaca,
Photo: G. Kohler Mexico). Photo: G. Kohler
141
 Iguanidae

Key to Ctenosaura
1 a Tail at the point of maximum circumfer-
ence wider than high (width-height ratio
1.66 - 1. 76); ratio of tail length/SVL less
than 1.5; maximum SVL to 200 mm; distal
dark tail bands not much longer than
proximal ones .......... .. .......... ..... ............. 2
b Tail at the point of maximum circumfer-
ence approximately as wide as high
(width-height ratio 0.90-1.11); ratio of tail
length/SVL greater than 1.6; maximum
SVL greater than 250 mm; posterior dark
tail bands twice as long as anterior ones 6
Fig. 394. Ctenosaura quinquecarinata (near
2 a Only about 2/3 of tail covered with whorls Teustepe, Boaco, Nicaragua). Photo: F. Schmidt
of enlarged spinous scales; parietal eye
clearly visibledwith naked eye; keels at the
base of the 3r toe not fused together ..... 3
b Entire tail covered with whorls of en-
larged spinous scales; parietal eye barely
visibl'awith naked eye; keels at the base of
the 3r toe fused together ....................... 5
3 a Adult males with middorsal crest spines
<2 mm high; usually gaps present along
the crest row in nuchal and anterior tho-
racic region; 2 or 4 postmentals; dorsal
coloration usually with some yellow ele-
ments ......... ... Ctenosaura flavidorsalis
b Adult males with middorsal crest spines
usually >2 mm high; nuchal and thoracic
crests continuous; 2 postmentals (Fig.
397 a); absence of yellow elements in dor-
sal coloration ......................................... ... 4

C. quinquecarinata

Fig. 393. Dorsal view of anterior Fig. 396. Mid-dorsal scales.

tail. Photos: G. Kohler Photos: G. Kohler

142
 Iguanidae

4 a Tail less spinous (paramedian caudal

scales spinous on proximal 3-7 [mean
value 4.5) caudal whorls); usually body
pattern of a broad pale middorsal band,
which may be fragmented into large
blotches; SVL to 170 mm in males, 124
mm in females ... Ctenosaura oaxacana
b Tail more spinous (paramedian caudal
scales spinous on proximal 4-13 [mean a. Ctenosaura quinquecarinata b. Ctenosaura similis
value 6.4) caudal whorls); usually no
broad pale middorsal band evident; SVL to Fig. 397. Chin scalation (postmentals orange).
195 mm in males, 145 mm in females
... ... ........ Ctenosaura quinquecarinata
5 a 27-29 tail whorls; ratio of tail length/SVL
0.79-0.85; rows of medial dorsals uniform
and reaching to pelvic region; parietals
well formed with small parietal eye; only
the first 0-3 interspaces between tail
whorls without complete rows of small flat
scales; SVL to 170 mm .......................
............ ....... Ctenosaura alfredschmidti
b 22-24 tail whorls; ratio of tail length/SVL
0.65-0.74; rows of enlarged medial dorsals
irregular and reduced; parietals reduced,
parietal eye not distinguishable; usually a. Ctenosaura similis b. Ctenosaura quinquecarinata
no complete rows of enlarged spiky scales
between the first eight tail whorls; SVL to Fig. 398. Dorsal view of anterior tail (inter-
145 mm ..... ....... .... Ctenosaura defensor calary scales orange)
6 a 31-47 dorsal spines; dewlap more or less
strongly developed .. .... ... .. ... .... .. ............ 7
b 66-106 dorsal spines; no dewlap, rather
just a transverse fold across the throat .. 9
7 a Two complete rows of small intercalary
scales between whorls of enlargethspinous
scales distally (starting after 7t to the
17th tail whorls); markings and coloration
of dorsal spines conform to that of adja-
cent dorsal scales; dorsum predominantly a. Ctenosaura alfredschmidti b. Ctenosaura similis
unicolor grey-brown Ctenosaura bakeri
b One complete row of small intercalary Fig. 399. Scalation on dorsal surface of shank.
scales between all whorls of enlarged
spinous scales (two rows of intercalary
scales present between most proximal 3
whorls in some specimens); markings and
coloration of dorsal spines do not conform
to that of adjacent dorsal scales; several
wide grey to black cross bands on dorsum,
separated by rows of small pale spots ... .. 8
8 a Ventral area predominantly black in color-
ation; 9-12 supralabials ... ........ ... .... ..... .
. . . . . . . . . . . . . . . ... . . . . . . ... Ctenosaura palearis
b Ventral area brown in coloration; 8-10
supralabials Ctenosaura melanosterna
9 a Enlarged heavily keeled scales on the dor-
sal surface of shank (Fig. 399a); snout of
older individuals blunt and rounded off Fig. 400. In older individuals of C. oedirhina
(Fig. 400); 9-15JemoralJ>ores; in the area the snout is blunt and rounded off.
between the 4tu and 5tu tail whorls, only Photo: G. Kohler

143
 Iguanidae
one complete row of small intercalary
scales ... ...... .. ....... Ctenosaura oedirhina
b Scales on the dorsal surface of shank not
enlarged and only slightly keeled (Fig.
399b); snout not blunt and rounded off; 4-
10 femoral pores; in the area between the
4th and sill tail whorls, at least two com-
plete rows of small intercalary scales ..... 10
lOa Scales of tail whorls very spiky (angle of
keels greater than 30 degrees); dorsal
crest usually interrupted in pelvic region;
in some specimens, rows of scales between
the 5th and 8th tail whorls reduced to one
complete row .... Ctenosaura acanthura
b Scales of tail whorls moderately spiky
(angle of keels less than 20 degrees) with
keels running diagonally (Fig. 401c);
dorsal crest in pelvic region interrupted or
not; always at least two complete rows of
scales between tail whorls ........... ... ..... 11
Ila Several dark crossbands on dorsum, which
usually have a pale center along the dor-
somedial line; dorsal crest continuous with
Fig. 401. Dorsal view of anterior tail.
Photo: G. Kohler
144
tail crest; in area between 3rd and 5th tail
whorls, two complete rows of small inter-
calary scales ...... ....... Ctenosaura similis
b No dark crossbands on dorsum; dorsal
crest interrupted in pelvic aegion or ~ot; in
the area between the 3r and 5t tail
whorls, 3 complete rows of small inter-
calary scales ....... Ctenosaura pectinata
Further Reading
BAILEY 1928, HENDERSON 1973, FITCH &
HENDERSON 1977, 1978, VAN DEVENDER 1982,
FITCH & HACKFORTH-JONES 1983, GICCA 1983,
DE QUEIROZ 1987a,b, 1990a,b, BUCKLEY &
Ax.TELL 1990, 1997, KOHLER 1991a, 1993a,c,
1994a,b, 1995a-e, 1996e, 1998a,b,e,g, KOHLER &
KLEMMER 1994, KOHLER & STREIT 1996, KOHLER
& VESELY 1996, KOHLER & RITTMANN 1998,
KOHLER et al. 2000, HAsBON & KOHLER 2001,
HAsBUN et al. 2001, KOHLER & HAsBUN 2001,
KOHLER 2002, RADACHOWSKY et al. 2004,
HAsBON et al. 2005, ZARZA et al. 2008
Ctenosaura
• C. similis
• C. acanthura
• C. pectinata
Ctenosaura • C. defensor
• C. quinquecarinata
.a. C. alfredschmidti
• C. melanosterna
!::> C. oedirhina
\1 C. palearis
 Iguanidae

Fig. 402. Enyalioides heterolepis (Lita - San Lorenzo, Esmeraldas, Ecuador). Photo: K.-H. Jungfer

Enyalioides Enyalioides heterolepis (BOCOURT 1874), Ann.

Sci. Nat. Paris (5) 19: 1; type locality: Vera-
gua, Panama. SVL to 138 mm. Panama to
From the primarily South American genus northwestern Ecuador, 100-600 m elevation.
Enyalioides, only one species, E. hetero-
lepis, reaches as far as southern Central Further Reading
America. So far, the existence of this spe- NICHOLSON 1998, LIPS 1999
cies has only been established at a few
localities in the Panamanian provinces of
Bocas del Toro, Colon, Code, Panama, and
San Blas (NICHOLSON 1998, LIPS 1999,
KOHLER unpubl. data). The species of this
genus are sit-and-wait predators in the
rain forest that primarily feed on insects
and spiders. At night, these lizards will
sleep at a height of 0.5-1.5 m on tree
trunks or lianas, sometimes even in the
rosettes of the aerial roots of palm trees.

.& Enyalioides heterolepis

145
 lguanidae

Iguana
The green iguana (Iguana iguana) is cer-
tainly the best known lizard in Central
and South America. The preferred habitat
of this magnificent giant lizard is lowland
forest, near streams, rivers, or lakes.
Whereas adult iguanas spend a lot of their
time in the treetops, juveniles are often
found closer to the ground in bushes.
However, I have also observed adult green
iguanas basking in the vegetation along
the shoreline in Tortuguero, Costa Rica.
Green iguanas are impressive swimmers
and divers, as they have demonstrated by Fig. 403. Adult male Iguana iguana rhinolo-
leaping into the water from as much as a pha (Belize). Photo: G. Kohler
10 m height when I approached within a
few meters of them. In the wild, green ig-
uanas consume an almost exclusively vege-
tarian diet, consisting mostly of leaves.

At the beginning of the dry season, the

female iguana will excavate a nesting site,
which, depending on how hard the ground
is, will be from 30 to 200 cm long (RAND
1968, RAND & DUGAN 1983). Clutch size
varies from 20 to 60 (seldom to 80) eggs,
primarily depending upon the size of the
female. The young will hatch following an
incubation period of approximately three
months, which usually corresponds with
the onset of the rainy season. Thus, the
hatchling iguanas are ensured a ready Fig. 404. Subadult Iguana iguana rhinolopha
supply of tender leaves as nourishment. (sleeping, Bartola, Nicaragua).Photo: G. Kohler

Iguana iguana (LINNAEUS 1758), Systema

Naturae, ed. 10: 206; type locality: "Indiis".
SVL to 550 mm. Sinaloa and Veracruz,
Mexico, across all of Central America as well
as large parts of South America, sea level to
1000 m elevation. Two subspecies: Iguana i.
iguana (Costa Rica southwards) and I. i. rhi-
nolopha (WIEGMANN 1834) (Mexico to Costa
Rica).

Further Reading
BURGHARDT & RAND 1982, KOHLER 1998d

• 1guanaiguana

146
 Iguanidae

Laemanctus

The casquehead iguanas, which are found

only in Central America and Mexico, are
remarkably well adapted for life in the
trees, which they leave only to lay eggs.
Predictably, casquehead iguanas are
difficult for the human observer to spot
amongst the green leaves. Several times a
year, Laemanctus females will produce a
clutch of 4-9 eggs, which hatch after only
two months at an incubation temperature
of 28-30 °C. Freshly hatched Laemanctus
longipes, on average, have a SVL of 50 mm
and a tail length of about 170 mm. Fig. 405. Laemanctus longipes. Photo: F. Riedel

Laemanctus longipes WIEGMANN 1834,

Reptilia and Batrachia. Biologia Centrali-
Americana: 46; type locality: Jalapa [=
Xalapa], Mexico. SVL to 150 mm. Central
Veracruz, Mexico, to Nicaragua, near sea
level to 1200 m elevation in moist forest and
rain forest. In Central America, the subspe-
cies Laemanctus l. deborrei BOULENGER 1877
(Isthmus of Tehuantepec to Nicaragua) and
L. l. waltersi SCHMIDT 1933b (northwestern
Honduras) occur.
Laemanctus serratus COPE 1864, Proc. Acad.
Nat. Sci. Philadelphia 16: 176; type locality:
Orizaba Valley, Mexico. SVL to190 mm.
Tamaulipas, Mexico, along the Atlantic coast
as far as northwestern Honduras, near sea
level to 1500 m elevation in dry forest. In
Central America, the subspecies L. s. alticoro- Fig. 406. Laemanctus serratus (female).
natus COPE 1864 (Yucatan Peninsula) and L . Photo: G. Kohler
s. mccoyi PEREZ-HIGAREDA & VOGT 1985
(southern Veracruz) occur.

Laemanctus longipes Laemanctus serratus

147
 lguanidae ..............................................................

Key to Laemanctus break into small irregular spots. According

1 a Posterior of head with spiky scales; dis-
tinct dorsal crest present; scales on dorsal
surface of snout large and regular
............. ... ............ Laemanctus serratus
b Posterior of head without spiky scales; no
dorsal crest present; scales on dorsal sur-
face of snout small and irregular .....
... .... ... ..... .. ......... Laemanctus longipes
Further Reading
McCOY 1968, LANG 1989, KOHLER 1999b,
MCCRANIE & KOHLER 2004a,b,c
to the observations of CORREDOR et al.
(1985), the animals live in burrows, which
they excavate in the floor of the rain
forest. During the day, these lizards will
sit motionless in front of the entrances to
their burrows attentively watching the
surrounding area. If they feel threatened,
they will disappear with lightning speed
into their burrows, blocking the entrance
with their spiky tails. Predators, such as
snakes, are thus prevented from forcing
their way into the burr.ows. Morunasaurus
annularis reproduces by laying eggs.

Morunasaurus Morunasaurus groi DUNN 1933a, Occ. Pap.

Boston Soc. Nat. Hist. 8: 76; type locality:
Valle de San Anton, Panama. SVL to 110 mm.
Three species make up the genus Central Panama and northwestern Colombia
Morunasaurus: M. annularis (premontane in tropical rain forest at 700-800 m elevation.
areas on the Amazon side of Ecuador and
Colombia), M. peruuianus (Amazonian Further Reading
Peru) and M. groi, which is distributed as DUNN 1933a, CORREDOR et al. 1985, KOHLER
far north as southern Central America. 199lb, KOHLER 2003
They are diurnal ground-dwellers of
stocky build that live in the shady areas of
the rain forest. Both species have a very
heterogeneous scalation on the dorsal and
lateral surfaces of the trunk, with large
conical scales scattered among the granu-
lar scales. The tail is equipped with regu-
larly arranged whorls made up of enlarged
spiky scales, which become increasingly
smaller distally and are barely distinguish-
able at the tail tip. Morunasaurus groi is
predominantly reddish brown in color. The
body has dark brown crossbands, which • Morunasaurus groi
reach to the midpoint of the sides and then

Fig. 407. Female of Morunasaurus groi (El Valle de Anton, Panama). Photo: W. E. Duellman
148
 Iguanidae

Phrynosoma

Of the twelve species of horned lizard that

are distributed across Mexico and the
USA, only the giant horned lizard
(Phrynosoma asio) reaches as far as the
north of Central America (REEVE 1952).
Their presence in Guatemala has not been
verified by any finds in this century. In
Oaxaca and Chiapas, the giant horned
lizard is found in sparse forest and rocky
semi-desert (BAUR & MONTANUCCI 1998,
KOHLER pers. obs. 1993). Phrynosoma asio
feeds on insects and spiders, with the larg- Fig. 409. Phrynosoma asio. Photo: G. Kohler
est portion of their diet, as with all horned
lizard species, consisting of ants (MONTA-
NUCCI 1989). This is one of the egg-laying
species of the genus, with clutch size rang-
ing from 15 to 24 eggs. Young measuring
32-37 mm SVL will hatch after an incuba-
tion period of 93 to 106 days. (BAUR & Polychrus
MONTANUCCI 1998).
Polychrus gutturosus, the only bush anole
species in Central America, lives in the
Phrynosoma asio COPE 1864, Proc. Acad. Nat. crown region of the lowland rain forest
Sci. Philadelphia 16: 178; type locality: and, apparently, only occurs in low popula-
Colima, Mexico. SVL to 124 mm. Colima, tion densities. Because they move very
Mexico, along the Pacific side of Mexico to
Chiapas, near sea level to 750 m elevation in little, bush anoles are difficult to detect in
savanna and dry forest the leafy tangle of the rain forest trees. On
the 9th of May, ROBERTS (1997) observed a
copulating pair of P. gutturosus about 2
Further Reading meters up a tree in La Selva, Costa Rica. A
REEVE 1952, MONTANUCCI 1987, 1989, BAUR &
MONTANUCCI 1998 second female with palpable eggs was
examined on the 24th of July (ROBERTS
1997).

Fig. 408. Morunasaurus groi (El Valle de

Anton, Panama). Photo: B. Akeret
149
 Iguanidae
Fig. 410. Adult male of Polychrus gutturosus
(Rio San Juan, Nicaragua). Photo: G. Kohler
Fig. 411. Adult female of Polychrus gutturosus
(Rio San Juan, Nicaragua). Photo: G. Kohler
Polychrus gutturosus BERTHOLD 1845, Nachr.
Georg-Augusts Univ. und K. Ges. Wiss. Got-
tingen 3: 38; type locality: western Colombia
(see MYERS & BOHME 1996). SVL to 165 mm.
Northern Honduras to northwestern Ecuador
in rain forest, sea level to 700 m elevation.
Further Reading
TAYLOR 1956
150
Sceloporus
In the dry regions of Mexico and Central
America, next to spiny-tailed iguanas,
spiny lizards are the dominant iguanine
group. The genus contains over 70 species,
with increasingly fewer varieties occur-
ring towards the south of Central America
(only three species occur south of
Guatemala). Spiny lizards are diurnal
small to mid-sized lizards, most of which
are covered with extremely spiky scales.
They feed on various arthropods, as well
as smaller vertebrates. Among the
Sceloporus species are ground-dwellers
and saxicolous species, as well as primari-
ly arboreal species. Most of the Sceloporus
species reproduce by laying eggs whereas
the species of the formosus group, as well
as S. serrifer (torquatus group), are live-
bearers.
The species of the variabilis group are
ground dwellers in the lowlands and, in
some areas, occur in high densities.
Frequently these animals can be seen on
tree stumps or rocks. The males, as with
all species of the genus, are highly territori-
al and attempt to intimidate their rivals
with push up-like movements of the upper
body. All species of this group lay eggs (2-7
eggs per clutch depending on species).
The systematics of the species of the for-
mosus group have yet to be satisfactorily
resolved, as the morphological variation,
as well as geographic distribution of cer-
tain species, is as yet unknown. Thus,
STUART (1971) claimed that comparisons
between populations of Sceloporus mala-
chiticus produce "(. .. ) only a hopeless jum-
ble of random variations". In his generic
revision, SMITH (1939a) considered mala-
chiticus and smaragdinus as subspecies of
formosus. Subsequently, both taxa have
been designated as full species (STUART
1971, VILLA et al. 1988, KOHLER 1990).
Also, the exact distribution of the various
species of the formosus group remains to
be well documented. SMITH et al. (2001)
 Iguanidae

stated that S. acanthinus ranges into El

Salvador, whereas KOHLER & HEIMES
(2002), partly based on advice from Eric N
Smith (pers. comm. to P. Heimes), referred
all malachite Sceloporus in El Salvador to
S. malachiticus, and I have used the latter
name for all El Salvadoran populations
herein.

With a vertical distribution of600-3800 m

above sea level, S. malachiticus and S.
smaragdinus are typical highland dwel-
lers (MA.RION & SEXTON 1971, KOHLER Fig. 412. Adult male of Sceloporus acanthinus
1990). They are found on trees, fence (near Guanagasapa, Escuintla, Guatemala).
posts, and on the roofs of houses (MERTENS Photo: G. Kohler
1952c, ROBINSON 1983). Usually only a
single male will inhabit a tree, which it
will defend from rivals (MERTENS 1952c).
Animals found at higher elevations tend
more towards a terrestrial lifestyle
(ROBINSON 1983). On Cerro de la Muerte
(Cordillera de Talamanca, Costa Rica,
3000 m above sea level), S. malachiticus
will inhabit natural stone walls and rocks
covered with mosses and ferns. At this ele-
vation, S. malachiticus is active only when
the sun shines. The air temperature mea-
sured 150 cm above the ground barely
reaches to 20 °C despite the sun. These
lizards, therefore, have adapted to optimal Fig. 413. Sceloporus taeniocnemis (male; near
use of the microclimate. They flatten their San Cristobal de las Casas, Chiapas, Mexico).
bodies and turn very dark while basking Photo: G. Kohler
on rocks protected from the wind. The
females of S. malachiticus produce one
brood of 3-10 young per year (MARION & found on individual stone blocks or on
SEXTON 1971, KOHLER 1990, 1993d), while piles of rocks. This species lays 4-5 eggs,
at lower elevations, births occur more or which hatch after 50-53 days at an incu-
less regularly throughout the year. In bation temperature of 29 °C. The young
populations above 2000 m elevation, measure from 17.0-18.5 mm SVL.
however, there is a highly seasonal repro-
ductive cycle, with births occurring prim- As the sole representative of the torquatus
arily in late January and February group in Central America, Sceloporus ser-
(MA.RION & SEXTON 1971). rifer reaches Guatemala and Belize. This
species can be seen in great numbers at
The siniferus group is represented in the Mayan ruins in Yucatan, although the
Central America by three species, all of animals are quite shy and will flee at the
which are ground dwellers in hot, dry slightest disturbance. These attractive
environments. In the Valle de Comayagua, lizards bask on trees, as well as on rocks
Honduras, S. squamosus is quite common- and brick walls, and will disappear into
ly seen (EISENBERG & KOHLER 1996). The the nearest crack or opening when threat-
spiny lizards in this area are generally ened.
151
 Iguanidae
Sceloporus acanthinus BOCOURT 1873, Ann.
Sci. Nat., Zool. (5) 17 (6): 24; type locality: San
Agustin, near Volcan de Atitlan, Guatemala.
SVL to 99 mm. Pacific versant of Chiapas,
Mexico, and Guatemala, 400-1500 m eleva-
tion.
Sceloporus carinatus H. SMITH 1936b, Proc.
Biol. Soc. Washington 49: 89; type locality:
near Tuxtla Gutierrez, Chiapas, Mexico. SVL
to 55 mm. Grijalva Valley of Chiapas, Mexico,
and adjacent Guatemala, 500-1000 m elevation.
Sceloporus chrysostictus COPE 1866, Proc.
Acad. Nat. Sci. Philadelphia 18: 125; type
locality: Yucatan, Mexico. SVL to 60 mm.
Yucatan Peninsula from central Belize and
north Guatemala northwards in dry forest,
sea level to 200 m elevation.
Sceloporus cozumelae JONES 1927, Occ. Pap.
Mus. Zool., Univ. Michigan 186: 1; type locali-
ty: Isla Cozumel, Quintana Roo, Mexico. SVL
to 54 mm. Coastal area of the northern
Yucatan Peninsula, including the offshore
islands Cozumel, Isla Contoy, and Isla
Mujeres in dry forest and open rocky areas,
sea level to 50 m elevation.
Sceloporus edwardtaylori H. SMITH 1936a,
Herpetologica 1: 6; type locality: Ixtepec (San
Geronimo), Oaxaca, Mexico. SVL to 107 mm.
Pacific coast of the Isthmus of Tehuantepec
(Oaxaca, Mexico) in dry forest, sea level to
750 m elevation.
Sceloporus internasalis H . SMITH &
BUMZAHEM 1955, Herpetologica 11: 118; type
locality: La Gloria, Oaxaca, Mexico. SVL to
100 mm. Los Tuxtlas region in southern Vera-
cruz, Mexico, southward through extreme
eastern Oaxaca to northwestern Chiapas, sea
level to 2000 m elevation in oak forest.
152
Sceloporus lunaei BocOURT 1873, Ann. Sci.
Nat., Zool. (5) 17 (10): 1; type locality: Plateau
of Guatemala. SVL to 95 mm. Motagua Basin
and Baja Verapaz, Guatemala, 150-1000 ele-
vation in dry forest.
Sceloporus lundelli H. SMITH 1939a, Field
Mus. Puhl. Zool. Ser. 26: 66; type locality:
Cohune Ridge, 20 miles SE Benque Viejo,
Belize. SVL to 100 mm. Yucatan Peninsula,
sea level to 300 m elevation in dry forest. Two
subspecies: Sceloporus l. lundelli (base of the
Yucatan Peninsula) and Sceloporus l. gaigeae
H. SMITH 1939 (northern part of the Yucatan
Peninsula).
Sceloporus malachiticus COPE 1864, Proc.
Acad. Nat. Sci. Philadelphia 16: 178; type
locality: "Arriba" ["this name apparently was
applied loosely to the Meseta Central" accord-
ing to SAVAGE 1974:77), Costa Rica. SVL to 98
mm. El Salvador and Honduras across
Nicaragua and Costa Rica to Panama, 600-
3800 m elevation in pine and cloud forest.
Sceloporus melanorhinus BocoURT 1876a,
Ann. Sci. Nat. Zool., Paris, (6) 3, art. 12: 2;
type locality: Isthmus of Tehuantepec,
Oaxaca, Mexico. SVL to 105 mm. Southern
Nayarit, Mexico, along the Pacific side of
Mexico to the Isthmus of Tehuantepec, as
well as in the Rio Grijalva Basin, Guatemala,
sea level to 2000 m elevation in dry forest. In
Central America, the subspecies Sceloporus
m. melanorhinus (Mexico) and S. m. stuarti
SMITH 1948 (Guatemala) occur.
Sceloporus serrifer COPE 1866, Proc. Acad.
Nat. Sci. Philadelphia 18: 124; type locality:
Yucatan, Mexico. SVL to 100 mm. Several iso-
lated populations from southern Tamaulipas,
Mexico, as far as Guatemala and Belize, as
well as in the north of the Yucatan Peninsula,
50-2300 m elevation in dry forest. In Central
America, the subspecies Sceloporus s. serrifer
(Yucatan Peninsula) and Sceloporus s. prezy-
gus H. SMITH 1942b (southwestern Guate-
mala) occur.
Sceloporus siniferus COPE 1870, Proc. Amer.
Phil. Soc. 11: 159; type locality: Pacific ver-
sant of Isthmus of Tehuantepec. SVL to 71
mm. From western Guerrero, Mexico, along
the Pacific Coast to eastern Chiapas, sea level
to 2000 m elevation in dry forest.
Fig. 414. Sceloporus carinatus (Rancho Alegre,
Chiapas, Mexico). Photo: G. Kohler
Fig. 415. Sceloporus chrysostictus (Telcha- Fig. 419. Sceloporus lundelli gaigeae (Coba,
quillo, Yucatan, Mexico). Photo: P. Heimes Quintana Roo, Mexico). Photo: J. C. Lee

Fig. 416. Sceloporus cozumelae (Telchac Fig. 420. Sceloporus malachiticus (Rio Sereno,
Puerto, Yucatan, Mexico). Photo: G. Kohler Panama). Photo: G. Kohler

Fig. 417. Sceloporus edwardtaylori (Oaxaca, Fig. 421. Sceloporus melanorhinus (Huautla,
Mexico). Photo: G. Kohler Morelos, Mexico). Photo: P. Heimes

Fig. 418. Sceloporus internasalis (Catemaco, Fig. 422. Sceloporus serrifer (Ruinas de
Veracruz, Mexico). Photo: P. Heimes Mayapan, Yucatan, Mexico). Photo: G. Kohler

153
 Iguanidae

Sceloporus smaragdinus BOCOURT 1873, Ann.

Sci. Nat., Zool. (5) 17 (10): 1; type locality:
Guatemala, in the vicinity of Solola,
Totonicapam, and Quetzaltenango. SVL to 85
mm. Highlands of Guatemala and Chiapas,
2000-4000 m elevation in pine and cloud
forest.
Sceloporus smithi HARTWEG & OLIVER 1937,
Occ. Papers Mus. Zool. Univ. Michigan 356: 1;
type locality: Quiengola Mountain, about 5
miles NW of the town of Tehuantepec,
Oaxaca, Mexico. SVL to 71 mm. Pacific side of
the Isthmus of Tehuantepec (Oaxaca,
Mexico), sea level to 600 m elevation in dry
forest.
Fig. 423. Sceloporus siniferus (near Mixte-
Sceloporus squamosus BocOURT 1874, Miss.
Sci. Mex., Rept.: 212; type locality: "Environs quilla, Oaxaca, Mexiko). Photo: G. Kohler
de Guatemala et de l'Antigua" (=Volcan
Antigua, Guatemala). SVL to 59 mm. From
western Chiapas, Mexico, along the Pacific
side of Central America into northwestern
Costa Rica, sea level to 1000 m elevation in
dry forest.
Sceloporus taeniocnemis COPE 1885, Proc.
Am. Phil. Soc. 22: 399; type locality:
Guatemala. SVL to 80 mm. Central plateau
of Chiapas, Mexico, to central Guatemala,
1200-2500 m elevation in pine and cloud
forest.
Sceloporus teapensis GUNTHER 1890, Reptilia
and Batrachia. Biologia Centrali-Americana:
75; type locality: Teapa, Tabasco, Mexico. SVL
to 70 mm. From Xalapa, Veracruz and north- Fig. 424. Sceloporus smaragdinus (Chiapas,
eastern Oaxaca, Mexico, across northern Mexico). Photo: A. Ramirez V.
Guatemala as far as Belize, sea level to 1000
m elevation in savanna and dry forest.
Sceloporus variabilis WIEGMANN 1834,
Herpetologica Mexicana: 51; type locality:
Mexico. SVL to 74 mm. From Texas, USA
along the Caribbean side of Mexico into
northwestern Costa Rica, sea level to 1700 m
elevation (to 2500 m elevation according to
JOHNSON 1989) in savanna and dry forest. In
Central America, only the nominate subspe-
cies occurs (of which olloporus H. SMITH 1937
is a synonym).

Fig. 425. Sceloporus smithi (near Mixtequilla,

Oaxaca, Mexico). Photo: G. Kohler

154
 Iguanidae

Sceloporus

.6. S. carinatus
.a. S. edwardtaylori
• S. chrysostictus
• S. squamosus
• S. siniferus

Fig. 426. Sceloporus squamosus (Santa Lucia, Sceloporus

Intibuca, Honduras). Photo: G. Kohler

.a. S. cozumelae
• S. lundelli
• S. variabilis
• S. teapensis
• S. smithi

S. acanthinus
S. internasalis
• S. lunaei
• S. malachiticus
S. smaragdinus
• S. taeniocnemis

Fig. 427. Sceloporus teapensis (Palenque,

Chiapas, Mexico). Photo: P. Heimes
Sceloporus

Sceloporus serrifer
• Sceloporus melanorhinus
Fig. 428. Sceloporus uariabilis (Canon de
Sumidero, Chiapas, Mexico). Photo: G. Kohler
155
 Iguanidae

Key to Sceloporus superciliaries

1 a Postfemoral pocket present (Fig. 431)
(variabilis group) ................................ 2
b Without postfemoral pocket ..................... 5
2 a Males uniform pale below, no colorful belly • •• loreal
patches; endemic to the north of the Yuca- mental
tan Peninsula ..... Sceloporus cozumelae
b Males with obvious ventral markings, con-
sisting of two large purple-colored belly
patches, usually with black borders ........ 3
3 a 49 or fewer dorsal scales from posterior of Fig. 429. Lateral view of head in Sceloporus
head to base of tail; 7-9 rows of enlarged malachiticus.
dorsal scales in the pelvic area; scales in
the inguinal region relatively large
........................... Sceloporus teapensis frontonasals supraoculars
b 50 or more dorsal scales from posterior of
head to base of tail; 10 or more rows of
enlarged dorsal scales in the pelvic area;
scales in inguinal region relatively small 4
4 a Ground color very dark, nearly black, with
very obvious yellow or white longitudinal
stripes (Fig. 432a), which are 2-2.5 scales rostral
wide; without obvious dark diagonal bars
on dorsum and body sides; keels of the dor-
sal scales form distinct fine combs; hatch-
lings with pink colored tail .................... .
. . . . . . . . . . . . . . . . . ... . . . . . . . . . .. Sceloporus smithi
b Ground color grey to brown, with less
distinct pale longitudinal stripes (Fig. Fig. 430. Dorsal view of head in Sceloporus
432b), which are 1.5-2 scales wide; distinct chrysostictus.
dark diagonal bars on dorsum and body
sides present; keels of the dorsal scales do
not form distinct fine combs; hatchlings
with grey brown tail .......................... .
.............................. Sceloporus variabilis b 12-17 femoral pores; 42-57 dorsal scales
from posterior of head to base of tail
5 a Males uniform pale below, no colorful belly ........... ..... ...... Sceloporus chrysostictus
patches (siniferus and chrysostictus
groups) .................................. ............ 6 9 a A distinct dark neckband present ......... 10
b Males with conspicuous colored belly pat- b No dark neckband present (spinosus
ches; if without colorful belly patches group) .................................................... 16
(Sceloporus edwardtaylori, only on the
Isthmus of Tehuantepec), then large, very lOa Dark neckband on both sides pale grey or
blue bordered (torquatus-group)
spiky-scaled species ............................... 9
............ .. ............... Sceloporus serrifer
6 a 1 canthal (Fig. 433b) ............ ................... 7 b Dark neckband not pale bordered (formo-
b 2 canthals (Fig. 433a) .............................. 8 sus-group) .......................................... 11
7 a 9-12 femoral pores; 38-44 dorsal scales lla In the snout area (between the nasals and
from posterior of head to base of tail between rostral and median frontonasal)
.......................... Sceloporus carinatus many small scales (Fig. 434a); usually two
b 3-7 femoral pores; 28-37 dorsal scales from scales between nasal and rostral
posterior of head to base of tail ........... . .......................... Sceloporus internasalis
. . . . . . ......... ....... .. . Sceloporus squamosus b In the snout region, a few large scales (Fig.
434b); usually 1 scale between nasal and
8 a 3-11 femoral pores; 31-46 dorsal scales rostral ..................................................... 12
from posterior of head to base of tail
............................ Sceloporus siniferus 12a Usually, only a single canthal ................ 13
b Usually two canthals ............................. 15

156
 Iguanidae

13a Supraoculars very large and arranged in a

row, usually one or more in contact with
medial head scales; 31-38 dorsal scales
from posterior of head to base of tail; low-
land species ............... Sceloporus lunaei
b Supraoculars smaller and arranged in two
Fig. 431. Post- rows, seldom in contact with medial head
fermoral pocket. scales; 30-51 dorsal scales from posterior
After SMITH (1939a). of head to base of tail; highland species
.. ................. ................. ....... ... .. ..... .... 14
14a 30-39 dorsal scales from posterior of head
to base of tail Sceloporus malachiticus
b 39-51 dorsal scales from posterior of head
to base of tail . Sceloporus smaragdinus
15a 34-39 dorsal scales from posterior of head
to base of tail .. .. Sceloporus acanthinus
a. S. smithi b. S. variabi/is b 39-47 dorsal scales from posterior of head
to base of tail Sceloporus taeniocnemis
Fig. 432. Dorsal coloration. Photos: G. Kohler
16a Males without colored ventral spots
............ ......... Sceloporus edwardtaylori
b Males with conspicuous colored ventral
spot ........ ..... .................. ....... ................. 17
17a 17 -24 femoral pores .. ........ .... ........ .... ..
1
...
~

......... ......... ... Sceloporus melanorhinus

b 9-12 femoral pores Sceloporus lundelli

Further Reading
a. S. siniferus b. S. squamosus H . SMITH 1939a (Revision of Mexican and
Fig. 433. Lateral view of head (canthals orange). Central American Sceloporus species); SITES &
DIXON 1982, H. SMITH et al. 1993, KOHLER
1994c, H. SMITH et al. 1995b (Sceloporus varia-
bilis group); H. SMITH & BUMZAHEM 1955,
STUART 1971, KOHLER 1990, 1993d, H. SMITH &
PEREZ-HIGAREDA 1992 (Sceloporus formosus
group); OLSON 1987, KOHLER et al. 1998;
KOHLER & HEIMES 2002 (summary of biology,
care and breeding)

a. S. internasalis b. S. malachiticus
Fig. 434. Dorsal view of head in Sceloporus.

157
 lguanidae

Urosaurus
The delicate, small lizards of the genus
Urosaurus are distributed predominantly
in the US and Mexico. Of the nine
Urosaurus species, only one reaches the
northern portion of Central America,
namely U. bicarinatus (MITTLEMAN 1942,
WIENS 1993). Phylogenetically, the
Urosaurus species are closely related to
the Spiny Lizards of the genus Sceloporus
(ETHERIDGE & DE QUEIROZ 1988).
Urosaurus bicarinatus is an inconspicuous
pale grey colored bush and tree dweller in Fig. 435. Urosaurus bicarinatus (near Mi.xte-
dry areas, usually found on the outer, thin- quilla, Oaxaca, Mexiko). Photo: G. Kohler
ner twigs. Thus, thorny bushes are a pre-
ferred habitat. The chest of the male is
adorned with two bright blue spots, while
the throat is yellowish in color. The threat
behavior of these territorial lizards con-
sists of pushup-like movements, in which
the whole body is pushed vertically
upright and back down again. The diurnal
Urosaurus feeds on small invertebrates
(insects and spiders, among others). For
egg laying, which, in Chiapas, occurs pri-
marily in June, the females come down
from the trees in order to bury their
• Urosaurus bicarinatus
clutches of around ten eggs (approx. 10x7
mm in size) (ALVAREZ DEL TORO 1960).

Urosaurus bicarinatus (DUMERIL 1858), Arch.

Mus. Hist. Nat. Paris 8: 549; type locality:
Mexico. SVL to 58 mm. Along the Pacific
Coast of Mexico from southern Sonora as far
as Chiapas, sea level to 1200 m elevation. The
subspecies Urosaurus b. anonymorphus
MITILEMAN 1940 (eastern Oaxaca northward)
and Urosaurus b. spinosus BuMZAHEM &
SMITH 1954 (Chiapas) occur in Central
America.

Further Reading
MITILEMAN 1942, WIENS 1993

Fig. 436. Urosaurus bicarinatus (near Mi.xte-

quilla, Oaxaca, Mexiko). Photo: G. Kohler
158
 Scincidae

Scincidae
From the large family of skinks, only five
genera with a total of nine species occur in
Central America. Most of the remaining
species are found in the Old World. Skinks
typically have an elongated head, body
and tail, with relatively short legs and
very smooth scales. Immediately under
the scales are small dermal bones (osteo-
derms).

Key to Skink Genera

1 a Without supranasals, therefore rostral in
complete contact with first unpaired scale Fig. 438. Mabuya unimarginata (Isla de
on dorsal surface of snout (Fig. 439a,b) .. 2 Guanaja, Honduras). Photo: E. Kohler
b Supranasals present, therefore rostral
entirely or partially separated from first
unpaired scale on dorsal surface of snout
(Fig. 439c,d) .. ..... .. ........ .... .................... 3
2 a One frontoparietal (Fig. 439b) ............... .
. . . .... . . . . . . . .......... .. .... . .. .. Sphenomorphus
b Two frontoparietals (Fig. 439a) ............... .
.............. .. .... ............... ...... ... ... Scincella
3 a Scales in temporal region same size as a. Scincella b. Sphenomorphus
body scales (Fig. 437b) ....... ......... Mabuya
b Scales in temporal region conspicuously
enlarged (Fig. 437a) .... .... ..................... .. 4
4 a Scales along the dorsomedial line not
obviously larger than paramedian dorsals;
with three pale longitudinal stripes, the
middle of which branches on the middle of c. Mabuya d. Eumeces
the head (Fig. 440a) ................... Eumeces
b Scales along the dorsomedial line obvious- Fig. 439. Dorsal head scalation in skinks
ly larger than paramedian dorsals; mark- (supranasals orange, frontoparietals brown).
ings different than those above (Fig.
440b,c) ..... ...... ... .. ... ... .. ..... Mesoscincus

a. Eumeces b. Mabuya a. E. sumichrasti b. M. managuae c. M. schwartzei

Fig. 437. Lateral view of head (temporals pale Fig. 440. Dorsal coloration in Eumeces and
brown). Mesoscincus.
159
 Scincidae

Eumeces

Based on a phylogenetic analysis of spe-

cies traditionally grouped under the gen-
eric name Eumeces, GRIFFITH et al. (2000)
provided evidence that this assemblage
was not a natural grouping. Consequently,
they split Eumeces sensu lato into several
genera. According to this new class-
ification, one Central American species (E.
sumichrasti) remains in the genus
Eumeces, whereas two species (M. mana-
guae and M. schwartzei) are assigned to
the genus Mesoscincus . Fig. 441. Juvenile Eumeces sumichrasti from
Honduras. Photo: J . R. McCranie
A clutch of 11 eggs from E. sumichrasti
was discovered in a freshly fallen bread-
fruit tree in Belize (MILLER 1997). The
eggs (15xll mm, a bit less than 1 g in
weight) lay in a knothole, which would
have been about 20 m above the ground
before the tree had collapsed. They were
buried in a substrate mix of fallen bark
and other organic material. A female E.
sumichrasti (92 mm SVL) stayed by the
clutch, possibly indicating that the species
practices brood care, as is the case with
other species of this genus (FITCH 1954,
VITT & COOPER 1985, SOMMA & FAWCETT
1989). Freshly hatched E. sumichrasti Eumeces sumichrasti
have a total length of 70 mm with a SVL
of 26 mm and a weight of 0.7 g (MILLER
1997). While adult E. sumichrasti have a
pale brown ground color with black longi-
tudinal stripes, the young are somewhat
more brightly colored. The latter have a
bright blue tail, and several orange spots
on the head, as well as three orange to yel-
low longitudinal stripes on the body
(MILLER 1997).

Eumeces sumichrasti (COPE 1867), Proc.

Acad. Nat. Sci. Philadelphia 1866: 321; type Mabuya unimarginata
locality: Orizaba, Veracruz, Mexico. SVL to 96
mm. Central Veracruz, Mexico, to northern
Honduras; also northeastern portion ofYucatan
Peninsula, sea level to 1000 m elevation.

Further Reading
TAYLOR 1935, GRIFFITH et al. 2000

160
 Scincidae

Mabuya

While earlier authors (e.g., TAYLOR 1956)

proposed that, in Costa Rica alone, three
species (alliacea, brachypoda, unimargi-
nata) of the genus Mabuya occurred,
recent authors conclude that only one spe-
cies, M. unimarginata, is distributed all
across Central America (SAVAGE & VILLA
1986, VILLA et al. 1988). However, the geo-
graphic variation of Central and South
American representatives of this genus
has hardly been examined. Therefore, the
current taxonomic division can only be Fig. 443. Mabuya unimarginata (Bartola, Rio
considered temporary. This insectivorous San Juan, Nicaragua). Photo: G. Kohler
skink. occurs commonly in many locations
in the lowlands of Central America. These
animals are generally found on the
ground, but they will climb tree trunks in
order to bask. Mabuya unimarginata is a
livebearer, with litter size varying from 2
to 7 young.

Mabuya unimarginata COPE 1862b, Proc.

Acad. Nat. Sci. Philadelphia 14: 187; type
locality: Panama. SVL to 90 mm. Mexico to
Panama, sea level to 1500 m elevation.

Further Reading
TAYLOR 1956, Webb 1958
Fig. 444. Mabuya unimarginata (Volcan
Maderas, Isla Ometepe, Nicaragua).
Photo: G. Kohler

Fig. 442. Mabuya unimarginata (near Fig. 445. Mabuya unimarginata (Ruinas de
Guanagasapa, Guatemala). Photo: G. Kohler Mayapan, Yucatan, Mexico). Photo: G. Kohler
161
 Scincidae
Fig. 446. Mesoscincus managuae (Morgans
Rock, Nicaragua). Photo: J . Sunyer
Mesoscincus
The genus Mesoscincus was recently pro-
posed for three species of skinks formerly
assigned to the diverse genus Eumeces
(GRIFFITH et al. 2000). Two of the species
included in Mesoscincus occur in Central
America, M. managuae and M. schwartzei.
These skinks are ground dwellers in dry
and seasonal rain forest, where they can
be seen basking on roots and fallen logs. In
July 1999, I discovered an adult specimen
of M. managuae at the foot of the Volcan
Cosiguina, Nicaragua, under a musty tree
trunk. Sometimes these lizards are also
Mesoscincus schwartzei
• Mesoscincus managuae
162
found under loose bark on upright tree
trunks. The Mesoscincus species are shy,
secretive lizards that are rarely encounter-
ed in nature.
Mesoscincus managuae (DUNN 1933c), Proc.
Biol. Soc. Washington 46: 67; type locality:
Aviation Field, Managua, Nicaragua. SVL to
117 mm. El Salvador to northwestern Costa,
near sea level to 530 m elevation in dry
forest.
Mesoscincus schwartzei (FISCHER 1885), Abh.
Naturwiss. Ver. Hamburg 8: 3; type locality:
small island in Laguna de Terminos,
Campeche; Mexico. SVL to 120 mm. Yucatan
Peninsula and bordering Tabasco, Mexico,
into central El Peten in Guatemala, sea level
to 250 m elevation.
Key to Mesoscincus
1 a With two wide pale longitudinal stripes,
which begin at the tip of snout and fuse
into a broad band in the anterior dorsal
area (Fig. 440c); 21 scales at midbody;
front and rear limbs touch, i.e., overlap,
when laid at the side of the body .......... ....
................... ... ... Mesoscincus schwartzei
b Dorsum brown with dark brown longitudi-
nal lines, which in some individuals break
up into spots (Fig. 440b); 17-19 scales at
midbody; front and rear limbs do not touch
when laid at the side of the body ............. .
.. .... ... ............... Mesoscincus managuae
Further Reading
TAYLOR 1935, 1956, CRUZ et al. 1979, REEDER
1990, GRIFFITH et al. 2000
Fig. 447. Mesoscincus managuae (Rio Sandillal,
Costa Rica). Photo: R. W. Van Devender
 Scincidae

Fig. 448. Mesoscincus schwartzei (19 km N

Temoz6n, Yucatan, Mexico).
Photo: Section of Herpetology, Natural History
Museum of Los Angeles County.
Fig. 449. Scincella gemmingeri (Hidalgo,
Mexico). Photo: G. Kohler

Scincella

According to the taxonomic division by

GREER (1974), one Central American skink
species, S. gemmingeri from the subfamily
Lygosominae, is included in the genus
Scincella, whereas the species assatus,
cherriei, incertus and rarus are placed in
the genus Sphenomorphus (MYERS &
DONNELLY 1991). According to HONDA et al.
(2003), the New World Scincella have a • Scincella gemmingeri
closer affinity with the New World
Sphenomorphus than with the Old World
Scincella. Scincella gemmingeri has 60-68
dorsal scales from the parietal to the base
of the tail (at the level of the posterior bor-
der of the thighs) and 25-28 longitudinal
rows of dorsal scales (H. SMITH 1949).

Scincella gemmingeri (COPE 1864), Proc.

Acad. Nat. Sci. Philadelphia 16: 180; type
locality: Orizaba, Veracruz, Mexico. SVL to 45
mm. Hidalgo, Veracruz and Oaxaca, Mexico,
200-2000 m elevation in pine forest.

Further Reading
TAYLOR 1937a, H. SMITH 1949, HONDA et al.
2003

Fig. 450. Scincella gemmingeri (Hidalgo,

Mexico). Photo: G. Kohler
163
 Scincidae

Sphenomorphus

The Central American herpetofauna in-

cludes three apparently closely-related,
small skink species from the subfamily
Lygosominae (assatus, cherriei, and incer-
tus) that, until the beginning of the 1970's,
were variously included in the genera
Lygosoma (e.g., STUART 1940), Scincella
(e.g., H . SMITH & TAYLOR 1950) or
Leiolopisma (e.g., TAYLOR 1956, STUART
1963, PETERS & DONOSO-BARROS 1970). In
his rev1s1on of genera related to Fig. 451. Sphenomorphus cherriei (Rio Sereno,
Leiolopisma, GREER (1974) placed the Chiriqui, Panama). Photo: G. Kohler
three above-named species in the genus
Sphenomorphus, a view which is widely
held today (FITCH 1983, VILLA et al. 1988).
Recently, MYERS & DONNELLY (1991)
described a fourth species from this genus
in Panama, S. rarus, that is known only
from a few specimens.

Whereas Sphenomorphus cherriei is a

diurnal ground dweller in the under-
growth of the rain forest, S. assatus is
mostly an inhabitant of subhumid areas.
The short legs are assisted in forward loco-
motion by snake-like movements of the
body, especially when increasing speed. At
air temperatures between 23 and 27 °C,
the animals are active primarily in the Fig. 452. Sphenomorphus assatus (near
morning, while they will hide from the Guanagasapa, Escuintla, Guatemala).
midday sun in cooler places. In the after- Photo: G. Kohler
noon, the agile Sphenomorphus will
undertake another period of activity
(FITCH 1983). Diet consists of small arthro-
pods (insects and spiders), which they
track primarily with their olfactory senses.
Large prey animals are preferred, and
even cannibalism is known among these
skinks (FITCH 1983). The species of the
genus Sphenomorphus reproduce ovipar-
ously, with females producing several
clutches per season, each with one to three
(seldom four) eggs. Whereas Sphenomo7-phus
in the constantly moist Caribbean low-
lands areas reproduce throughout the
year, populations at higher altitudes or
areas with distinct dry periods are highly Fig. 453. Hatchlings of Sphenomorphus cherriei
seasonal in their egg production. Thus, (Quintana Roo, Mexico). Photo: H. Bahena B.

164

FITCH (1983) reported that the reproduc-
tive activities of S. cherriei in the region of
Turrialba, Costa Rica, stagnate in the
months from November to April. With an
SVL of approx. 22 mm, the hatchlings of S.
cherriei are already 40 % of their adult
size. Within seven months, the young are
sexually mature at an SVL of 48 mm. The
average duration of a generation of S.
cherriei is approximately one year.
Countless predators (primarily snakes,
birds and mammals) prey upon these
small sk.inks. All Sphenomorphus species
will discard their tails if they are seized.
Sphenomorphus assatus (COPE 1864), Proc.
Acad. Nat. Sci. Philadelphia 16: 179; type
locality: Volcan Izalco, El Salvador. SVL to 50
mm. Colima, Mexico, to El Salvador and
southern Honduras (MCCRANIE & KOHLER
1999a), sea level to 1500 m elevation (to 2500
m elevation according to JOHNSON 1989).
Sphenomorphus cherriei (COPE 1893), Proc.
Amer. Phil. Soc. 31: 340; type locality: El
Palmar, Costa Rica. SVL to 66 mm. Tabasco,
Mexico, to western Panama, from sea level to
1300 m elevation in lowland and premontane
rain forest.
Sphenomorphus incertus (STIJART 1940), Occ.
Pap. Mus. Zool., Univ. Michigan 421: 10; type
locality: Volcan Tajumulco, Guatemala. SVL
to 67 mm. Guatemala and Honduras.
Sphenomorphus rarus MYERS & DONNELLY
1991, Amer. Mus. Nov. 3027: 2; type locality:
SW headwaters of Rio Guabo, 780 m eleva-
tion, 8°47'N, 82°1l'W, Bocas del Toro,
Panama. SVL to 52 mm. Known only from the
type locality.
Fig. 454 Sphenomorphus rarus (holotype).
Photo: C. W. Myers
Scincidae
Key to Sphenomorphus
1 a Without prefrontal scale, therefore,
contact occurs between frontonasal and
frontal ........ ....... Sphenomorphus rarus
b Prefrontal scale present, frontonasal not
in contact with frontal ............................. 2
2 a 30-36 scales at midbody; front and rear
extremities touch when laid at side of
body; tail brown in adult specimens, blue
in juveniles .... Sphenomorphus cherriei
b 24-31 Scales at midbody; front and rear
extremities do not touch when laid at side
of body; tail red brown in adult specimens,
intense red in juveniles ...................... .. .. 3
3 a 67-77 dorsal scales between parietal and
base of tail (at level of posterior border of
thighs); 27-31 scales at midbody ............... .
. ..... ................ Sphenomorphus assatus
b Fewer than 65 dorsal scales between
parietal and base of tail (at level of poster- Ill
ior border of thighs); 26 or fewer scales at
midbody ........ Sphenomorphus incertus 1...
N
~
Further Reading
TAYLOR 1937a, SroART 1940, H. SMITH 1946,
1951, GREER 1974, WILSON & MCCRA.NIE 1994,
MCCRANIE & KOHLER 1999a, TOWNSEND 2005
• Sphenomorphus assatus
• Sphenomorphus incertus
165
 Teiidae

Teiidae
The lizards of the family Teiidae, including
ten genera with approximately 110 species,
are widespread in temperate, subtropical,
and tropical areas of the USA, Mexico,
Central America, South America, and the
Antilles. Most species are active terrestrial
lizards, and all are diurnal. All teiids are
oviparous; parthenogenesis is prevalent
among Aspidoscelis.

Key to genera of Teiidae Fig. 457. Ameiua undulata (El Imposible

1 a One row of greatly enlarged scales on the National Park, Ahuachapan, El Salvador).
outside of the upper arm (Fig. 456a); dor- Photo: G. Kohler
sal pattern with five or fewer longitudinal
stripes; a fleshy sheath enclosing base of
tongue (Fig. 455a) ................... .. ... Ameiva

f•
b Three or more rows of moderately en-
larged scales on the outside of the upper
arm (Fig. 456b); dorsal pattern usually
with six or more pale longitudinal stripes,
or with pale spots; no fleshy sheath en-
closing base of tongue (Fig. 455b) .. ........ 2
2 a One interparietal and 4 parietals (Fig.
4 71a) ...... .. ... .... .... . .... .. Cnemidophorus
b One interparietal and 2 parietals (Fig.
471b,c) ....... .... .... ................. Aspidoscelis

Fig. 458. Ameiua quadrilineata (Rio Sereno,

Panama). Photo: G. Kohler

a. Ameiva b. Aspidosce/is

Fig. 455. Floor of mouth. Note tongue sheath

(arrow) in Ameiua.

~-:_~~,
~ b. Cnemidophorus

Fig. 459. Ameiua leptophrys (Manuel Antonio,

Fig. 456. Foreleg (enlarged scales shaded). Costa Rica). Photo: D. M. Dehling

166

Fig. 460. Ameiva undulata (San Julian,
Suchitepequez, Guatemala). Photo: G. Kohler
Fig. 461. Ameiva ameiva. Photo: R. D. Bartlett
Fig. 462. Ameiva festiva (Nusagandi, Comarca
de San Blas, Panama). Photo: G. Kohler
Teiidae
Ameiva
In Central America, there are six ameiva
species, of which A. festiva and A. undula-
ta are the most widely distributed
(ECHTERNACHT 1971). Ameivas are diurnal,
very agile lizards that are common in
many places, primarily in open secondary
forests along the paths and in clearings.
The sunlight and heat loving ameivas keep
their body temperature around 37 °C during
their active phase (HIRTH 1963, HILLMAN
1969, FITCH 1973a). Cooler and rainy days
are spent in burrows in the ground.
Ameivas are tenacious hunters, who will
wander around their territory looking for
prey, scratching in the leaves, digging in
the ground, and searching in the under-
growth and under bark, consuming any
creature they can overpower, primarily
insects and arachnids. The lizards are con-
stantly flicking their tongues, checking
scents in their surroundings. Although
these lizards are ground dwellers, they
will also climb tree trunks and low vegeta-
tion to bask or to reach prey. Important
predators of the ameivas are the diurnal,
quick colubrids, including Conophis linea-
tus, Mastigodryas melanolomus, Drymo-
bius margaritiferus, and Masticophis men-
tovarius.
The Central American ameiva species pro-
duce several clutches of eggs per year, with
clutch size dependent upon species and
the size of the female (A ameiva 1-9, A.
festiva 2-4, A. undulata l-7; ECHTERNACHT
1983, KOHLER 1997). At 28-30 °C, the incu-
bation period for the eggs of A. ameiva
lasts 129-137 days. The juveniles of A fest-
iva, A. quadrilineata, and A. undulata
have pale blue-colored tails, just like the
juvenile whiptail Aspidoscelis deppii . Up
to three ameiva species can occur in a
single area (sympatric) (e.g., the species
festiva, leptophrys, and quadrilineata in
southeastern Costa Rica), with each
making use of a slightly different ecologi-
cal niche to reduce the competition
(ECHTERNACHT 1971, 1983, FITCH 1973a).
167
 Teiidae

Ameiva ameiva (LINNAEUS 1758), Systema

Naturae, ed. 10: 202; type locality: Brazil.
SVL to 197 mm (males) and 157 mm (fe-
males). Costa Rica Panama as well as large
parts of tropical South America, sea level to
700 m elevation. In Panama, the subspecies
Ameiva a. praesignis (BAIRD & GIB.ARD 1852)
occurs.
Ameiva chaitzami STUART 1942a, Proc. Biol.
Soc. Washington 55: 143; type locality: along
Cahab6n-Lanquin trail about 2 km N Finca
Canihor, about 38 km ENE Cohan, Alta
Verapaz, Guatemala. SVL to 85 mm (males) • Ameiva ameiva
and 75 mm (females). Eastern Chiapas,
Mexico, as well as eastern and central
Guatemala, 800-2000 m elevation.
Ameiva festiva (LICHTENSTEIN & VON MARTENS
1856), Nomenclator Reptilium et
Amphibiorum Musei Zoologici Berolinensis:
13; type locality: Veragua, Panama. SVL to
114 mm (males) and 129 mm (females).
Tabasco, Mexico, across all of Central
America as far as Colombia, sea level to 1200
m elevation. In Central America, three sub-
species occur: Ameiva f. festiva (Panama and
northern Colombia); Ameiva f. edwardsii
BocoURT 1873 (Isthmus of Tehuantepec to
Nicaragua); Ameiva f. occidentalis TAYLOR • Ameiva leptophrys C!. ..
1956 (Costa Rica).
Ameiva leptophrys COPE 1893, Proc. Amer.
Phil. Soc. 31: 341; type locality: Buenos Aires, • Ameiva quadrilineata
Costa Rica. SVL to 133 mm (males) and 129
mm (females). Southeastern Costa Rica to
eastern Panama, sea level to 700 m elevation.
Ameiva quadrilineata {HALLOWELL 1861),
Proc. Acad. Nat. Sci. Philadelphia 12: 483;
type locality: Nicaragua. SVL to 88 mm
(males) and 82 mm (females). Southeastern
Nicaragua to western Panama, sea level to
1050 m elevation.
Ameiva undulata (WIEGMANN 1834),
Herpetologica Mexicana: 27; type locality:
Mexico. SVL to 129 mm (males) and 111 mm
(females). Nayarit, Mexico, to Costa Rica, sea
level to 1500 m elevation. In Central
America, six subspecies occur: Ameiva u. gai-
geai SMITH & LAUFE 1946 (northern Yucatan
Peninsula), Ameiva u. hartwegi SMITH 1940
(Base of the Yucatan Peninsula and northern
Honduras), Ameiva u. miadis BARBOUR &
LOVERIDGE 1929a (Isla del Maiz Grande),
Ameiva u. parva BARBOUR & NOBLE 1915
(Pacific side of Central America), Ameiva u.
pulchra HALLOWELL 1861 (Honduras to Costa
Rica along the Caribbean side), Ameiva u.
thomasi SMITH & LAUFE 1946 (upper Rio
Grijalva Basin, Chiapas).
• Ameiva festiva

168
 Teiidae

Key to Ameiva
1 a 10-12 longitudinal rows of ventral scales;
no obvious enlarged throat scales (Fig. a. A. ameiva
463a), rather gradual increase in size
posteriorly; scales on the underside of the
neckband (mesoptychial scales) not
obviously enlarged ......... Ameiva ameiva
b 8 longitudinal rows of ventral scales; 2-5
throat scales enlarged; scales on the
underside of the neckband (mesoptychial b. A. festiva
scales) obviously enlarged (Fig. 463b-d) .. 2
2 a Anterior throat scales smaller than pos-
terior throat scales (Fig. 463c) ................. 3
b Anterior and posterior throat scales of
equal size (Fig. 463b,d) ......... ................... 4
3 a Parietals and frontoparietals separated by
one or more scales (Fig. 464a); scales at c. A. leptophrys
mid-throat greatly enlarged ................ .
................................... Ameiva leptophrys
b Parietals in contact with frontoparietals
(Fig. 464b); scales at mid-throat only
slightly enlarged Ameiva quadrilineata
4 a Scales at mid-throat greatly enlarged and d. A. quadrilineata
not arranged in a longitudinal row (Fig.
463b); a pale dorsomedial longitudinal
stripe present (may be absent in very
large specimens) ............. Ameiva festiva
b Scales at mid-throat greatly enlarged and
arranged in a longitudinal row or only
slightly enlarged and irregularly arranged
(Fig. 463d); without pale dorsomedial e. A. undulata
longitudinal stripe .... .............................. 5
5 a SVL to 85 mm (males) and 75 mm (fe-
males); paravertebral stripes very narrow;
dorsolateral spots in males fused with pale Fig. 463. Ventral view of head inAmeiva.
dorsolateral stripes ... Ameiva chaitzami
b SVL to 129 mm (males) and 111 mm
(females); paravertebral stripes wide; dor-
solateral spots in males, if present, not
fused with pale dorsolateral stripes ......... .
. . ............................... ... Ameiva undulata

a. A. leptophrys b. A. quadrilineata
Fig. 464. Dorsal view of head inAmeiva
(frontoparietals orange, parietals brown).

• Ameiva chaitzami
• Ameiva undulata
Further Reading
ECHTERNACIIT 1971.
169
 Teiidae

Aspidoscelis

Whiptails are the most common lizards in

the dry areas of Mexico and Central
America. REEDER et al. (2002) proposed a
new classification of the whiptails based
on a phylogenetic analysis. Because the
diverse assemblage of species that was
traditionally grouped under the generic
name Cnemidophorus was demonstrated
to be paraphyletic, REEDER et al. (2002)
split the group into several genera. The
lemniscatus group retained the generic
name Cnemidophorus, whereas the other
North American and Central American
species were transferred to the resurrec-
ted genus Aspidoscelis. The genus
Aspidoscelis contains at least 87 currently
recognized bisexual and unisexual taxa
and is distributed throughout most of
North America (except Canada), south Fig. 466. Aspidoscelis deppii (male above, fe-
male below) at Playa Tamarindo, Costa Rica.
through Mexico into lower Central
Photo: G. Kohler
America. In Central America, the genus is
represented by six species (DUELLMANN &
WELLMAN 1960, VILLA et al. 1988, LEE human observer, these lizards seem tre-
1996). mendously restless, as they are virtually
in constant motion. As heat-loving ani-
Whiptails are active hunters that will eat mals, the whiptails are out in the open pri-
· everything that they can overpower, most- marily on hot, sunny days, whereas they
ly insects and other arthropods. To the retreat to their burrows when the sky is
cloudy. Most of the 189 Aspidoscelis deppii
examined by FITCH (1973b) had body tem-
peratures from 29-42 °C. Aspidoscelis are
most active in the morning. Most of the
Aspidoscelis species produce 2-5 clutches
per year, each with 1-5 eggs, which they
bury superficially in the sand. In the dry
season, reproductive activity comes almost
entirely to a standstill, whereas the egg-
laying period begins with the onset of the
rainfall. The incubation period lasts from
60-80 days. Aspidoscelis deppii becomes
sexually mature at the age of 5-6 months
(ECHTERNACHT 1983). Worthy of note is the
fact that several Aspidoscelis species,
including the Central American A. cozu-
mela and A. rodecki, reproduce partheno-
Fig. 465. Aspidoscelis cozumela maslini (Banco genetically. In these two species, only
Chinchorro Reef, Quintana Roo, Mexico). females, which produce fertile eggs with-
Photo: H. Bahena B. out any previous copulation, are known.
170
 Teiidae

Aspidoscelis angusticeps (COPE 1877), Proc.

Arner. Phil. Soc. 17: 95; type locality: Yucatan,
Mexico. SVL to 115 mm. Yucatan Peninsula,
sea level to 100 m elevation. Two subspecies
are recognized: Aspidoscelis a. angusticeps
(northern part of the Yucatan Peninsula) and
Aspidoscelis a. petenensis BEARGIE & McCOY
1964 (base of the Yucatan Peninsula).
Aspidoscelis cozumela (GADOW 1906), Proc.
Zool. Soc. London 1906: 316; type locality: Isla
Cozumel, Quintana Roo, Mexico. SVL to 83
mm. Yucatan Peninsula, sea level to 100 m
elevation. Two subspecies are recognized:
Aspidoscelis c. cozumela (Isla Cozumel) and
Aspidoscelis c. maslini (FRITTS 1969)
(Yucatan Peninsula). Fig. 468. Aspidoscelis motaguae (male, nr. Sta.
Aspidoscelis deppii (WIEGMANN 1834), Cruz, Zacapa, Guatemala). Photo: G. Kohler
Herpetologica Mexicana: 28; type locality:
Mexico. SVL to 93 mm. Northern Veracruz
and Guerrero, Mexico to Costa Rica, sea level
to 1200 m elevation. Only the nominate sub-
species occurs in Central America.
Aspidoscelis guttata (WIEGMANN 1834),
Herpetologica Mexicana: 29; type locality:
Mexico. SVL to 145 mm. Mexico, Veracruz on
the Atlantic side and from Guerrero to
western Chiapas on the Pacific side, sea level
to 1200 m elevation.
Aspidoscelis motaguae (SACKETT 1941),
Notulae Naturae 77: 1; type locality: Motagua
River, Zacapa, Guatemala. SVL to 145 mm.
Oaxaca, Mexico, to El Salvador and
Honduras, 500-1200 m elevation.
Aspidoscelis rodecki (McCoy & MAsLIN 1962),
Copeia 1962: 620; type locality: Isla Mujeres,
Quintana Roo, Mexico. SVL to 70 mm. The Fig. 469. Aspidoscelis motaguae (male, nr. Sta.
northeastern portion of the Yucatan Cruz, Zacapa, Guatemala). Photo: G. Kohler
Peninsula, sea level to 50 m elevation.

Fig. 467. Aspidoscelis angusticeps. Fig. 470. Aspidoscelis deppii (male right,
Photo: R. Cedeno female left) Santa Rosa National Park, Costa
Rica. Photo: D. M. Dehling
171
 Teiidae

Key to Aspidoscelis
1 a 4 supraoculars ...... .. .............. ... ............. 2
b 3 supraoculars ...... .................................. 3
2 a Dorsum of juveniles with six pale
longitudinal stripes on a dark brown back-
ground; in adults (SVL > 100 mm), dorsal
stripes usually completely replaced by
spots ................. A.spidoscelis motaguae
b Dorsum of juveniles with six or seven pale
longitudinal stripes on a black back-
ground; adult pattern variable, but some
trace of striping usually retained ......... .
....................... A.spidoscelis angusticeps Fig. 4 72. Aspidoscelis c. cozumela (Isla
Cozumel, Mexico). Photo: R. Cedeiio
3 a Frontoparietals divided from parietals by
one or more scales .................................... 4
b Frontoparietals in contact with parietals
................................................................... 5 Aspidoscelis

4 a Dorsal pattern with indistinct para-

vertebral stripes, discontinuous or absent
on the posterior body half ........................ .
. . . . . . . . ................. .. .. A.spidoscelis rodecki
b Dorsal pattern with four distinct contin-
uous paravertebral stripes ... ......... ...... ... .
. . . . . . .. . . . ........ ....... A.spidoscelis cozumela
5 a Fewer than 140 scales around midbody,
not counting ventral scales .................... . ""' A. cozumela
............................... A.spidoscelis guttata • A. deppii
b More than 165 scales around midbody, not
counting ventral scales ........................ .
................................. A.spidoscelis deppii Aspidoscelis

""'A.rodecki
• A. angusticeps
• A. motaguae

a. C. lemniscatus b. A. motaguae c. A. deppii

Fig. 471. Dorsal head scalation (supraoculars

orange; parietals brown) in Aspidoscelis and
Cnemidophorus.

Further Reading
BURT 1931, DUELLMAN & WELLMAN 1960,
DUELLMAN & ZWEIFEL 1962, MAsLIN & SECOY
Aspidoscelis guttata
1986, WRIGHT & VITT 1993, REEDER et al. 2002

172
 Teiidae

Fig. 473. Adult male ofCnemidophorus Fig. 4 74. Adult female of Cnemidophorus
lemniscatus (Isla de Utila, Honduras). lemniscatus (Isla de Utila, Honduras).
Photo: G. Kohler Photo: G. Kohler

Cnemidophorus Cnemidophorus lemniscatus (LINNAEUS

1758), Systema Naturae, ed. 10: 209; type
locality: "Guinea" [probably in error for
REEDER et al. (2000) proposed a new Guiana]. SVL to 104 mm. Central America
classification for the whiptail lizards and from Guatemala as far as northern South
transferred the majority of species from America east of the Andes, including many
offshore islands, as well as various Caribbean
Cnemidophorus to Aspidoscelis. As cur- islands (e.g., Trinidad and Tobago), sea level to
rently recognized, C. lemniscatus is the at least 1000 m elevation. Only the nominate
only Central American representative of subspecies occurs in Central America.
the genus Cnemidophorus.
Further Reading
Cnemidophorus lemniscatus is a typical KOHLER 1996d, COLE & DESSAUER 1993, Wright
diurnal inhabitant of the open coastal & Vitt 1993, REEDER et al. 2002, Mo.ncA et al.
areas, but will also advance inland along 2003
large rivers, hardly ever penetrating the
forest. These active lizards are often seen
in great numbers. They run a bit, stop ab-
ruptly and execute rapid turning motions
with their forelegs, which has earned them
the name "Shaky-Paw" amongst the inha-
bitants of the Islas de la Bahia, Honduras
(WILSON & HAHN 1973, KOHLER 1996d,
1998f). Although Cnemidophorus lemnis-
catus feed primarily on insects and other
arthropods, juveniles have also been
observed eating cactus blossoms (Opuntia
wentiana) (MIJARES-URRUTIA et al. 1997).
C. lemniscatus produce several clutches
per year, each with 1-5 eggs, which hatch
after about two months (KOHI.ER 1997).
173
 Xantusiidae

Xantusiidae
The family Xantusiidae consists of three
genera (Xantusia, Lepidophyma, Crico-
saura), with only Lepidophyma occuring
in Central America (BEZ¥ & CAMARILLO
2002). The distribution ranges of most
night lizards are extremely fragmented,
and populations are often isolated from
one another by hundreds of kilometers.
Such disjunct distribution has been taken
as an indication of the great geological age
of the group (BEZ¥ 1972).

Fig. 475. Lepidophyma smithii (El Refugio,

Ahuachapan, El Salvador). Photo: G. Kohler
Lepidophyma

The 17 species of the genus Lepidophyma

are distributed from Nuevo Leon in north-
eastern Mexico to as far south as Panama.
They are bizarre-looking mid-size lizards,
which are secretive ground-dwellers that
are seldom observed outside their hiding
places (beneath decaying tree stumps and
logs, in rock crevices and caves). They
have no eyelids, rather having, similar to
many gecko species, so-called "spectacles".
The sides of the body are decorated with
countless, irregularly arranged tubercular
scales, whereas the tail is equipped with
whorls of slightly enlarged scales.

During the day, these lizards hide under

rocks and fallen logs, which provide a Fig. 476. Lepidophyma mayae (Huehuetenan-
moist microclimate. These nocturnal ani- go, Guatemala). Photo: J. A. Campbell
mals are sit-and-wait predators whose
prey consists mostly of insects and spi-
ders. The remarkable thing about this
genus is the livebearing method of repro-
duction. There are also populations that • L. tuxtlae
"" L. chicoasense
consist entirely of females that reproduce v L. lipetzi
parthenogenetically. During the months • L. mayae
from April to July, the females of L. fiaui- • L. reticulatum
maculatum bear 2-5 (seldom as many as
8) young, which have a SVL of 35-38 mm
at birth (ALVAREZ DEL TORO 1960, TELFORD
& CAMPBELL 1970).

Lepidophyma

174
 Xantusiidae

Lepidophyma chicoasense ALVAREZ & Key to Lepidophyma

VALENTIN 1988, Anales de la Escuela 1 a 16 or more divided lamellae under the 4th
Nacional de Ciencia Biologia Mexico 32: 125; toe ... ...... .... Lepidophyma chicoasense
type locality: Canon de Sumidero, 16.3 km N, b 15 or fewer divided lamellae under the 4th
1.6 km E Tuxtla Gutierrez, 600 m, Chiapas, toe ................................ ........ .......... ........... 2
Mexico. SVL to 78 mm. Known only from the
type locality. 2 a Paravertebral scale row relatively homo-
geneous, composed of tubercular scales
Lepidophyma fiavimaculatum A. DUMERIL that are similar in size and nearly contin-
1851, Cat. Meth. Coll. Rept. Mus. Paris: 137; uous ................ Lepidophyma tuxtlae
type locality: El Peten, Guatemala. SVL to 95
mm. Veracruz, Mexico, along the Caribbean b Paravertebral scale row heterogeneous,
composed of tubercular scales of various
side of Central America as far south as the sizes interspersed with small granular
Panama Canal, sea level to 750 m elevation.
scales ........................................................ 3
Lepidophyma lipetzi SMITH & ALVAREZ DEL 3 a Parietal eye not visible in adults (Fig.
TORO 1977, J. Herpetol. 11: 37; type locality:
Lago de Mal Paso, headwaters of Rio de la 477a), occasionally present in juveniles
Venta, 30 km N Cintalapa (straight line), with a SVL less than 75 mm ....... .
Chiapas, Mexico. SVL to 55 mm. Known only .............................. Lepidophyma smithii
from the type locality. b Parietal eye visible (Fig. 477b) ................ 4
Lepidophyma mayae BEZ¥ 1973, Los Angeles
County Mus. Contrib. Sci. 239: 1; type locali-
ty: near Chinaja, 140 m elevation, Alta
Verapaz, Guatemala. SVL to 65 mm.
Southern El Peten and the lower slopes of the
Sierra de los Cuchumatanes and the Sierra
Xucaneb (Departments of Huehuetenango
and Alta Verapaz), Guatemala; also Maya
Mountains of Belize, 100-300 m elevation.
Lepidophyma reticulatum TAYLOR 1955, Univ.
Kansas Sci. Bull. 37: 551; type locality: Agua
Buena, Puntarenas, Costa Rica. SVL to 92
mm. Pacific side of Costa Rica, sea level to
1250 m elevation.
a. L. smithii b. L. navimacu/atum
Lepidophyma smithii BOCOURT 1876, Journ.
Zool. Paris 5: 402; type locality: Tehuantepec
Fig. 477. Dorsal head scalation in two species
and western Guatemala. SVL to 81 mm.
Guerrero, Mexico, along the Pacific side of of Lepidophyma.
Central America as far south as El Salvador,
sea level to 1200 m elevation.
Lepidophyma tuxtlae WERLER & SHANNON
1957, Herpetologica 13: 119; type locality:
lower slopes ofVolcan San Martin, 2500 feet,
Veracruz, Mexico. SVL to 97 mm. Isthmus of
Tehuantepec (discontinuously from the
Sierra de las Tuxtlas, Veracruz, Mexico, to the
El Ocote region, Chiapas); sea level to 1500 m
elevation.

Fig. 478. Lepidophyma fiavimaculatum (El

Ocote region, Chiapas, Mexico).
Photo: G. Kohler
175
 Xenosauridae
a. L. mayae b. L. flavimaculatum
Fig. 479. Lateral head scalation in two species
of Lepidophyma. Scales between 2nd postorbi-
tal supralabial and postocular orange.
4 a Dorsal scales (occiput to base of tail) 179
or fewer; more than 10 longitudinal rows
of ventrals on posterior half of body
........... ................... Lepidophyma lipetzi
b Dorsal scales usually 180 or more; if fewer
than 180, 10 or fewer longitudinal rows of
ventrals on posterior half of body
................................................................... 5
5 a 0-1 Scales between second postorbital
supralabial and postoculars (Fig. 4 79a);
lateral tubercular scales arranged in 33-
45 rows .......... ....... Lepidophyma mayae
b 2 or more scales between second post-
orbital supralabial and postocular scales
(Fig. 479b); lateral tubercular scales
arranged in 24-32 rows ............................ 6
6 a Throat region with obvious dark reticular
markings ..... Lepidophyma reticulatum
b Throat region without obvious dark retic-
ular markings, although some diffuse
shading may be present ............... ..... ... .. .
.. ........... Lepidophyma flavimaculatum
Further Reading
TELFORD & CAMPBELL 1970, BEZV 1972, 1984,
1989, BEZV & CAMARILLO 2002
Fig. 480. Xerwsaurus grandis (near Cordoba,
Veracruz, Mexico). Photo: R. E. Ballinger
176
Xenosauridae
The family of knob-scaled lizards has a
fragmented distribution in Mexico and the
northern portion of Central America, as
well as in China, which shows it to be a
relictual group, known from fossils since
the Upper Cretaceous.
Xenosaurus
In Central America, only one knob-scaled
lizard species occurs, Xenosaurus grandis.
The ability to autotomize and regenerate
the tail is not present in the species of the
genus Xenosaurus. The typical habitat of
Xenosaurus grandis is open rocky land
with hiding places in the form of rock crev-
ices. Actually, all the X grandis observed
by BALLINGER et al. (1995) in Veracruz,
Mexico, were in rock crevices; none were
found in the open. Only one knob-scaled
lizard lives in each crevice. Social rela-
tionships and interactions could not be
determined in field studies, which leaves
the suspicion that these animals are very
solitary. Horizontal rock crevices are pre-
ferred over vertical ones (BALLINGER et al.
1995). The assumption in earlier literature
that knob-scaled lizards are nocturnal
(KING & THOMPSON 1968) could not be sub-
stantiated by more recent field studies. At
night, all of the observed X grandis slept
 Xenosauridae

Xenosaurus grandis (GRAY 1856), Ann. Mag.

Nat. Hist. (2) 18: 270; type locality: Cordoba,
Veracruz, Mexico. SVL to 116 mm. Disjunct
populations from central Veracruz and
Oaxaca, Mexico, to Guatemala, 750-1800 m
elevation. Two subspecies occur in Central
America: Xenosaurus g. grandis (Veracruz
and Oaxaca) and Xenosaurus g. rackhami
STUART 1941b (Chiapas and Guatemala).

Fig. 481. Xenosaurus grandis.

Photo: R. D. Bartlett

with their eyes closed and did not react to

light stimulation (BALLINGER et al. 1995).
The first activities (e.g., opening the eyes,
looking out of the rock crevice) were deter-
mined to take place about two hours after
sunrise. Xenosaurus grandis belongs to
the thigmothermic lizards, which make
use of the heat in the surrounding sub-
strate, but not the heat of the sun's rays.
Fig. 482. Xenosaurus grandis (El Ocote region,
The body temperature of active knob-scaled
Chiapas, Mexico). Photo: A. Ramirez V.
lizards corresponds to the temperature of
the surrounding rock and lies between
20.0-25.8°C (average 22. 7°C); (BALLINGER
et al. 1995). In general, these lizards are
not shy and seem almost lethargic. The
radius of activity is reported to be less
than 1 m (ALVAREZ DEL TORO 1960). If a
knob-scaled lizard is grabbed, it will spray
a considerable amount of fluid from the
cloacal bladder as a defense mechanism,
similar to many frogs and toads. Knob-
scaled lizards feed primarily on many
types of insects, but will occasionally eat
small lizards (PRESCH 1981, BALLINGER et
al. 1995). Xenosaurus grandis reproduces
ovoviviparously, with litters containing 3-7 Xenosaurus grandis
~~~~~~-"-=-~"'--~~-
young (FRITTS 1966). ALVAREZ DEL TORO
(1960) reported that births among X. gran-
dis occur in Chiapas, Mexico, from March Further Reading
to July, and that newborns have a total FRITTS 1966, PRESCH 1981, BALLINGER et al.
length of 45 mm. 1995

177
 Amphisbaenia
Worm Lizards (Amphisbaenia)
Although the taxonomic position of the
worm lizards has yet to be satisfactorily
resolved, this group is generally regarded
as an independent suborder related to the
lizards (Sauria) and snakes (Serpentes).
Evidence exists that the worm lizards had
already separated from the snakes and
lizards (order Squamata) during the
Cretaceous period, even prior to the rise of
modem lizard families (GANS 1969, 1978).
Earlier authors had considered the worm
lizards to be a lizard family (VANZOLINI
1951). A number of anatomical characters
distinguish the worm lizards from both
the snakes and the lizards. These include
the reduction of the right lobe of the lung
and the existence of a middle pre-maxil-
lary tooth, as well as a characteristic skull
and middle ear morphology (GANS 1969).
Worm lizards are the true burrowers
among the reptiles. Living in self-con-
structed, permanent tunnel systems,
amphisbaenians have an elongated body
and, with the exception of one genus
(Bipes), no external limbs. Rather than use
already existing tunnels as worm snakes
will, or staying in loose soil or sand, they
are in the habit of tunneling through com-
pact earth. They seldom come to the sur-
face and then only at night. Frequently,
Amphisbaena fuliginosa
"" Amphisbaena spurrelli
178
they can be found in the underground colo-
nies ofleaf-cutter ants (GANS 1969). Worm
lizards are equally able to crawl back-
wards and forwards in their tunnels.
Amphisbaenids are skilled hunters, prey-
ing upon insects, spiders, and rodents, as
well as worm snakes and other reptiles
(GANS 1969).
Amphisbaena alba LINNAEUS 1758, Systema
Naturae, ed. 10: 229; type locality: America.
SVL to 670 mm. Tropical South America east
of the Andes. Occurrence in Panama ques-
tionable.
Amphisbaena fuliginosa LINNAEUS 1758,
Systema Naturae, ed. 10: 229; type locality:
America. SVL to 500 mm. Panama, tropical
South America east of the Andes, as well as
western Colombia and Ecuador.
Amphisbaena spurrelli (BOULENGER 1915),
Proc. Zool. Soc. London 1915: 659; type locali-
ty: Andagoya, at junction of Rios Condoto and
San Juan, Colombia. SVL to 300 mm.
Panama, northern Colombia, and Venezuela.
Fig. 483. Amphisbaena fuliginosa (Peru).
Photo: E . Lehr
 Amphisbaenia

Key to Amphisbaenia
1 a 65 or more scales at mid body .................. .
. . . . . . . . . .. .................. .. .. Amphisbaena alba
b 55 or fewer scales at midbody .. .... .... ..... . 2
2 a Irregular pale-dark spotting; more than
four precloacal pores .. .... ...... ........ ........ . .
.. . . . . . . . . . . . . . . . . . . Amphisbaena fuliginosa
b Without spotting; four precloacal pores
......... ... ............. Amphisbaena spurrelli
Further Reading
VANZOLINI 1951, GANS 1962a, b, 1967, HooG-
MOED 1973, GANS & MATHERS 1977, GANS 1978

Fig. 485. Amphisbaena alba.

Photo: R. W. Van Devender

nas;r~efrontal
ocular temporal

rostral · •
supralabials . .
nasal infralabials
prefrontal

ocular
I

- Fig. 486. Amphisbaena alba.

ll"FF?~~~ precloacal
pores
nN=Fr=?=l=l-+-t11 precloa cals

......,_~__,_......,__,_,_~
--
cloaca
postcloaca ls
:i:;
~~ l
>la oJ~

I
)
}'
'/
J
Photo: R. W. Van Devender

I
Is

-
>->--
-
--
"t-rl-~
~1 1 -r. J
m1 I I 1/ IJI
,

Fig. 487. Amphisbaena fuliginosa (Nusagandi,

Fig. 484. Amphisbaena spurrelli. Modified Comarca de San Blas, Panama).
from BOULENGER (1915). Photo: G. Kohler
179
 Serpentes
Snakes (Serpentes)
anterior temporal postoculars
parietal
Fig. 488. Scalation
characters on the
head of a colubrid
snake.
infralabials
internasal
In Central America, the snakes are well
represented by 290 different species.
Key to Snake Families
1 a Tail conspicuously flattened laterally (Fig.
843, p. 316) .............................................. .
............... Elapidae (in part: sea snakes)
b Tail more or less round in cross-section 2
2 a Ventral scales not distinctly enlarged,
scales around body all approximately of
equal size ........ .............................. .. ......... 3
b Ventral scales distinctly larger than dorsal
scales .......... .... .... ................................... 5
3 a 14 scale rows at midbody .............. ...... ..
.......... .... ... .. ............ .. Leptotyphlopidae
b 18 or more scale rows at mid body ........... 4
4 a 18-20 scale rows at midbody; one preocu-
lar; no subocular ..... ........ .. Typhlopidae
b 21-22 scale rows at midbody; preocular
present or not; subocular present or not
.. . . . ..... .... .. ... .... ..... . ..... Anomalepididae
5 a One tube-like loreal pit between nostril
and eye .............................. ........ Viperidae
b Without tube-like loreal pit between
nostril and eye ........ ........ .... .. .. ................ 6
180
supraocular
rostral
prenasal
u.-~-n•-- mental
supra labials
1st dorsal row
ventrals
chin shields
,r---r-.,-.4-...+--1
mental
mental groove
6 a Ventrals do not extend across the entire
body width (one or more dorsal rows on
the ventral side); usually all or most sub-
caudals undivided (exception: Loxocemus);
often with cloacal spurs on either side of
the cloaca ................. ... ... ....... ................ 7
b Ventrals extend across the entire body
width (no dorsals on the ventral side);
usually all or most subcaudals paired
(exception: Pseudoboa); no cloaca! spurs
present .. ................................................... 9
7 a All or most subcaudals paired .... .... ............
............ ... ..... .... ... ............... Loxocemidae
b All or most subcaudals undivided ........... 8
8 a Fewer than ~ dorsal rows at midbody
.................... .... ............... Tropidophiidae
b More than 35 dorsal rows at midbody ..... .
..................... ......... ........ ............... Boidae
9 a Upper jaw with 2 immobile, enlarged
teeth, otherwise, upper jaw without teeth;
no loreal; body pattern with complete
black rings ...................................................
........... Elapidae (in part: coral snakes)
b Upper jaw with many teeth, none conspic-
uously enlarged in front of upper jaw; lore-
al present or not; coloration and markings
variable ... ..... ............ .... ....... Colubridae

Blind Snakes (Scolecophidia)
eye
rostral
ocular
mental ----1•~~~':::CS.D.L
Fig. 489. Scalation characters on the head of a
blind snake.
Three families of comparatively primitive
snakes are grouped under the term blind
snakes (Scolecophidia). They are charac-
terized by shiny, smooth scales of fairly
uniform size surrounding the entire body.
Fig. 490. Anomalepis mexicanus (Saslaya
National Park, Nicaragua). Photo: A. Quintana
Anomalepididae
The family Anomalepididae is made up of
four genera (Anomalepis, Helminthophis,
Liotyphlops, and Typhlophis; the last not
in Central America), with a total of 15 spe-
cies. Little is known about the habits of
these small underground-dwelling snakes.
In Central America, the genera
Ano malepis, Helminthophis, and
Liotyphlops are each represented by a
single species.
Scolecophidia
Key to ~alepididae
1 a Dorsal surface of head with two large pre-
frontals behind the rostral, broadly in
contact with each other and followed
posteriorly by a large frontal (Fig. 491a) ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... Anomalepis
b Scalation on dorsal surface of head diffe-
rent than that above ................................ 2
2 a Prefrontals behind the rostral in contact
with each other (Fig. 491b) Helminthophis
b Prefrontals not in contact with each other,
separated by conspicuously large rostral
(Fig. 491c) .... ...... .... ............ Liotyphlops
a. Anomalepis b. Helminthophis
Fig. 491. Dorsal head scales
(prefrontals yellow;
parietal brown).
c. Liotyphlops
Anomalepis
The genus Anomalepis includes four spe-
cies (KOFRON 1988a), one of which occurs
in southern Central America. One indivi-
dual of A. mexicanus was collected under a
log on a bank above a stream and another
specimen was inside a rotten log on a hill-
side (MCCRANIE 2004a). This snake is ovi-
parous, as verified by the dissection of a
female, which contained two eggs (KOFRON
1988a).
181
 Anomalepididae

Anomalepis mexicanus JAN 1861, Icon. Gen. Liotyphlops

Ophid., Livr. 1: pl. 5, fig. 1, pl. 6, fig. 1; type
locality: "Mexico" [in error). TL to 153 mm.
Honduras to Panama; also in Peru, 300-500 m
elevation; distribution in Colombia and
Ecuador not yet demonstrated (KOFRON
1988a).
Further Reading
TAYLOR 1939, DUNN 1941, ROBB & SMITH 1966,
KOFRON 1988a, KOHLER et al. 2004, MCCRANIE
2004a.
According to the most recent revision of
Liotyphlops by DIXON & KOFRON (1984),
this genus is represented in Central
America solely by the species L. albiros-
tris. The other seven species of this genus
are distributed in South America.
Liotyphlops albirostris (PETERS 1857),
Monats. Akad. Wiss. Berlin 1857: 402; type
locality: Panama. TL to 223 mm. Southern
Central America (southeastern Costa Rica,
Panama) and northwestern South America
(Ecuador, Colombia, Venezuela), as well as on
Cura~ao (Netherlands Antilles), near sea
level to 1640 m elevation.

Further Reading
DIXON & KOFRON 1984

Fig. 492. Helminthophis frontalis (San Jose,

Costa Rica). Photo: R. D. Bartlett

Helminthophis
Fig. 493. Liotyphlops albirostris (Isla Pachaca,
Little is known about Helminthophis fron- Pearl Islands, Panama). Photo: C. W. Myers
talis , other than that it lives as a secretive
ground dweller. It is distributed in south-
ern Central America.

Helminthophis frontalis (PETERS 1860),

---· "' Anomalepis mexicanus
..,. Helminthophis frontalis
Monats. Akad. Wiss. Berlin 1860: 517; type Liotyph/ops albirostris
locality: Costa Rica. TL to 160 mm. Central
Costa Rica and sout~estern Panama, 90-
1440 m elevation.

Further Reading
AMARAL 1924a, WALLACH & GONTHER 1997

182
 Leptotyphlopidae

Leptotyphlopidae
The family Leptotyphlopidae contains two
genera (Leptotyphlops and Rhinoleptus)
with a total of 80 species worldwide. Two
species of Leptotyphlops occur in Central
America.

Leptotyphlops

The taxonomy of the Central American

Leptotyphlops is poorly understood.
KOHLER (2001a,b) and SAVAGE (2002) treat- Fig. 494. Leptotyphlops goudotii (Isla de
ed Leptotyphlops nasalis and L. ater, re- Guanaja, Honduras). Photo: G. Kohler
spectively, as valid species. However, these
decisions appear to be only weakly support-
ed and I prefer to not recognize these taxa
until a comprehensive review of the
Central American Leptotyphlops has been
published. Leptotyphlops nasalis has been
described based on a single specimen from
Managua, Nicaragua, which differs from
the widely distributed species L. goudotii
by a single distinguishing scale character,
which might also be due to individual
anomaly. Since L. nasalis was first describ-
ed over 60 years ago, no further specimens
that could be assigned to this taxon have
been found . Leptotyphlops goudotii is the
most common Leptotyphlops found in
Central America. These not even pencil- Fig. 495. Leptotyphlops goudotii can be found
thick snakes have a bright yellow spot on in termite nests. Photo: G. Kohler
both the anterior and the posterior end. As
the very small head is not well differentia-
ted from the body, it is difficult at first
glance to distinguish the head from the tail
tip. These blind snakes live secretively in
rotting logs and in the mounds of termites
or ants, which make up the bulk of their
food intake (KOHLER 1998£). These little
snakes most often eat their prey whole.
However, they also have been observed
sucking out the soft body contents of insects
and leaving behind empty exoskeletons
(Sco'IT 1983c). Leptotyphlops goudotii are
oviparous and produce clutches of 8-12
eggs (ALVAREZ DEL TORO 1983).

Fig 496. Leptotyphlops macrolepis (Cordoba,

Colombia). Photo: M. Lundberg
183
 Leptotyphlopidae
Leptotyphlops goudotii (DUMERIL & BIBRON
1844), Erp. Gen. 6: 330; type locality: Valley of
Rio Magdalena, Colombia. TL to 185 mm.
Colima and Veracruz, Mexico, through
Central America to Colombia and Venezuela,
sea level to 1600 m elevation in dry forest and
wet forest.
Leptotyphlops macrolepis (PETERS 1857),
Monats. Akad. Wiss. Berlin 1857: 402; type
locality: Caracas and Puerto Cabello,
Venezuela. TL to 400 mm. Panama and north-
ern South America.
Key to Leptotyphlops
1 a Without pale longitudinal lines (Fig. 496);
supraoculars about as large as prefrontal
and frontal ... Leptotyphlops macrolepis
b Pale longitudinal lines present (Fig. 495);
supraoculars somewhat larger than pre-
frontal and frontal ............................... .
.......................... Leptotyphlops goudotii
Further Reading
HAHN 1979, SCOTT 1983c, KOHLER 2000b
a. R. braminus b. T. costaricensis c. T. tenuis
Fig. 497. Lateral head scalation in Typhlops
and Ramphotyphlops (rostral brown, prefron-
tals yellow).
• Leptotyphlops goudotii
• Leptotyphlops macro/epis
184
Typhlopidae
The family Typhlopidae is represented by
two genera in Central America, one of
which (Ramphotyphlops) has been intro-
duced in several countries. The species in
this family only have teeth on the upper
jaw and not on the lower jaw, unlike the
Leptotyphlopidae and Anomalepididae.
Similar to these species, however, they are
also small, worm-like snakes that live very
secretively underground. Their dorsal and
ventral scales are similarly smooth and of
equal size.
Key to Typhlopidae
1 a More than 345 dorsal scales between
rostral and tail spike; scale suture between
nostril and rostral on a diagonal (Fig.
497b,c) ............... .... .... ...... .... .. Typhlops
b Fewer than 345 dorsal scales between
rostral and tail spike; scale suture between
nostril and rostral almost vertical (Fig.
497 a) ............ .. .............. Ramphotyphlops
Fig. 498. Dorsal head scalation in Typhlops
costaricensis (rostral brown, prefrontals
yellow).
• Typhlops tenuis
~ . Typh/ops stade/mani
• Typhlops microstomus
""" Typhlops costaricensis
,.. Ramphotyphlops braminus
 Typhlopidae

Typhlops

The genus Typhlops is found throughout

the tropics and subtropics worldwide. In
Central America, there are four species of
this genus, one of which (T. stadelmani)
has only recently been resurrected
(MCCRANIE & WILSON 2001b). These blind
snakes live very secretively buried under-
ground, where they feed on termites, ants,
and other small insects. The Central
American species of this genus are ovi-
parous.

Fig. 499. Ramphotyphlops braminus.

Photo: R. D. Bartlett

Ramphotyphlops

Ramphotyphlops braminus is a species

native to Southeast Asia, but has been
introduced throughout the tropics world-
wide. In Central America, this species has
been reported in Mexico, El Salvador and
Guatemala, where it has recently become
indigenous (DIXON & HENDRICKS 1979, LEE
1996, DUENAS et al. 2001). It is assumed
that this small snake is transported into Fig. 500. Typhlops stadelmani (near Quebrada
foreign countries along with the soil on Grande, Copan, Honduras). Photo: G. Kohler
plants. As it is parthenogenetic, only a
single female is required to establish a new
population (NUSSBAUM 1980). Rampho-
typhlops braminus produces clutches of 2-
3, possibly up to 6 eggs (NUSSBAUM 1980).

Ramphotyphlops braminus (DAUDIN 1803),

Hist. Nat. Rept. VII: 279; type locality:
Vizagapatam, India. TL to 173 mm.
Southeast Asia, Africa, Madagascar, Indian
Ocean, Hawaii, South Pacific and Australia;
introduced in several locations in Central
America.

Further Reading
DIXON & HENDRICKS 1979, NUSSBAUM 1980, OTA Fig. 501. Typhlops microstomus (Quintana Roo,
et al. 1991 Mexico). Photo: H. Bahena B.
185
 Typhlops

Typhlops costaricensis JIMENEZ & SAVAGE Key to Typhlops

1962, Rev. Biol. Trop. 10: 199; type locality:
Monteverde, Sierra de Tilaran, Puntarenas,
Costa Rica, 1500 m. TL to 360 mm. Central
Honduras to Costa Rica, .
Typhlops microstomus COPE 1866, Proc. Acad.
Nat. Sci. Philadelphia 18: 125; type locality:
Yucatan, Mexico. TL to 366 mm. Central
Veracruz, Mexico, as well as the Yucatan
Peninsula.
Typhlops stadelmani SCHMIDT 1936a, Proc.
Biol. Soc. Washington 49: 48; type locality:
Subirana Valley, Yoro, Honduras. TL to 310
mm. Northwestern Copan and southwestern
Yoro, Honduras, 850 to 1370 m elevation in
wet forest.
Typhlops tenuis SALVIN 1861a, Proc. Zool. Soc.
London 1860: 545; type locality: Cohan, Alta
Verapaz, Guatemala. TL to 326 mm.
Veracruz, Mexico, to Guatemala, sea level to
1500 m elevation in dry forest and wet forest.
1 a 20 scales at midbody ...................... ......... . .
............................ Typhlops costaricensis
b 18 scales at midbody ..... ...... .. ....... ......... 2
2 a More than 4 70 dorsal scales between
rostral and tail spike; subocular present
..... .............. .. .... Typhlops microstomus
b Fewer than 450 dorsal scales between
rostral and tail spike; subocular absent ... 3
3 a 361-441 dorsal scales between rostral and
tail spike; body with brown spots ............. .
... ............... ... ...... ... .... ... Typhlops tenuis
b 341-369 dorsal scales between rostral and
tail spike; body solid color pink (in life) and
pale brown (in preservative) ................. .... .
. ..... ...................... . Typhlops stadelmani
Further Reading
VILLA 1978, DIXON & HENDRICKS 1979,
MCCRANIE & WILSON 200lb

Macrostomata

According to MCDIARMID et al. (1999), six

families of early snakes (i.e., Boidae,
Bolyeriidae, Loxocemidae, Pythonidae,
Tropidophiidae, and Xenopeltidae) are
included in the Macrostomata.

Boidae
The species of the family Boidae are wide-
ly distributed in the tropics and subtropics
of the Old and New World. Characteristic
of the boids are the head, which is clearly
differentiated from the neck, vertically slit
pupils, remnants of the pelvic girdle
(which in some species is recognizable by Fig. 502. Boa constrictor sabogae (Isla Saboga,
externally visible cloaca} spurs) as well as Panama). Photo: G. Kohler
the presence of functional lungs on both
the left and right sides. All boids are non-
venomous and kill by constriction, using
the muscular coils of their body to suffo-
cate prey. The jaws are equipped with Further Reading
strong teeth. All boids are live bearers. KLUGE 1991, ZAHER 1994
186

Fig. 503. Boa constrictor imperator (Reserva Foresta} Fortuna, Panama).
Key to Boidae
1 a All scales on dorsal surface of head small
and irregular, no enlarged scales on the
snout; no labial pits ....... .. ... .. .......... .. Boa
b Snout with several enlarged shields; labi-
al pits present .. ..... .. ...... .... ........ .. ... .... 2
2 a More than 245 ventrals; more than 65 sub-
caudals; usually 3, rarely 2 loreals ......... .. .
...................... ............ ................. . Corallus
b Fewer than 240 ventrals; fewer than 65
subcaudals; usually one large loreal, sel-
dom 2 loreals ............... ...... ...... Epicrates
Boa
Boa constrictor is potentially the largest
as well as the best-known snake in
Central America. It is a large, heavy-
bodied snake that can be found both on the
ground as well as in the branches of large
trees. Although it is predominantly cre-
puscular and nocturnal, it is occasionally
encountered outside of its hiding places
during the day. Typical daytime hiding
places include hollow trees, as well as piles
Boidae
Photo: G. Kohler
of leaf litter among the buttress roots of
large trees. Boas are not too particular
about their vertebrate prey and will con-
sume any lizard, bird, or mammal they
can swallow. If they feel threatened, these
well-armed snakes will hiss with their
mouths wide open, and they are capable of
inflicting bloody and painful injuries with
their powerful jaws. Often pieces of the
long teeth will break off and remain
lodged in the wound. Boa constrictor is a
live bearer and produces litters of 10-64
young (GREENE 1983). On September 2,
1997, I discovered a female Boa constrictor
slightly more than 180 cm in total length
in the hollow stump of a dead coconut
palm on Utila. The snake was coiled up
and had obviously just given birth, as
there were 13 newborn snakes, with the
remains of umbilical cords, in the back of
the hole. The total length of the young
boas was between 49 and 52 cm, their
weight around 52 g.
187
 Boidae
Boa constrictor LINNAEUS 1758, Systema
Naturae, ed. 10: 215; type locality: "India" [in
error] . TL to 4450 mm (WATKINS-COLWELL &
LEENDERS 2003); however, most specimens
measure less than two meters). Tamaulipas
and Sonora, Mexico, to Peru, Bolivia, and
Argentina, as well as in the Lesser Antilles,
sea level to 1000 m elevation in dry forest,
wet forest, and rain forest as well as man-
groves. In Central America, the subspecies
Boa c. imperator DAUDIN 1803 (Mexico,
Central America excluding Toboga Island,
Panama, and northwestern South America)
and Boa c. sabogae (BARBOUR 1906) (Toboga
Island, Panama) occur.
Further Reading
STULL 1935, LAzELL 1964, GREENE 1983, WILSON
& MEYER 1985
• Boa constrictor
Corallus
There are seven different species in the
genus Corallus, all of which are crepuscu-
lar and nocturnal tree dwellers in rain
forest areas. The two Central American
species are extremely variable in terms of
coloration and pattern; this is particularly
the case with Corallus ruschenbergerii.
Corallus species will consume frogs,
lizards, small mammals, and birds, which
they kill by constriction. Corallus are
usually very aggressive, and a bite from
their particularly long front teeth can pro-
duce a deep wound. These snakes are live-
bearers, producing litters of 7 to 30 young
(Ross & MARZEC 1990).
188
Corallus annulatus (COPE 1876), J . Acad. Nat.
Sci. Philadelphia (2) 8: 129; type locality:
Costa Rica. TL to 1335 mm. Disjunct from
eastern Guatemala to southwestern Ecuador,
sea level to 400 m elevation in rain forest.
Corallus ruschenbergerii (COPE 1876), J.
Acad. Nat. Sci. Philadelphia (2) 8: 129; type
locality: Panama. TL to 2310 mm. South-
western Costa Rica to Panama, as well as
across northern South America east of the
Andes, sea level to 1000 m elevation in rain
forest.
Key to Corallus
1 a Nasal shields separated from one another
by rostral; scales posterior to nasal shields
not enlarged; fewer than 90 subcaudals
.......... .. .......... ......... Corallus annulatus
b Nasal shields in contact with one another;
scales posterior to nasal shields usually
distinctly enlarged; more than 90 sub-
caudals ....... ... Corallus ruschenbergerii
a. C. annulatus b. C. ruschenbergerii
Fig. 504. Snout scalation in Corallus (nasal
shields orange).
Further Reading
HENDERSON 1993a-c, STAFFORD & HENDERSON
1996, HENDERSON 1997, E. SMITH & ACEVEDO
1997, HENDERSON 2002.
""' '-/"'---~~
=
• Coral/us annu/atus
Coral/us rusch'enbergerii
 Boidae

Fig. 505. Juvenile Corallus annulatus (lzabal,

Guatemala). Photo: L. Melendez

Fig. 508. Corallus annulatus (near Tortuguero,

Limon, Costa Rica). Photo: M. Burger

Fig. 506. Corallus ruschenbergerii (Rio Sierpe,

Limon, Costa Rica). Photo: R. W. Van Devender

Fig. 507. Corallus annulatus (Limon Province, Fig. 509. Corallus ruschenbergerii (Costa Rica).
Costa Rica). Photo: R. W. Van Devender Photo: T. Leenders
189
 Boidae

Epic rates

Only one species of the genus Epicrates is

found in southern Central America, E. cen-
chria, whose common name, "Rainbow
Boa", comes from its iridescent coloration.
It is a slim, arboreal, crepuscular, and noc-
turnal species found in the rain forest. It is
a livebearer (15-28 per litter; Ross &
MARzEC 1990) that will eat frogs, lizards,
and small mammals.

Epicrates cenchria (LINNAEUS 1758), Systema

Naturae, ed. 10: 215; type locality: Surinam.
TL to 1500 mm. Costa Rica and Panama, as
well as widely distributed through tropical
South America; sea level to 500 m elevation
in rain forest. The subspecies Epicrates c.
maurus GRAY 1849a occurs in Central
America.
Fig. 511. Loxocemus bicolor from Costa Rica.
Insert: L. bicolor from Mexico (Playa Azul, Oaxaca).
Further Reading Photos: G. Kohler
KLUGE 1989

Fig. 510. Epicrates cenchria (Guanacaste, Costa Loxocemus bico/or

Rica). Photo: R. W. Van Devender
190
 Tropidophiidae

Loxocemidae Trachyboa

Loxocemus Little is known about the two species of

The Mexican burrowing python
(Loxocemus bicolor) is the only species in
the genus. It lives in dry forest and savan-
na habitat and can be seen both during the
day as well as at night outside of its hiding
place (personal observation). It digs
underground passages with its raised and
pointed snout and hunts whiptails
(Aspidoscelis) and small mammals. This
snake is also well known for successfully
tracking down and raiding iguana nests
(MORA 1987). Loxocemus bicolor reprodu-
ces oviparously, with the largest documen-
ted clutch containing four eggs (Ross &
MARzEC 1990).
the genus Trachyboa. In Ecuador, together
with R. SEIPP, I found a specimen of T.
boulengeri that was lying well camoufla-
ged and motionless on the ground during
the day. The animal behaved quite calmly
and made no attempt to flee. Rather, it
seemed to stiffen into whatever shape it
was manipulated. The species is live bear-
ing and produces 5-7 young per litter
(Ross & MARzEc 1990).
Trachyboa boulengeri PERACCA 1910, Ann.
Mus. Zool. Univ. Napoli 3 (12): 1; type locality:
unknown. TL to 430 mm. Choc6 region of
Panama through Pacfic Colombia to western
Ecuador, sea level to 750 m elevation in rain
forest.

Loxocemus bicolor COPE 1861c, Proc. Acad.

Nat. Sci. Philadelphia 13: 77; type locality:
unknown. TL to 1530 mm. Nayarit, Mexico,
along the Pacific coast to northwestern Costa
Rica; isolated occurrences also on the
Caribbean side, sea level to 600 m elevation
in dry forest and savanna.

Further Reading
NELSON & MEYER 1967

Tropidophiidae
The species of the family Tropidophiidae
are small snakes that are distributed in Fig. 512. Trachyboa boulengeri from Ecuador.
southern Mexico, Central America to Photos: G. Kohler
northwestern Pacific South America, and
on the West Indies. Contrary to an earlier
arrangement in which these snakes were ~ Ungaliophis continentalis
"-""-,..~---• Unga/iophis panamensis
included as a subfamily (Tropidophiinae)
in the Boidae, the tropidophiids are now
recognized as a distinct family (GREENE
1997, MCDIARMID et al 1999).

Key to Tropidophiidae
1 a Dorsals keeled; more than 27 dorsal rows
at midbody ....... .......... ......... . Trachyboa
b Dorsals smooth; fewer than 27 dorsal rows • Trachyboa
at midbody ... ... .... .. .. .. ....... Ungaliophis
191
 Tropidiphiidae

Ungaliophis

The dwarf boas of the genus Ungaliophis

are represented by two species, which are
distributed from Mexico across Central
America and into northeastern Colombia.
Little is known about these secretive
snakes. In Selva Negra, Nicaragua, about
two hours after dusk, I discovered an U.
continentalis on a tree overgrown with
tillandsias. The animal was flicking its
tongue as it hung about two meters above
the ground. Other observers have reported
Ungaliophis as a nocturnal tree-dweller
that prefers to live in dense epiphytic
growth (CORN 1974). These are attractive
snakes that are completely harmless and
make no attempt to bite when handled. A
specimen of U. continentalis that was kept
at Senckenberg Museum for several years Fig. 514. Ungaliophis continentalis (Selva
ate new-born mice. Ungaliophis continen- Negra, Matagalpa, Nicaragua).
talis bears 5-6 young per litter (Ross & Photo: G. Kohler
MARzEC 1990).
Ungaliophis continentalis MOLLER 1882,
Verh. Naturforsch. Ges. Basel 7: 142; type
locality: Retalhuleu, Guatemala. TL to 482
mm. Chiapas, Mexico, to northern Nicaragua,
1000-2130 m elevation in wet forest and
cloud forest.
Ungaliophis panamensis SCHMIDT 1933a,
Smithson. Misc. Coll. 89 (1): 12; type locality:
Cerro Brujo, Panama. TL to 4 70 mm.
Southern Nicaragua to western Colombia,
sea level to 2100 m elevation in rain forest
and cloud forest.

Key to Ungaliophis
1 a Rostral and prefrontal broadly in contact,
therefore, internasals separate from one
another; blotches on body sides oval; 25
dorsal rows at midbody, 15 dorsal rows one
headlength anterior of cloaca ....... .. ..........
...... ............... Ungaliophis continentalis
b Rostral and prefrontal not in contact,
internasals in contact with one another;
blotches on body sides triangular; 19-23
dorsal rows at midbody, 17 dorsal rows one
headlength anterior of cloaca .. .. .. ..
..... ... .. . . . . . . ..... Ungaliophis panamensis

Fig. 513. Ungaliophis panamensis (Guyacan de Further Reading

Siquirrez, Limon, Costa Rica). BOGERT 1968a, CORN 1974, MERAHTZAKIS 1988,
Photo: G. Kohler VILLA & WILSON 1990
192
 Colubridae

Colubridae
Approximately 80% of the known snake
species belong to the family Colubridae.
This family is distributed worldwide and,
on almost all continents, it is the dominant
family in terms of both the number of spe-
cies and their frequency. The one exception
is Australia, where the cobras, kraits and
elapids (family Elapidae) are most com-
monly represented. There is some contro-
versy regarding the intrafamilial structure
of Colubridae. In Central America, 64
genera from this family are known. The
many species (divided among 270 genera Fig. 515. Dipsas brevifacies (southeast of Coba,
Quintana Roo, Mexico). Photo: J.C. Lee
worldwide) of the family Colubridae dis-
play tremendous morphological and ecolog-
ical variety. Thus, in Central America,
there are slenderly built climbing species
(Oxybelis spp., Leptophis spp., Imantodes
spp.), ground-dwellers with stocky
(Xenodon rabdocephalus) or very slim body
shape (Leptodrymus pulcherrimus) and
species that live in streams or swamps
(Tretanorhinus nigroluteus, Hydromor-
phus concolor, Nerodia rhombifera). Some
species reach over two meters in total
length (Clelia clelia, Leptophis ahaetulla,
Masticophis mentovarius), while others are
fully grown at less than 30 cm (e.g.,
Geophis hoffmanni, Tantilla schistosa,
Tantillita lintoni).

Some Colubrids have enlarged grooved Fig. 516. Oxybelis aeneus (Isla de Roatan,
fangs in the back of the upper jaw and pro- Honduras). Photo: G. Kohler
duce a secretion that is moderately venom-
ous (Conophis lineatus, Oxybelis spp.).
Although the colubrid snakes of Central
America do not pose any mortal danger to
humans, their bite can lead to serious
localized swelling and considerable pain.

Fig. 517. Ninia sebae (near Santiago, Costa

Rica). Photo: G. Kohler
193
 Colubridae

Key to Colubridae
1 a Dorsal surface of head covered with small,
irregular scales (Fig. 518) ..... .. Nothopsis
b Dorsal surface of head covered with large
symmetrical shields (Fig. 519) ................. 2
2 a Rostral shield protruding and pointed (Fig.
521) ........ .......... .......... .. ...... ..... ............ ... 3
b Rostral shield normal (Fig. 519) ............... 4
3 a Fewer than 160 ventrals; cloacal scute
divided (Fig. 523b); no loreal .. ...... Ficimia
b More than 165 ventrals; cloaca} scute Fig. 518. Nothopsis rugosus; note the small,
undivided; loreal present ........ Phimophis irregular scales on top of head.
4 a Dorsal scales in even number of rows at Photo: R.W. Van Devender
midbody (compare Fig. 522) ....... ......... .. 5
b Dorsal scales in uneven number of rows at
midbody ..... ...... ............ .. ....... ..... ......... 6
5 a 10-12 dorsal rows at midbody; fewer than
170 ventrals; cloaca} scute usually divided
(Fig. 523b) ................ ......... .... Chironius
b 14-18 dorsal rows at midbody; more than
190 ventrals; cloaca} scute undivided (Fig.
523a) ... .......................... ................ Spilotes
6 a Without longitudinal mental groove (Fig.
590) ........... ..... ................................. Dipsas
b Longitudinal mental groove present ....... . 7 Fig. 519. Mastigodryas melanolomus; note the
7 a Body extremely slender; head large and large, symmetrical shields on top of head.
distinct from thin neck (Fig. 520a); with Photo: G. Kohler
large protruding eyes; 195-262 ventrals;
106-178 subcaudals ................. lmantodes
b Combination of characters different from
that above ............ .. .... ...... .... .. .......... ...... 8
a. lmantodes
8 a Number of dorsal rows constant from mid-
body to cloaca, no reduction ... ..... ..... .. .... 9
b Number of dorsal rows one headlength
anterior to cloaca at least two rows fewer
than at midbody ....... ... ..... .. .. ...... .. ... ..... 36 b. Oxyrhopus
Fig. 521. Ficimia
9 a Cloacal scute undivided (Fig. 523a) ....... 10 Fig. 520. Head shape publia (from DUMERIL
b Cloaca} scute divided (Fig. 523b) .. .......... 16 in two colubrids. et al. 1870-1909).
lOa No anterior temporal .......... ....... ............. 11
b One or two anterior temporals ............... 12
l la 13 dorsal rows at midbody Sibon (in part)
b 15-17 dorsal rows at midbody ...... Geophis
12a Dorsal scales strongly keeled ........... Ninia
b Dorsal scales smooth or only slightly
keeled ............ ... .... .............................. ... 13
13a Unicolor black (adults) or red with pale
neck band (juveniles) ...... Clelia (in part)
b Coloration different from above ............. 14
Fig. 522. How to count dorsal scale rows; the
14a 13-15 dorsal rows at midbody ................ . example shows 21 dorsal scale rows in a
............... ..... ... . . . ... . . . . . . . . .... Sibon (in part) Rhadinaea .
194 b 17-19 dorsal rows at midbody ................ 15
 Colubridae

15a Sixth supralabial not conspicuously en- Fig. 523.

larged, not in contact with postocular, an- Subcaudals
terior and posterior temporal; fewer than can be paired
40 subcaudals ............................. Atractus undivided
(as shown) or
divided unpaired.
b Sixth supralabial enlarged and in contact
with postocular, anterior and posterior
temporal; more than 40 subcaudals ..........
.. ............. ......... .... ............ Tropidodipsas subcaudals

16a Dorsal scales keeled (Figs. 527c,d) Storeria b.

b Dorsal scales smooth (Figs. 527a,b) ....... 17
17a Prenasal fused with bordering internasal Fig. 524. Stenorrhina
(Fig. 524) .. .... ...... ... ...... ... ..... .... ........... 18 degenhardtii; note that
the prenasal is fused to
b Prenasal not fused with bordering interna- the adjacent internasal
sal .................. .... ... .. ..... ...... .. ...... .. ..... 19 (orange).
18a 17 dorsal rows at midbody; fewer than 55
subcaudals ......... ...... .... ...... Stenorrhina
b 15 dorsal rows at midbody; more than 90
subcaudals .... ..... .. Symphimus (in part) Fig. 525. N inia sebae;
the loreal (orange)
19a Body with complete black rings ....... ...... 20 reaches to the eye;
b Body without complete black rings, at most there is no preocular.
dark bands that are not closed on the ven-
tral side ...................................... ... .... .. .. 22
20a 17 dorsal rows at midbody; fewer than 150
ventrals ......... ... .... .................. Pliocercus
b 15 dorsal rows at midbody; more than 160
ventrals ... ................... ..... .... .... ....... ....... 21
21a One posterior temporal; black rings at
regular intervals; in life, a bright orange-
red dorsal blotch between each of the black
rings (Fig. 526); in preservative, without
these blotches, only with black and white
rings ........... ... ... .. .. .. ... .. ... .... Scolecophis
b Two posterior temporals; black rings not at
regular intervals, either a pair of black
rings or rings set somewhat irregularly or Fig. 526. Scolecophis atrocinctus (Nicaragua).
partially along the vertebral line; without Photo: G. Kohler
orange-red blotch dorsally between the
black rings in life ........... Erythrolamprus
22a 15 dorsal rows at midbody .. ... ....... ......... 23
b 17-21 dorsal rows at midbody ................ 28
23a Preocular and loreal present, therefore, at
least three scales between nostril and eye a. Stenorrhina b. Mastigodryas
.......... ..... ....... .. ... .. ... ... .. ...... .............. 24
b Either preocular or loreal missing, there-
fore, only two scales between nostril and
eye .. ............... ....... ......... .. .... ... .... ...... 25
24a Fewer than 160 ventrals; fewer than 80
subcaudals ...... ...... Trimetopon (in part) c. Senticolis d. Thamnophis
b More than 160 ventrals; more than 90 sub-
caudals ...... ..... ...... Symphimus (in part) Fig 527. Dorsal scales
25a No preocular; loreal reaches to eye ........ 26 a. smooth, without apical scale pits
b. smooth, with two apical scale pits
b Preocular present; no loreal ................... 27 c. keeled, with two apical scale pits
d. keeled, without apical scale pits
195
 Colubridae

26a Fewer than 160 ventrals; fewer than 60

subcaudals ........... ................. Adelphicos
b More than 170 ventrals; more than 80 sub-
caudals ............ ...... .... ............ Enuliophis
27a Fewer than 21 maxillary teeth, the poster-
iormost pair grooved; more than 126 ven-
trals or, if fewer than 126 ventrals, then
without pale spots on supraoculars and
prefrontals ................. .... ......... Tantilla Fig. 528. Hydromorphus concolor; internasals
(brown) and prefrontals (yellow) are fused,
b More than 21 maxillary teeth, the poster- respectively.
iormost pair not grooved; fewer than 116
ventrals or, if 116-126 ventrals (T. brevissi-
ma), then with small pale spots on supra-
oculars and prefrontals ............ Tantillita
28a 19-21 dorsal rows at midbody ........ ........ . .
. . . . . . . . . . . . . . ................ Rhadinaea (in part)
b 17 dorsal rows at midbody .......... ..... .... 29
29a Anterior portion of body dorsally patterned
similar to coral snakes (paired black bands
with pale spaces in between on red back- Fig. 529. Oxybelis fulgidus; the loreal is absent;
ground); posterior portion of body either the prefrontals (orange) contact the supralabials.
colored like anterior or predominantly uni-
color brown) ............... Scaphiodontophis
b Coloration different than above ............. 31
30a Number of dorsal rows one headlength
posterior of head two rows fewer than at
midbody ............................... Chapinophis
b Number of dorsal rows constant from head Fig. 530. Dorsal
to cloaca ........... .. .. .. ... ......... .... ................ 31 scales in
Diaphorolepis
31a No preocular; loreal reaches to eye (Fig. (from BOGERT 1964).
525) ...................................... .. ........ ...... 32
b Preocular present; loreal present or not 33
32a Anterior temporal present; ratio of tail
length I TL > 20% .. .. .. ... ... ..... ...... Enulius
b No anterior temporal; ratio of tail length I
TL/< 10% ....... .. ... .. ...... ... . Omoadiphas
33a Dorsal scales in neck area with apical pits
(Fig. 527b); dorsum of head noticeably
paler in color than dorsum of body; with Fig. 531. Enulius bifoveatus.
distinct canthus rostralis ... Amastridium
b Without apical pits (Fig. 527a); head colo-
ration variable; canthus rostralis not
distinct .... ..... ... ..................................... 34
36a Body with complete black rings .... ......... 37
34a Usually obvious white line along the first b Body without complete black rings, at
dorsal rows ... ... ..... ... ... .. ..... .... Urotheca most with dark bands that are not closed
b No obvious white line along first dorsal on the ventral side ............. .. .... ........... 38
rows .... ...... ... .... .. ...... .. .......................... 35
37a Each shield on the dorsal surface of head
35a Prefrontals not fused into a single shield; black with sharply contrasting pale edging
usually more than 50 subcaudals .............. . (Fig. 532); more than 100 subcaudals; cloa-
......... ..... ... .. ......... Rhadinaea (in part) cal scute divided ............ Rhinobothryum
b Prefrontals fused into a single shield or b Head coloration different than above;
not; if not, then usually fewer than 50 sub- fewer than 80 subcaudals; cloacal scute
caudals .... ........ .. .. Trimetopon (in part) undivided ......... ...... ..... .... Lampropeltis

196
 Colubridae

38a Scales of the vertebral row with two keels

(Fig. 530) ............... .... ....... Diaphorolepis
b Scales of the vertebral row smooth or with
only one keel ..... .. ............ .. ... ................... 39
39a Prefrontals fused into a single shield (Fig.
528), or three prefrontal shields present
...................... ................... . Hydromorphus
b Two prefrontals shields present .............. 40
40a No loreals, prefrontals in contact with
supralabials (Fig. 529) ......... ...... ........... 41 Fig. 532. Rhinobothryum bovallii.
b One or more loreals, prefrontals not in Drawing: M. Vesely
contact with supralabials .. ....... ... ......... 44
41a Fewer than 85 subcaudals; 19 (rarely 17)
dorsal rows at midbody ....... .. .. Manolepis
b More than 85 subcaudals; 15-17 dorsal
rows at midbody ....... ... ......................... 42
42a 17 dorsal rows at midbody ............. .. ... .. ... .
. . . ... . . . . . . . . . . . . . . . . . . .......... Oxybelis (in part)
b 15 dorsal rows at midbody ........ ............. 43
43a Cloaca} scute undivided; 13 dorsal rows
one headlength anterior of cloaca .. ...... .... .
. . . ... . . . . . . . . . . . . . . . . . . . . . . . . ... Oxybelis (in part)
b Cloaca} scute divided; 11 dorsal rows one
headlength anterior of cloaca .............. ..... .. Fig. 533. Dorsal scales in Xenodon .
.......... .... ... .. ... .. ........... Leptophis (in part)
44a Three loreals; an obvious lyre-shaped
marking dorsally behind the head; 25-27
dorsal rows at midbody .... Trimorphodon 51a Subcaudals not paired ........... Pseudoboa
b One or two loreals; no obvious lyre-shaped b Subcaudals paired .... .. ....... .................. 52
marking dorsally behind the head; number
of dorsal rows at midbody variable ........ 45 52a Dorsum with coral snake-like pattern of
black and red or reddish orange cross
45a Cloaca} scute undivided ............... ..... ..... 46 bands ...................................... Oxyrhopus
b Cloaca} scute divided ....... ....... ......... .... 56 b Dorsum without coral snake-like pattern
of black and red or reddish orange cross
46a Dorsal scales keeled ................. .. ........... 47 bands ................................ ...................... 53
b Dorsal scales smooth .............................. 50
53a 56-93 subcaudals ................... ..... ..... ..... 54
47a More than 25 dorsal rows at midbody ........ . b 105-128 subcaudals ............................... 55
......... ..... .................................... Pituophis
b 25 or fewer dorsal rows at mid body ....... 48 54a Dorsal surface of head reddish to dark
brown, followed by a white or yellow occi-
48a 19-25 dorsal rows at midbody; more than pital collar and a long postnuchal black
180 ventrals ........................ ........ Pseustes band; body red to reddish brown with
b 17-19 dorsal rows at midbody; if 19 dorsal small black blotches or narrow bands ......
rows, then fewer than 180 ventrals ........ 49 ..... ......................................... Tripanurgos
b Coloration not as above ......... Siphlophis
49a 17 dorsal rows at midbody ......... .... ..... .
. . . . . . . . . . . . . . . ... .... Dendrophidion (in part) 55a Unicolor black (adults) or red with pale
b 19 dorsal rows at midbody .. Thamnophis neck band (juveniles); 17 or 19 dorsal scale
rows at midbody ............. Clelia (in part)
50a Fewer than 55 subcaudals; dorsal scale b Coloration different than above; 17 dorsal
rows run obliquely towards the middorsum scale rows at midbody .... .... Drymarchon
(Fig. 533) ........................ ... ..... .... Xenodon
b More than 55 subcaudals; dorsal scale 56a 25 or more dorsal rows at midbody ....... 57
rows run longitudinally .............. .... ...... 51 b 23 or fewer dorsal rows at midbody ....... 61

197
 Colubridae

57a More than 230 ventrals ............... .... ...... 58

b Fewer than 220 ventrals ........................ 59
58a Usually 8 supralabials, two bordering the
eye; dorsal blotches, if present, brown or
gray .......... ..................... .. .......... Senticolis
b Usually 9 supralabials, three bordering the
eye; dorsal blotches reddish .......... Elaphe Fig. 534. Pattern oflateral head in Conophis
59a Dorsal scales keeled .................... Nerodia lineatus (left) and Crisantophis nevermanni
(right). Drawings: M. Vesely
b Dorsal scales smooth ............................. 60
60a Body with dark spots ........... ................. .. ..
.. .. .. . . . . . . . . . .. .. . ......... Leptodeira (in part)
b Body with dark longitudinal stripes ........ .
.. .. .. .. .. . . .. . .. .. ...... Coniophanes (in part) f. ~ l ~~
~~
61a Dorsal scales keeled ........... ...... ..... ...... 62 ~
~ , l14~' ~~
~ ~
b Dorsal scales smooth ............................ 65 l ~~
~t
, I!l i~
~
62a Nostrils directed dorsolaterally ...... ....... .. .
. . . . . . . . . . . . . . . . . .... . . . . . . .. .. ...... Tretanorhinus I~ ~
't
~
ft
,
1'
.•l
ft
,,~
~
~
b Nostrils directed laterally ....... .. ... .. ...... 63 ~ ~ ! ~
63a 15 dorsal rows at midbody ......... ....... ... ..
............................... Leptophis (in part)
"'
Fig. 535. Pattern of dorsal body in Conophis
lineatus (left) and Crisantophis nevermanni
b 17 dorsal rows at midbody .................. 64 (right). Drawings: M. Vesely
64a Body with narrow diagonal stripes, poster-
ior area longitudinally striped or complete-
ly unicolor (Figs. 577-582); rear maxillary
teeth obviously enlarged .......................... .. 70a More than 194 ventrals; more than 140
............. ............ Dendrophidion (in part) subcaudals; three postoculars ...................
.. .. .... ......... ... ...................... Leptodrymus
b Body unicolor green in life (blue in preser-
vative) or with large dark spots or a pale b Fewer than 197 ventrals; fewer than 140
spot on each dorsal scale so that body sur- subcaudals; two postoculars .................. 71
face appears entirely speckled (Figs. 598- 71a One small presubocular below the preocu-
604); all maxillary teeth approximately the lar ......................... .. ...... .............. Coluber
same size ............ ....... .... ........ Drymobius
b No small presubocular below the preocu-
65a 13 dorsal rows one headlength anterior of lar ..................................... Mastigodryas
the cloaca ................................................. 66
72a Pupils vertically slit; dorsum with large
b 15 or more dorsal rows one headlength dark blotches or broad dark diagonal
anterior of the cloaca ............................ 67 bands ................... Leptodeira (in part)
66a Body without dark longitudinal stripes b Pupils round; dorsum longitudinally
......... ...... ..... ..... ....... ........ .. Masticophis striped, unicolor or with dark diagonal
b Body with dark longitudinal stripes bands ..................... ...... ........................... 73
........ .......................... .......... Salvadora 73a Dorsal scales with apical pits, at least at
67a Two anterior temporals ................ ........ 68 anterior portion of the body; body with
dark diagonal bands or, if longitudinally
b 0-1 anterior temporals ............................ 72 striped, then with two pairs of longitudi-
68a 19 dorsal rows at midbody ............ ......... 69 nal stripes, with the pa:i:amedians.running
along the adjacent 7th and 8th dorsal
b 17 dorsal rows at midbody .................... 70
rows, wheri;,as the lateral stripes run
69a Body unicolor or with several dark longitu- along the 5t row ...................... Liophis
dinal stripes on pale background (Figs. b Dorsal scales without apical pits; body
570-575) .................................... Conophis with dark longitudinal stripes, arranged
b Body dark with several pale longitudinal differently than above or unicolor (Figs.
stripes Figs. 576) ................ Crisantophis 557-569) ....... .... Coniophanes (in part)

198

Adelphicos
The genus Adelphicos is represented in
southern Mexico and northern Central
America by six species (CAMPBELL & FORD
1982, CAMPBELL & BRODIE 1988). These are
small and highly secretive snakes. They
are found in rain, cloud, and pine forest at
elevations up to approximately 2200 m
above sea level. By chance, while digging a
hole about 30 cm deep, I came upon a spe-
cimen of Adelphicos quadrivirgatus at Pico
Bonito, Honduras. Otherwise, they are
usually found by turning over rotting logs
or decomposing plant material. LANDY et
al. (1966) reported finding an A quadrivir-
gatus in the leaflitter at the base of a large
tree. These nocturnal snakes feed predom-
inantly on earthworms (LANDY et al. 1966).
The species of this genus reproduce by lay-
ing eggs. A clutch of three A quadrivir-
gatus eggs was found in a nest of tree ter-
mites about one to one and a half meters
above the ground (PEREZ-HIGAREDA &
SMITH 1989). Two weeks later, a snake, 86
mm TL, hatched out of one of the eggs. In
Guatemala, the eggs are laid in the early
rainy season (May and June) (CAMPBELL
1998a).
Fig. 536. Adelphicos nigrilatus (Chiapas,
Mexico). Photo: P. Heimes
Colubridae
Adelphicos daryi CAMPBELL & FORD 1982,
Occ. Pap. Mus. Nat. Hist. Univ. Kansas 100: 3;
type locality: San Jorge Muxbal, 1844 m ele-
vation, 5.5 km W San Jose Pinula, Dpto.
Guatemala, Guatemala. TL to 574 mm.
Southeastern Guatemala, 1830-2135 m eleva-
tion in pine-oak forest.
Adelphicos ibarrorum CAMPBELL & BRODIE
1988, Herpetologica 44: 416; type locality: 3
km SSE Chichicastenango, 2100 m elevation,
in valley between Paxot and Caminibal,
Quiche, Guatemala. TL to 521 mm. Southern
and central Guatemala, 2000-2100 m elevati-
on in pine-oak forest.
Adelphicos latifasciatus LYNCH & SMITH 1966,
Trans. Kansas Acad. Sci. 69: 66; type locality:
Sierra Madre north of Zanatepec, Oaxaca,
Mexico. TL to 437 mm. Western Sierra Madre
de Chiapas in Oaxaca, Mexico, 1500-2000 m
elevation in cloud forest.
Adelphicos nigrilatus SMITH 1942a, Proc.
Rochester Acad. Sci. 8: 182; type locality: San
Cristobal de las Casas, Chiapas, Mexico. TL
to 451 mm. Meseta Central in Chiapas,
Mexico, 2200-2900 m elevation in cloud forest
and pine-oak forest.
Adelphicos quadriuirgatus JAN 1862, Arch.
Zool. Anat. Fis. 2: 18, type locality: Mexico. TL
to 390 mm. Central Veracruz to northern
Honduras on the Caribbean versant; in
eastern Chiapas and western Guatemala on
the Pacific versant. Sea level to 1500 m ele-
vation in rain forest and pine-oak forest.
Adelphicos ueraepacis STUART 1941c, Occ.
Pap. Mus. Zool. Univ. Michigan 452: 5; type
locality: Finca Samac, 7 km W Cohan, Alta
Verapaz, Guatemala. TL to 461 mm. Central
Guatemala (Montaiias de Cuilco, Sierra de
las Minas), 1200-2200 m elevation in cloud
forest and pine-oak forest.
Adelphicos .& A. daryi
17 T A. ibarrorum
• A. latifasciatus
• A. nigrilatus
A. quadrivirgatus
t::. A. veraepacis
199
 Colubridae

Key to Adelphicos
1 a Third infralabial absent or much reduced;
chin shields greatly expanded toward lip
(Fig. 537) .... Adelphicos quadrivirgatus
b Third infralabial barely or not at all re-
duced; chin shields not greatly expanded 2
2 a More than 42 (males) or 32 (females) sub-
caudals; vertebral stripe also encloses
paramedian scale row; venter unpigment-
ed ................... Adelphicos latifasciatus
b Fewer than 42 (males) or 32 (females) sub-
caudals; vertebral stripe, if present, only
on median scale row; venter unpigmented
or not ....... ........ ....... .... ..... ... ................. 3
3 a Venter unpigmented or only darker along
the midline; dorsal ground color pale Fig. 538. Adelphicos quadrivirgatus (Bosawas,
brown or bright red-orange .................. .. . 4 Nicaragua). Photo: J. Sunyer
b Anterior borders of ventral shields conspic-
uously dark; dorsal ground color gray or
brown ................. .............. .. .. ................ 5
4 a 113-128 (males) or 123-135 (females) ven-
trals; ratio of snout length I eye diameter
1.4-2.6 .... .......... .. Adelphicos nigrilatus
b 135-138 ventrals (females) (data for males
not available); ratio of snout length I eye
diameter 2.9-3.2 Adelphicos ibarrorum
5 a Venter not conspicuously darker along
midline; ventrals 120 (only known male) or
128-132 (females); subcaudals 28 (males)
or 19-22 (females); frontal at least as wide
as long ...... ................ ... Adelphicos daryi
b Venter noticeably darker along the mid-
line; ventrals 120-133 (males) or 132-142
(females); subcaudals 29-41 (males) or 24-
31 (females); frontal usually longer than Fig. 539. Adelphicos daryi (Depto. Guatemala,
wide .................... Adelphicos veraepacis Guatemala). Photo: J . A. Campbell

Further Reading
H. SMITH 1942a, CAMPBELL & FORD 1982,
CAMPBELL & BRODIE 1988

Fig. 537. Ventral view of head in Adelphicos

quadrivirgatus; note the conspicuously large Fig. 540. Adelphicos veraepacis (Depto. Baja
chin shields. Verapaz, Guatemala). Photo: J. A. Campbell
200

Fig. 541. Adelphicos quadrivirgatus (Chiapas,
Mexico). Photo: A. Ramirez V.
Amastridium
Contrary to the earlier view of the genus
Amastridium as monotypic (WILSON &
MEYER 1969, WILSON & ROBINSON 1971,
WILSON 1988c), the current view is that
there are two different species, one of
which is distributed in northern Central
America (A sapperi), the other in the
south (A veliferum) (SAVAGE 2002). This
small colubrid is rarely found, thus little is
known about its habits. It is a diurnal gro-
und-dweller in the leaf litter of the rain
forest, but it is also found on coffee planta-
tions (WILSON & MEYER 1969). Sparse
observations on the nutrition of A. velifer-
um indicate that frogs and lizards, as well
as invertebrates, such as centipedes, play a
role in their diet (MARTIN 1955, BLANEY &
BLANEY 1978, CAMPBELL 1998a).
Fig. 542. Amastridium sapperi (holotype).
From WERNER (1903).
Colubridae
Fig. 543. Amastridium veliferum (La Selva,
Costa Rica). Photo: G. Kohler
Amastridium sapperi (WERNER 1903a), Abh.
Bayer. Alcad. Wiss. 22: 349; type locality:
Guatemala. TL to 725 mm. Nuevo Leon and
Oaxaca, Mexico, to Honduras, 100-1000 m
elevation.
Amastridium veliferum COPE 1861a, Proc.
Acad. Nat. Sci. Philadelphia 12: 370; type
locality: Cocuyas de Veraguas, N. Grenada
[= Cocuyas, Panama) . TL to 725 mm. Nica-
ragua to Colombia, 200-800 m elevation.
Key to Amastridium
1 a Fewer than 140 ventrals; no loreal ............ .
........................ Amastridium veliferum
b More than 140 ventrals; loreal present ....
................ ........... Amastridium sapperi
Further Reading
DUNN 1924, WILSON & MEYER 1969, WILSON &
ROBINSON 1971, WILSON 1988c, RUEDA
ALMONACID & RUEDA MARTiNEZ 2004
"" Amastridium sapperi
• Amastridium veliferum
201
 Atractus Atractus hostilitractus MYERS 2003, Amer.
Mus. Novit. 3391: 22; type locality: "Morti
Hydro", 8°52 '28 " N, 77°54'19 ' 'W, about 100-
Only a few of the approximately 70 species 200 m, Rio Morti, Darien, Panama. TL to 374
in the almost exclusively South American mm. Known only from the type locality.
genus Atractus occur in Central America, Atractus imperfectus MYERS 2003, Amer. Mus.
where these are extraordinarily rare. Novit. 3391: 25; type locality: Piedras-Pacora
Recently, MYERS (2003) described four new Ridge, Panama, Panama. Known only from
species of Atractus from Panama. Two of the type locality.
the Panamanian species are known from
the low uplands east of the Canal Zone and
three are from the Darien. The species of
the genus Atractus are small to medium
size fossorial snakes that exhibit shiny
bodies and tiny eyes. This Atractus key is
based on MYERS (2003).

Atractus clarki DUNN & BAILEY 1939, Bull.

Mus. Comp. Zool. 86: 8; type locality: Mine at
Santa Cruz de Cana ["approx. 7°46'N,
77°41 'W, 500 m" according to MYERS 2003],
Darien, Panama. TL to 313 mm. Eastern Abb. 545. A. Atractus clarki (holotype, MCZ
Panama and the Colombia Choc6. 28800); B. A. imperfectus (holotype, MCZ
Atractus darienensis MYERS 2003, Amer. Mus. 50213). Photos: Museum of Comparative
Novit. 3391: 16; type locality: north end of the Zoology, Harvard University. Scale bars = 1 cm
Serrania de Pirre, approx. 8°00'N, 77°43'W,
500 m, Darien, Panama. TL to 346 mm.
Known only from the type locality.
Atractus depressiocellus MYERS 2003, Amer.
Mus. Novit. 3391: 20; type locality: Cerro Azul
[Cerro Jefe], approx. 9°14 'N, 79°23'W, 200-
800 m, Panama, Panama. TL to 750 mm.
Known only from the type.locality.

Abb. 546. A. Atractus darienensis (holotype,

KU 110274); B. A. hostilitractus (holotype,
AMNH 130330). Photos: C. W. Myers

• Atractus clarki
Y Atractus darienensis
v Atractus hostilitractus
• Atractus imperfectus
• Atractus depressiocel/us

Abb. 544. Atractus clarki (holotype, MCZ

28800). Photo: Museum of Comparative
Zoology, Harvard University

202
Key to Atractus Chapinophis
1 a Venter pale; lower scale rows with pale
centers; eye length noticeably greater than
distance to lip ....... ... ...... Atractus clarki The genus Chapinophis, containing the
b Venter black or clouded with dark pig- species C. xanthocheilus, was described in
ment (Fig. 547b); no pale dashes on lower 1998 (CAMPBELL & SMITH 1998). This colu-
scale rows; eye length less than or scarce- brid is a diurnal ground-dweller from the
ly greater than distance to lip ......... ........ 2 cloud forest that so far is known from only
2 a Neck brown like body, with black trans- three specimens.
verse lines not greatly wider than on rest
of body (Fig. 547c); eye very small, contai-
ned about three times in loreal plate
........................ Atractus depressiocellus Chapinophis xanthocheilus CAMPBELL &
SMITH 1998, Herpetologica 54 (2): 210; type
b Neck black with pale rings, or with black locality: 5 km ENE Finca Miranda on trail to
bands several times wider than those Aldea Vega Larga, 2300 m elevation, Baja
posteriorly on body; eye size moderate, Verapaz, Guatemala. TL to 590 mm. Western
going less than 2.5 times in loreal plate 3 portion of Sierra de las Minas in central
3 a Snout rounded in profile; loreal approaching Guatemala, 1830-2300 m elevation in cloud
or touching internasal; labials mostly forest.
black ...... .. ........ Atractus hostilitractus
b Snout bluntly pointed in profile (Fig.
546b); loreal well separated from interna-
sal; labials mostly white .......................... 4
4 a Neck with black saddles (Fig. 547a);
rostral plate narrowly visible in dorsal
view; eye length less than 90% of distance
to lip ..................... Atractus darienensis
b Neck black with pale bars or ringlike
markings; rostral plate well visible in dor-
sal view; eye length about equal to
distance to lip ...... Atractus imperfectus

Further Reading
DUNN & BAILEY 1939, MYERS 2003
Fig. 548. Chapinophis xanthocheilus (near
Chilasco, 1855 m, Sierra de las Minas, Baja
Verapaz, Guatemala). Photo: E. N. Smith

•
\
J
,-
i /f
/ 0--
1"'..r ~ ' • / '-- \r ·

j
Abb. 547. A. and B. Atractus darienensis; J
I
C. Atractus depressiocellus; D. Atractus hostili- _/ ,
T Chapinophis xanthochei/us
tractus (all holotypes). Photos: C. W. Myers J
 Colubridae

Chironius

The species of the genus Chironius are

large, predominantly tree-dwelling, harm-
less colubrids. They are the only
Neotropical snakes with 10 or 12 dorsal
scale rows at midbody. Only one species in
this genus has advanced as far north as
Honduras, whereas the other 12 species
are distributed in South America and in
southern Central America, where three
species occur in Panama and Costa Rica.
For the most part, they live in primary rain
forest, but can also be found in areas with
secondary growth. When they are handled, Fig. 549. Juvenile Chironius grandisquamis
they will usually respond with gaping (Bartola, Rio San Juan, Nicaragua).
mouths and defensive strikes. Photo: G. Kohler

Large eyes with round pupils are charac-

teristic of these diurnal colubrids. Their
primary food source is frogs (mostly tree
frogs), followed by lizards and, less fre-
quently, salamanders (TAYLOR 1951, DIXON
& SOINI 1977' DUELLMAN 1978, DIXON et al.
1993). At night they can be found coiled up
asleep in shrubs and trees. All species in
this genus are oviparous and produce
clutches of 4-15 eggs (DIXON et al. 1993,
GOLDBERG 2007).

Chironius carinatus (LINNAEUS 1758),

Systema Naturae, ed. 10: 223; type locality:
"lndiis." TL to 2055 mm. Costa Rica and Fig. 550. Chironius exoletus (Pavones de
Panama, also distributed across tropical Turrialba, Cartago, Costa Rica).
South America, sea level to 750 m elevation Photo: R. W. Van Devender
in rain forest and on the edges of mangrove
forest. In Central America, the subspecies
Chironius c. flauopictus (WERNER 1910)
occurs.
Chironius exoletus (LINNAEUS 1758), Systema
Naturae, ed. 10: 223; type locality: "Indiis" [in
error] . TL to 1545 mm. Costa Rica and
Panama; also distributed across tropical and
subtropical South America, 300-2170 m ele-
vation in rain forest and cloud forest.
Chironius grandisquamis (PETERS 1868a),
Monatsber. Kong. Akad. Wiss. Berlin 1868:
451; type locality: Costa Rica. TL to 2720 mm.
Northern Honduras to northwestern
Ecuador, sea level to 1600 m elevation in rain
forest and cloud forest . Fig. 551. Chironius carinatus (Prov. Punta-
renas, Costa Rica). Photo: R. W. Van Devender
204
 Colubridae

Key to Chironius
1 a 10 dorsal rows ~t m~dbody ···· ·:······ ·······;·
..................... Chironius grandisquamis
b 12 dorsal rows at midbody ........... .......... 2
2 a Dorsal color olive brown with whitish or
yellow spots on most of the dorsal scales
........ .. ........ ....... ... Chironius carinatus
b Dorsal color unicolor olive brown or olive
green (adult specimens) or with pale nar-
row cross bands running diagonally
(juveniles) ................ Chironius exoletus
Clelia
At present, three species of the genus
Clelia are recognized in Central America.
Of these, C. clelia is differentiated from the
other two species by the presence of 19
rather than 17 dorsal rows at midbody, a
character that is possibly more variable
than previously thought (CAMPBELL
1998a).

Clelia clelia is a very large snake with a

relatively calm temperament. Juveniles
Fig. 552. Adult have completely different coloration (red
Chironius gran- with a yellow neck band) than the adults,
disquamis. which are unicolor shiny black. The eyes
Photo: R. W.
Van Devender are small and dark and the head is not
clearly distinct from the neck. C. clelia
feeds primarily on other snakes, but also
on lizards (predominately basilisks and
ameivas), small mammals, and possibly
Further Reading frogs (STAFFORD 1999). Other potential prey
DIXON et al. 1993, GOLDBERG 2007
items include large pitvipers, as verified by
the case of a 180 cm long Bothrops asper
that was swallowed by a 2 meter long C.
clelia (CAMPBELL 1998a). C. clelia is pre-
dominantly nocturnal; thus, this species is
Chironius grandisquamis seldom seen outside of its hiding place
during the day (STAFFORD 1999a).

=Chironius carinatus
Chironius exoletus

Fig. 553. Juvenile Clelia clelia (KrinKrin,

Nicaragua). Photo: J. Sunyer
205
 Colubridae

Key to Clelia
1 a 19 dorsal rows at midbody ... Clelia clelia
b 17 dorsal rows at mid body ...................... 2
2 a Dorsal scales with two apical pits; ventrals
203-216 (males) or 218-221 (females); sub-
caudals 78-90 (males) or 75-80 (females)
...... .... .......................... .... Clelia scytalina
b Dorsal scales without apical pits; ventrals
202-207 (males) or 200-217 (females); sub-
caudals 75-80 (males) or 54-64 (females)
.... ......... ................ Clelia equatoriana

Fig. 554. Adult Clelia clelia (upper Raspaculo Further Reading

River, Maya Mountains, Belize). GREER 1965, BAILEY 1970a, ScoTr 1983a, ZAHER
Photo: P. Stafford 1996, STAFFORD 1999

Clelia clelia (DAUDIN 1803), Hist Nat. Rept. 6:

330; type locality: Surinam. TL to 2500 mm.
Guatemala and Belize to northwestern
Ecuador as well as to Argentina, sea level to
1000 m elevation. In Central America, the
nominate subspecies occurs.
Clelia equatoriana (AMARAL 1924), J.
Washington Acad. Sci. 14: 201; type locality:
Guayaquil, Ecuador. TL to 1575 mm. Costa
Rica through Panama and Colombia to north-
western Ecuador.
Clelia scytalina (COPE 1867b), Proc. Acad.
Nat. Sci. Philadelphia 18: 320; type locality:
Near Tabasco, Mexico. TL to 1600 mm.
Southern Mexico, Guatemala, and Belize; sea
level to 1000 m elevation.
Fig. 556. Adult Clelia clelia (KrinKrin,
Nicaragua). Photo: J. Sunyer

Clelia clelia
"" Clelia equatoriana
• Clelia scytalina

Fig. 555. Juvenile Clelia scytalina (Tapanti,

Cartago, Costa Rica). Photo: R. W. Van Devender

206

Fig. 557. Coluber c. constrictor (North
Carolina, USA). Photo: P. Kornacker
Coluber
Coluber constrictor is a slender, fast, diur-
nal hunter that prefers savannas and open
dry forest; it feeds on insects (especially as
a juvenile), lizards, frogs, and rodents
(WRIGHT & WRIGHT 1957). This snake
reproduces oviparously, producing clutches
of 2-16 eggs, which hatch in about two
months (FITCH 1963, 1970).
Coluber constrictor LINNAEUS 1758, Systema
Naturae, ed. 10: 216; type locality: "America
septentrionale." TL to 700 mm. Distributed
widely across North America and Mexico; in
Central America south to Belize and north-
ern Guatemala, sea level to 800 m elevation.
In Central America, the subspecies Coluber c.
oaxaca (JAN 1863) occurs.
Further Reading
0RTENBURGER 1928, FITCH 1963, WILSON 1978
Co/uber constrictor
Colubridae
Coniophanes
The species of the predominantly Neo-
tropical genus Coniophanes are small to
medium size colubrids that are highly
secretive. While many species are particu-
larly nocturnal (e.g., C. bipunctatus, C.
quinquevittatus; see CAMPBELL 1998a), at
least C. fissidens, C. imperialis and C. picei-
vittis are also active during the day (MYERS
1969, personal observation). They are
included among the rear-fanged (opistho-
glyphous) snakes, but are usually not
aggressive. Caution is still recommended,
as their bite can lead to localized swelling
and pain (BROWN 1939). Most species are
ground-dwellers in forest areas, while
some (e.g., C. bipunctatus, C. quinquevitta-
tus) are believed to be semi-aquatic (LEE
1996, CAMPBELL 1998a). The Coniophanes
species are known to feed on frogs, sala-
manders, lizards, and snakes, as well as
reptile and bird eggs. They will also con-
sume invertebrates, such as earthworms
and insect larvae (MINTON & SMITH 1960,
LANDY et al. 1966, SEIB 1985, PLATT &
RAINWATER 1998). To escape predators,
these colubrids will throw pieces of their
tail if they are seized. This explains why
many specimens are found with incom-
plete tails (ZUG et al. 1979). The species of
this genus are oviparous and lay 1-10 eggs
per clutch, usually under rotting logs or
roots (H. SMITH 1940, WERLER 1949,
LIVEZEY & PECKHAM 1953, FITCH 1970, ZUG
et al. 1979, CAMPBELL 1998a).
207
 Colubridae

Coniophanes alvarezi CAMPBELL 1989, Proc.

Biol. Soc. Washington 102 (4): 1036; type lo-
cality: 11.3 km ESE Teopisca, 2073 m eleva-
tion, Chiapas, Mexico. TL to 521 mm. In the
area ofTeopisca, Chiapas, Mexico, 2010-2135
m elevation in pine-oak forest.
Coniophanes bipunctatus (GONTHER 1858),
Cat. Snakes Brit. Mus. 1858: 36; type locality:
unknown. TL to 750 mm. Southern Veracruz,
Mexico, to Panama, sea level to approximate-
ly 400 m elevation in rain forest, wet forest
and dry forest.
Coniophanes fissidens (GONTHER 1858), Cat.
Snakes Brit. Mus. 1858: 36; type locality:
Mexico. TL to 795 mm. Southern Veracruz
and Oaxaca, Mexico, through Central Abb. 559. Coniophanes bipunctatus (Catemaco,
America to Colombia and Ecuador, sea level Veracruz, Mexico). Photo: P. Heimes
to approximately 1800 m elevation in rain
and wet forest. In Central America, the sub-
species Coniophanes f. fissidens (along the
Caribbean coast except for the Yucatan
Peninsula), Coniophanes f. obsoletus MINTON
& SMITH 1960 (Volcan de Buenos Aires and
Turrialba, Puntarenas, Costa Rica), and
Coniophanes f. punctigularis COPE 1860d
(along the Pacific coast) occur.
Coniophanes imperialis (KENNICOTT 1859), in
BAIRD: Reptiles of the Boundary: 23; type
locality: Matamoros, Tamaulipas, Mexico. TL
to 550 mm. Southern Texas, USA, along the
Caribbean coast of Mexico to northern
Central America (Yucatan Peninsula and
northern Honduras), sea level to approxim-
ately 2000 m elevation im rain, wet, and dry
forest. In Central America, the subspecies
Coniophanes i. clavatus (PETERS 1864) occurs. Abb. 560. Coniophanes bipunctatus
(Guatemala). Photo: G. Kohler

Fig. 558. Coniophanes alvarezi (El Laurel, Abb. 561. Coniophanes bipunctatus
Chiapas, Mexico). Photo: A. Ramirez V. (Guatemala). Photo: G. Kohler
208

Fig. 562. Coniophanes imperialis (Isla de Utila, Honduras): juvenile (left) and adult (right).
Fig. 563. Coniophanes fissidens (near
Guanagasapa, Escuintla, Guatemala).
Photo: G. Kohler
Fig. 564. Coniophanes joanae (holotype).
Photo: C. W. Myers
Colubridae
Photos: G. Kohler
Coniophanes joanae MYERS 1966, Copeia
1966: 665; type locality: Cerro Pirre, Serrania
de Pirre, Darien, Panama. TL to 369 mm.
Panama east of the canal, 500-1440 m eleva-
tion, in montane rain forest and cloud forest.
Coniophanes meridanus SCHMIDT & ANDREWS
1936, Field Mus. Nat. Hist. Puhl. Zool. Ser. 20:
179; type locality: Merida, Yucatan, Mexico.
TL to 375 mm. Northern half of the Yucatan
Peninsula, sea level to approximately 200 m
elevation in dry forest.
Coniophanes piceivittis COPE 1870, Proc.
Amer. Phil. Soc. 11: 149; type locality:
Chihuitan, Oaxaca, Mexico. TL to 570 mm.
Tamaulipas and Guerrero, Mexico, to Costa
Rica, sea level to approximately 1000 m ele-
vation in dry forest.
Coniophanes quinquevittatus (DUMERIL, ~
BIBRON & DUMERIL 1854), Erp. Gen. 7: 975; ~
type locality: unknown. TL to 500 mm. ·~
Southern Veracruz, Mexico, to northern J3
Guatemala, sea level to approximately 200 m
elevation in rain forest.
Coniophanes schmidti BAILEY 1937, Occ. Pap.
Mus. Zool. Univ. Michigan 362: 1; type locali-
ty: Chichen Itza, Yucatan, Mexico. TL to 400
mm. Northern Chiapas, Mexico, as well as the
Yucatan Peninsula, sea level to approximate-
ly 300 m elevation in dry forest and wet
forest.
209
 Colubridae

Key to Coniophanes b At outer end of each ventral scale a large

1 a 17 Dorsal rows at midbody ............ .. ......... 2 dark spot, such that a symmetrical row of
spots runs along each ventral edge (Fig.
b 19 or more dorsal rows at midbody .......... 3 565b) ............. Coniophanes bipunctatus
2 a More than 70 subcaudals; pale, dark-bor- 8 a Dorsum unicolor brown without longitu-
dered stripe running along side of head at dinal stripes ........ Coniophanes alvarezi
eye height from snout tip to posterior tem-
poral scale; pale neck band, which may be b Dorsum with longitudinal stripes .... ........ 9
incomplete ventromedially, present .. ......... 9 a Pale, dark-bordered stripes running along
......... ........ .. ... Coniophanes meridanus sides of head from snout tip above eye to
b Fewer than 60 subcaudals; pale, dark-bor- posterior temporal s~ales ·········:··········:····:·
dered stripe running along side of head at ...... ...... ....... ... Coniophanes impenalis
mouth height from behind lower eyelid to b Head markings different than those above
neck; without pale neck band ....... . . (Fig. 566a), if a dark-bordered stripe is
..... .. .... .. ... ....... ..... Coniophanes joanae present, it is limited to the temporal region
3 a More than 150 ventrals .............. ... .......... 4 ....... ... .. Coniophanes fissidens (in part)
b Fewer than 150 ventrals ........................... 6
4 a At outer end of each scale a large dark Further Reading
spot, such that a symmetrical row of spots BAILEY 1939, McCoy 1969, MYERS 1969, SEIB
runs along each ventral edge; subcaudals 1985, CAMPBELL 1989, SOLORZANO 2001
63-70 ..... Coniophanes quinquevittatus
b Venter unicolor pale color; subcaudals 78-
115 ................ ............................................. 5
5 a Lateral dark longitudinal stripes at sides
4-5 scales wide, lower edge color sharply
contrasting from colo~ of venter· ·;····;··:····;·
... ..... ..... ........... Coniophanes picewittis
b Lateral dark longitudinal stripes 0.5-1.5
scales wide, lower edge color grades into
ventral coloration Coniophanes schmidti
6 a 21 dorsal rows at mid body ..... .... ......... .. .... 7
b 19 dorsal rows at midbody .... .......... ........ 8
7 a Venter unicolor pale or with irregularly
distributed fine dark spots (Fig. 565a) .......
............ Coniophanes fissidens (in part)

Coniophanes bipunctatus

, A Coniophanes alvarezi
"' Coniophanes joanae

_ Coniophanes
Coniophanes fissidens = quinquevittatus
• Coniophanes meridanus Coniophanes imperialis

210
 Colubridae

• • ..
a. C. fissidens
-
•

. - - - --
b. C. bipunctatus

Fig. 565. Ventral pattern in Coniophanes .

Fig. 567. Coniophanes meridanus (Chetumal,

Qintana Roo, Mexico).
Photo: P. Charruau & J . R. Cedeno-Vazquez

Fig. 566. Lateral head in Coniophanes.

Drawings: M. Vesely

Fig. 568. Coniophanes piceivittis (Volcan

Maderas, Isla Ometepe, Nicaragua).
Photo: G. Kohler

Coniophanes piceivittis Fig. 569. Coniophanes schmidti (near

• Coniophanes schmidti
Macanche, Peten, Guatemala).
Photo: G. Kohler
211
 Colubridae
Conophis
Recently, P:EREZ-HIGAREDA et al. (2002) pro-
posed to restrict the usage of the name
Conophis lineatus to the populations in
central Veracruz and elevate the Central
American populations to species level,
assigning the name C. concolor to them.
However, because their proposal is not sub-
stantiated by data or specimens examined,
I prefer to use the name C. lineatus for all
populations until a comprehensive revision
of Conophis has been published.
Both species of the genus Conophis are
medium-sized, very irritable rear-fanged
snakes, with a very painful bite. They are
diurnal and hunt mostly lizards
(Sceloporus and Aspidoscelis) and other
species of snakes, which are injected with
powerful venom when bitten by the long
rear fangs of these predators. Although the
bite of a Conophis is not critically dange-
rous to humans, it can still cause painful
localized swelling and bleeding (MERTENS
1952c, WELLMAN 1963, SCOTT 1983b,
JOHNSON 1988, LEE 1996). Captive-held
Conophis will eat frogs and toads
Fig. 570. Conophis lineatus (Guanacaste
National Park, Costa Rica). Photo: M. Franzen
212
(MITTLEMAN 1944), as well as snakes
(Thamnophis and Storeria). Rodents are
also part of the diet of these snakes, as
demonstrated by the finding of a mouse in
the stomach of a dissected C. lineatus
(WELLMAN 1963). Both species reproduce
oviparously, with a clutch size of 5 to 10
eggs (ALVAREZ DEL TORO 1960, LEE 1996,
personal observation).
Conophis lineatus (DUMERIL, BIBRON &
DUMERIL 1854), Erp. Gen. 7: 936; type locali-
ty: Mexico. TL to 1170 mm. Oaxaca and
Veracruz, Mexico, as far as Costa Rica, sea
level to approximately 1000 m elevation in
dry forest and savanna. In Central America,
the subspecies Conophis l. concolor COPE
1867 (Yucatan Peninsula) and Conophis l.
dunni SMITH 1942b (southeastern Oaxaca,
Mexico, to Costa Rica) occur. Conophis pul-
cher COPE 1869 is regarded to be a synonym
of C. lineatus according to WILSON & MEYER
(1985).
Conophis vittatus PETERS 1860, Monatsber.
Kong. Akad. Wiss. Berlin 1860: 519; type lo-
cality: unknown. TL to 840 mm. Pacific side of
Mexico from southern Durango as far south
as Chiapas, sea level to approximately 1000
m elevation in dry forest and savanna.
Fig. 571 Conophis vittatus (near Mixtequilla,
Oaxaca, Mexico). Photo: G. Kohler
 Colubridae

Key to Conophis
1 a Body with 3 or 4 longitudinal black stripes,
each at least two scald rows tff de; usually
7 supralabials, the 3r and 4t of these in
contact with eye ....... . Conophis vittatus
b Body with 6 to 10, rarely 4, longitudinal
black stripes, that are usually less than
two scale rows wide, or entirely without
longitudinal strifiis; usual~ 8 supralabi-
als, with the 4 and 5t of these in
contact with eye ....... Conophis lineatus

Fig. 574. Conophis lineatus (Morgans Rock,

Nicaragua). Photo: J. Sunyer

Fig. 572. Conophis lineatus (nr. Sta. Cruz, Fig. 575. Conophis lineatus (Valle de Otoro,
Zacapa, Guatemala). Photo: G. Kohler Honduras). Photo: G. Kohler

• Conophis lineatus
_ Conophis vittatus

Further Reading
Fig. 573. Conophis lineatus (Rus Rus, H. SMITH 1941a, 1942b, SAVAGE 1949, WELLMAN
Honduras). Photo: J. R. McCranie 1963, SCOTT 1983b
213
 Colubridae
Crisantophis
Crisantophis nevermanni was earlier
included in the genus Conophis (WELLMAN
1963), but at the beginning of the 1970's,
was placed in its own genus (VILLA 1971).
This fast colubrid lives in dry areas, prefer-
ably close to a source of water where it can
hunt frogs and lizards.
Crisantophis nevermanni (DUNN 1937a),
Copeia 1937: 214; type locality: Rio Poas de
Aserri [= a few miles south of San Jose],
Costa Rica. TL to 1007 mm. Southeastern
Guatemala to Costa Rica; sea level to approx-
imately 1250 m elevation in dry forest and
savanna.
Fig. 576. Crisantophis nevermanni feeding on
a race runner (Ometepe, Nicaragua).
Photo: P. van Beest
• Crisantophis nevermanni
Further Reading
WELLMAN 1963, VILLA 1969b, 1971, 1988,
SOL6RZANO 2003
214
Dendrophidion
These medium-sized, slender colubrids are
distributed from southern Mexico through
Central America and far into tropical
South America. They are diurnal forest-
dwellers that feed primarily on lizards and
frogs. Dendrophidion species reproduce
oviparously; females of D. percarinatum
and D. vinitor in lower Central America
have an extended or possibly continuous
reproductive cycle and may produce more
than one clutch per year. Clutch sizes
range from 3-5 in D. vinitor to 4-6 in D. per-
carinatum and 5-8 in D. nuchale (STAFFORD
2003).
Dendrophidion nuchale (PETERS 1863c),
Monatsber. Kong. Akad. Wiss. Berlin 1863:
285; type locality: Caracas, Venezuela. TL to
1530 mm. Belize and central Guatemala to
northwestern Ecuador, 125-1500 m elevation
in rain forest. Recently demonstrated to occur
in Nicaragua (STAFFORD 2002). Dendrophi-
dion clarki DUNN 1933a is regarded to be a
synonym of D. nuchale according to LIEB
(1988).
Dendrophidion paucicarinatum (COPE 1894),
Proc. Acad. Nat. Sci. Philadelphia 46: 202;
type locality: La Candelaria, Bruno
Carranza, Costa Rica. TL to 1380 mm .
Cordillera Central in Costa Rica and
Cordillera de Talamanca in Costa Rica and in
western Panama, 1000-1600 m elevation in
montane rain forest and cloud forest.
Fig. 577. Dendrophidion percarinatum (Rio
Patuca, Olancho, Honduras). Photo: G. Kohler
 Colubridae

Dendrophidion percarinatum (COPE 1893),

Proc. Amer. Philos. Soc. 31: 344; type locality:
Boruca and Buenos Aires, Costa Rica. TL to
1130 mm. Northern Honduras to north-
western Ecuador, 30-500 m elevation in rain
forest.
Dendrophidion vmitor SMITH 194ld, Proc.
Biol. Soc. Washington 54: 74; type locality:
Piedras Negras, Guatemala. TL to 980 mm.
Southern Veracruz, Mexico, to western
Colombia, 50-1300 m elevation in rain forest.

Key to Dendrophidion
1 a Reduction of scales on dorsal side of tail
froff: eight to six rows occurs before the
25t subcaudal scale ......... ..... .............. 2
Fig. 578. Dendrophidion paucicarinatum
b Reduction of scales on the dorsal side of (Fortuna, Panama). Photo: G. Kohler
taibfrom eight to six rows occurs after the
25t subcaudal scale ............... .. ........... 3
2 a 179-195 ventrals; 119-139 subcaudals ... .
... . . . ... Dendrophidion paucicarinatum
b 153-166 ventrals; 147-163 subcaudals
............. Dendrophidion percarinatum
3 a 132-163 subcaudals; cloaca! scute usually
divided; first dorsal scale row strongly
keeled; dark-bordered pale cross bands on
neck, if present, less than one scale row
wide; primary color of tail abruptly paler
than that of body ............. ...... .. .... ... .
. ............. ... . . . . . . . Dendrophidion nuchale
b 111-128 subcaudals; cloacal scute undivi-
ded; first dorsal scale row smooth or only
slightly keeled; dark-bordered pale cross
bands on neck more than one scale row
wide; primary color of tail not paler than
that of body ....... Dendrophidion vinitor

Further Reading Fig. 579. Dendrophidion nuchale (Cuxta Bani,

LIEB 1988, 1991a-c, 1996 upper Raspaculo River, Cayo District, Belize).
Photo: P. Stafford

Dendrophidion nuchale Dendrophidion percarinatum

=: Dendrophidion paucicarinatum

215
 Colubridae

Diaphorolepis

The poorly known genus Diaphorolepis

consists of two species, of which one, D.
wagneri, occurs in eastern Panama.
Nothing is known about the habits of this
species.

Diaphorolepis wagneri JAN 1863b, Elenco

Sistema Ofidi: 98, type locality: Andes of
Ecuador. TL to 600 mm. Known to occur in
Ecuador and in eastern Panama.

Fig. 580. Dendrophidion vinitor (north of El Further Reading

Cope, Cocle, Panama). Photo: C. W. Myers BOGERT 1964

Fig. 581. Dendrophidion vinitor (Rio San Juan,

Nicaragua). Photo: D. M. Dehling
Fig. 583. Diaphorolepis wagneri. From BOGERT
(1964).

• Dendrophidion vinitor
.& Diaphorolepis wagneri

Fig. 582. Dendrophidion percarinatum (Los

Cusingos, Costa Rica). Photo: G. Kohler
216

Dipsas
Approximately 30 species are included in
the genus Dipsas, eight of which occur in
Central America. Recently, D. nicholsi (for-
merly treated as a subspecies of D. varie-
gata) has been raised to species level
(CADLE & MYERS 2003). The species of
Dipsas are small to mid-size, harmless
colubrids that can be found at dusk and at
night crawling around in vegetation, parti-
cularly in the rain. They feed (exclusively?)
on snails and slugs. These small colubrids
have been observed flicking their tongues
and following the slime trail of snails over
several meters (SAZIMA 1989). Slugs are
grabbed in the middle, pulled out of the
substrate and usually swallowed tail first.
Snails are also pulled out of the substrate
first, and held tightly in a coil. The snake
will grab the foot of the snail with the teeth
of the lower jaw, press the shell against the
upper jaw and then use the lower jaw to
pull the soft body parts out of the shell. Up
to a third of the snake's lower jaw is shoved
into the shell during this procedure. After
the snail is pulled out, the snake will drop
the shell and swallow the soft body parts.
All Dipsas species are oviparous and pro-
duce clutches of 1-5 (usually 2-3) eggs
(FITCH 1985, SCHUMACHER 1996, CAMPBELL
1998a). In captivity, the eggs of Dipsas arti-
culata require just three months to hatch.
(SCHUMACHER 1996 [misidentified as D.
tenuissima]).
Dipsas
A D. temporalis
,,. D. nicholsi
.._ D. articulata
Fig. 584. Dipsas bicolor (Siquirres, Costa
Rica).
Colubridae
Dipsas articulata (COPE 1868), Proc. Acad.
Nat. Sci. Philadelphia 20: 135; type locality:
Veraguas, "Costa Rica" [actually in Panama].
TL to 712 mm. Southeastern Nicaragua to
western Panama, sea level to approximately
500 m elevation in rain forest.
Dipsas bicolor (GUNTHER 1895), Biologia
Centrali-Americana. Reptilia and Batrachia:
178; type locality: Chontales Mines,
Nicaragua. TL to 482 mm. Northeastern
Honduras to northeastern Costa Rica, sea
level to approximately 500 m elevation in
rain forest.
Dipsas brevifacies (COPE 1866), Proc. Acad.
Nat. Sci. Philadelphia 18: 127; type locality:
Yucatan, Mexico. TL to 528 mm. Northern
half of the Yucatan Peninsula (southern
Campeche, Mexico, and central Belize north-
wards), sea level to approximately 300 m ele-
vation in dry forest.
Dipsas maxillaris (WERNER 1909), Zool.
Jahrb. 28: 279; type locality: Tabasco, Mexico.
TL to 405 mm. Tabasco, Mexico, sea level to
100 m elevation in rain forest.
Dipsas nicholsi (DUNN 1933), Copeia 1933:
193; type locality: "Mid-basin of Chages River
and mouth of Pequeni River, Panama" [=jun-
ction of Rio Pequeni with Rio Boquer6n, in
upper drainage of Rio Chagres at 9°2l'N,
79°33'W according to CADLE & MYERS 2003].
TL to 821 mm. Disjunct populations in
Panama, sea level to 855 m elevation in rain
forest.
Photo: R. D. Bartlett
217
 Colubridae

Fig. 585. Dipsas brevifacies (Quintana Roo, Fig. 587. Dipsas brevifacies (Corrillo Puerto,
Mexico). Photo: J. C. Lee Quintana Roo, Mexico). Photo: A. Ramirez V.

Dipsas temporalis (WERNER 1910), Mitt.

Naturhist. Mus. Hamburg 26: 241; type lo-
cality: Esmeralsdas, Ecuador. TL to 680 mm.
Central Panama to Ecuador.
Dipsas tenuissima TAYLOR 1954, Univ.
Kansas Sci. Bull. 36: 771; type locality: on
Dominica} Road in swamp, approx. 15. km
WSW San Isidro del General, Costa Rica. TL
to 555 mm. Pacific side of southern Costa
Rica and of western Panama, sea level to
approximately 500 m elevation in rain forest.
Dipsas viguieri (BOCOURT 1890), Bull. Soc.
Philom. Paris (7) 8: 136; type locality:
Isthmus of Darien, Panama. TL to 637 mm. Fig. 588. Dipsas temporalis (Rio Jaque I Rio
Eastern Panama and western Colombia. Imamad6, Darien, Panama).
Photo: C. W. Myers

Fig. 586. Dipsas bicolor (Bosawas, Nicaragua). Fig. 589. Dipsas articulata (Cahuita, Costa
Photo: S. Travers Rica). Photo: R. W. Van Devender

218
 Colubridae

Key to Dipsas
1 a Dorsum with large lateral dark brown
spots that alternate and do not meet along
the vertebral line ..... ....... Dipsas nicholsi
b Dorsum with regular dark brown bands
that reach from one side to the other .. .... 2
2 a Two or three large unpaired chin shields
(Fig. 590a) .............. .......... Dipsas bicolor
b Two or three pairs of chin shields (Fig.
590b) ............... ............ ............ ... .............. . 3
3 a Fewer than 188 ventrals; fewer than 101
subcaudals ........................ ........ ............... . 4
b More than 188 ventrals; more than 101
subcaudals ............................. ................. .. 5
4 a 8 or more supralabials ...................... .. .. ... .
. .. . . . . ... .... .... .. ..... ........ Dipsas brevifacies
b Fewer than 8 supralabials ............. .. ... .. ... .
. . . . . .. . . . .. . . .... . .............. Dipsas maxillaris
5 a Mental shield in contact with first pair of
chin shields (first pair of infralabials not in
contact with each other) .... .. ..... ...... . . Fig. 591. Dipsas articulata (Bartola, Rio San
..... .... ... .... ... .. .... ... ... Dipsas temporalis Juan, Nicaragua). Photo: G. Kohler
b First pair of infralabials in contact with
each other, thereby separating mental
shield from first pair of chin shields .... .... 6
6 a No preoculars; usually 2 postoculars .........
....... ... . .. . . . .. . . . . . ........... Dipsas articulata
b One preocular; usually 3 postoculars ....... 7
7 a More than 210 ventrals; dorsal surface of
head unicolor dark brown ......... ...... .. ...... .
.. . .. . . . . . . . . . .. . .. . . . . . . ...... Dipsas tenuissima
b Fewer than 210 ventrals; dorsal surface of
head with fine white spottin.g ····· ···;· ···; ··· :
...... ................. .......... ... .. Dipsas viguien

Fig. 592. Dipsas articulata (near Santa Fe,

Panama). Photo: S. Lotzkat

b. 0. articulata

Further Reading
Fig. 590. Scalation of chin region in Dipsas PETERS 1960, KOFRON 1982, KOHLER & VIEL-
(chin shields orange). METTER 2002, CADLE & MYERS 2003, MYERS 2007
219
 Colubridae

Drymarchon

Indigo snakes (Drymarchon) comprise a

widespread group of large colubrid snakes,
distributed in south-eastern North
America, Central America and parts of
South America. Throughout most of the
20th century, Drymarchon has been con-
sidered monotypic (e.g., AMARAL 1929c,
MCCRANIE 1980). Recently, two former sub-
species (i.e., couperi and melanurus) were
raised to the status of full species and a
new species, D. caudomaculatus, was Fig. 593. Drymarchon corais unicolor (Santa
described from Venezuela (COLLINS 1991, Rosa National Park, Guanacaste, Costa Rica).
Photo: M. Franzen
WOSTER et al. 2001). However, I prefer to
recognize melanurus as a subspecies of D.
corais until the systematics of the Indigo
snakes is better understood.

Indigo snakes are quick, diurnal colubrids

that feed on a variety of vertebrates (e.g.,
fish, frogs, turtles, lizards, rodents, and
small birds). Drymarchon corais reprodu-
ces oviparously, producing clutches of 4-11
eggs, which hatch in 70-80 days (KOHLER
1997).

Drymarchon corais (BOIE 1827), Isis von

Oken 1827: 537; type locality: America. TL to
2950 mm. Southern USA on the Caribbean
side and Sinaloa, Mexico, on the Pacific side
to Argentina, sea level to approximately 1500 Fig. 594. Drymarchon corais melanurus (Isla
m elevation in dry forest, wet forest, and rain de Roatan, Honduras). Photo: G. Kohler
forest. In Central America, the subspecies
Drymarchon c. melanurus (DUMERIL, BIBRON
& DUMERIL 1854) (Veracruz, Mexico, along
the Caribbean side to northwestern South
America; in Costa Rica and Panama also on
the Pacific side), Drymarchon c. rubidus
SMITH 1941c (Sinaloa, Mexico, along the
Pacific side to the Isthmus of Tehuantepec,
Mexico, as well as in Rio Grijalva-Basin of
Chiapas) and Drymarchon c. unicolor SMITH
1941c (Chiapas, Mexico, along the Pacific side
to northwestern Costa Rica) occur.

Further Reading
AMARAL 1929c, MCCRANIE 1980

Fig. 595. Drymarchon corais unicolor (Morgans

Rock, Rivas, Nicaragua). Photo: J. Sunyer
220
 Colubridae

Drymobius

The genus Drymobius includes only a few

Fig. 596. Drymarchon corais unicolor (San
Marcos Palajunoj, Quetzaltenango, Guate-
mala). Photo: G. Kohler
species that are distributed in Mexico,
Central America, and northwestern South
America. They are slender, quick, and
usually rather irritable snakes that are
diurnal. They live as ground-dwellers in
savanna and forest, in both primary and
secondary vegetation. Whereas D. chlorot-
icus is mostly a highland species, the
others live predominantly in the lowlands,
usually close to a body of water. Despite its
bright green coloration, D. chloroticus is
quite difficult to spot amongst the debris
and contours of the forest floor. Although
their diet consists primarily of frogs and
toads, (DUELLMAN 1963, HENDERSON &
HOEVERS 1977b), they will also eat lizards,
reptile eggs and rodents. All Drymobius
species are oviparous. Drymobius margarit-
iferus produces 4-7 eggs per clutch, with
young hatching in 64-68 days (WERLER
1949, 80L6RZANO & CERDAS 1987). In
Guatemala, freshly hatched snakes of this
species can be seen from June until
October (CAMPBELL 1998a).

Fig. 597. Drymarchon corais melanurus

(Bosawas, Nicaragua). Photo: J. Sunyer

Fig. 598. Drymobius melanotropis (Cerro El

Drymarchon corais Toro, Atlantico Norte, Nicaragua).
Photo: G. Kohler
221
 Colubridae

Fig. 600. Drymobius rhombifer (Ecuador). Fig. 602. Drymobius margaritiferus (Bartola,
Photo: G. Kohler Rio San Juan, Nicaragua). Photo: G. Kohler

Fig. 599. Drymobius chloroticus (Selva Negra, Fig. 603. Drymobius margaritiferus (Los
Matagalpa, Nicaragua). Photo: G. Kohler Guatuzos, Nicaragua). Photo: J . Sunyer

Fig. 601. Drymobius chloroticus (Selva Negra, Fig. 604. Drymobius margaritiferus (David,
Matagalpa, Nicaragua). Photo: G. Kohler Panama). Photo: J. Sunyer
222

Drymobius chloroticus (COPE 1886a), Proc.
Amer. Phil. Soc. 23: 278; type locality:
Guatemala; restricted by COPE (1887) to
Cohan, Alta Verapaz, Guatemala. TL to 1200
mm. San Luis Potosi and Oaxaca, Mexico, to
central Nicaragua, 500-2500 m elevation in
pine-oak forest, montane rain forest and
cloud forest.
Drymobius margaritiferus (SCHLEGEL 1837),
Essai Physion. Serpens 2: 184; type locality:
"New Orleans" [in error]. TL to 1340 mm.
Southern Texas, USA, and Sonora, Mexico, as
far as Colombia, sea level to 2000 m elevation
in savanna, wet and rain forest. In Central
America, the subspecies Drymobius m. mar-
garitiferus (Texas, USA, along the Caribbean
coast as far as South America), Drymobius m.
maydis VILLA 1968 (Great Corn Island,
Nicaragua) and Drymobius m. occidentalis
BocoURT 1890 (Chiapas, Mexico, along the
Pacific coast to El Salvador) occur.
Drymobius melanotropis (COPE 1876), Journ.
Amer. Phil. Soc. (2) 8: 134; type locality: Costa
Rica. TL to 760 mm. Eastern Honduras to
southeastern Costa Rica, sea level to 1400 m
elevation in rain forest.
Drymobius rhombifer (GONTHER 1860), Proc.
Zool. Soc. London 1860: 236; type locality:
Esmeraldas, Ecuador. TL to 1270 mm.
Nicaragua, Costa Rica, and Panama, as well
as northern South America (Peru, Ecuador,
and Venezuela).
Drymobius chloroticus
• Drymobius melanotropis
Colubridae
Key to Drymobius
1 a Dorsum green in life (blue in preservative),
without large dark blotches or yellow-
green spots ............. ...... ............................. 2
b Dorsum with large dark blotches or yellow-
green spots ................... .... .. ... ........... .... 3
2 a 107-125 subcaudals; keels of the middle
three dorsal scale rows not black ............... .
............ .... .. ......... Drymobius chloroticus
b 91-94 subcaudals; keels of the middle three
dorsal scale rows black ...... ....................... .. .
............. ......... Drymobius melanotropis
3 a On each dorsal scale, a small yellow or yel-
low-green spot, so that the dorsum appears
covered with fine speckles; 103-138 sub-
caudals ........ Drymobius margaritiferus
b Dorsal pattern consists of large dark
blotches; 84-102 subcaudals .. ............. ..... .
.. ....... ... .... ........... Drymobius rhombifer
Further Reading
WILSON 1970, 1974, 1975a-c, KOHLER 2003
Drymobius margaritiferus
=Drymobius rhombifer
223
 Colubridae

Elaphe

The species of the genus Elaphe are

distributed predominantly in the Northern
Hemisphere. Many of them have very
attractive markings, as does the yellow-red
rat snake (Elaphe fiavirufa), which occurs
in Central America. This species is semi-
arboreal and lives in savanna, wet forest
and dry forest, hunting at night predom-
inantly for rats, but also birds and lizards.
In captivity, a female produced a clutch of
five eggs, which was incubated at 28 °C
and hatched within 65-66 days (ENTZEROTH Fig. 606. Elaphe flauirufa pardalina (Calak-
1991). mul, Campeche, Mexico). Photo: H. Bahena B.

Elaphe flauirufa (COPE 1867), Proc. Acad.

Nat. Sci. Philadelphia 19: 319; type locality:
Yucatan. TL to 1650 mm. Tamaulipas and
Chiapas, Mexico, to Nicaragua, sea level to
1500 m elevation in seasonal rain and dry
forest. In Central America, the subspecies
Elaphe fiauirufa flauirufa (Tamaulipas,
Mexico, to the base of the Yucatan Peninsula),
Elaphe f pardalina (PETERS 1868b) (Alta
Verapaz, Guatemala, and Belize to Nicara-
gua), Elaphe f matudai SMITH 1941e (Pacific
versant of Oaxaca and Chiapas, Mexico), and
Elaphe f phaescens DOWLING 1952 (the north
of the Yucatan Peninsula) occur.

Further Reading
DOWLING 1952, ENTZEROTH 1991, SCHULZ 1996
Fig. 607. Elaphe flauirufa phaescens (Cancun,
Quintana Roo, Mexico). Photo: A. Ramirez V.

• Elaphe flavirufa
Fig. 605. Elaphe flauirufa pardalina (Isla de = Enuliophis sclateri
Utila, Honduras). Photo: G. Kohler
224

Enuliophis
The genus Enuliophis was recently estab-
lished for the species sclateri which was
previously included in the genus Enulius
(MCCRANIE & VILLA 1993). Not much is
known about this species other than that it
is highly secretive and lives in savanna,
dry forest and wet forest (TAYLOR 1951,
PEREZ-SANTOS & MORENO 1988). It has
been described as both diurnal and noctur-
nal; two specimens were active in leaf lit-
ter at night (MCCRANIE 2004b).
Enuliophis sclateri (BOULENGER 1894a), Cat.
Snakes Brit. Mus. 2: 251; type locality: South
America. TL to 504 mm. northeastern
Honduras to Colombia, sea level to 1285 m in
savanna, dry forest, and wet forest.
Further Reading
DUNN 1938a, MCCRANIE & VILLA 1993,
MCCRANIE 2004b
Fig. 608. Enuliophis sclateri (La Selva,
Heredia, Costa Rica). Photos: G. Kohler
Colubridae
Enulius
The species of the genus Enulius are small,
harmless colubrids that are highly secre-
tive. They inhabit the rotten limbs of dead
trees and coconut palms, which termites
have left sponge-like, riddled with holes
that make ideal hiding places for the little
snakes. Termites and ants are their pri-
mary source of food, although SCOTT
(1983c) maintains that E. flavitorques lives
on a specialized diet of reptile eggs.
Enulius bifoveatus MCCRANIE & KOHLER
1999b, Carib. J. Sci. 35: 15; type locality: be-
tween Savannah Bight and East End,
l6°29.19'N, 85°50.30'W, less than 10 m eleva-
tion, Isla de Guanaja, Islas de la Bahia,
Honduras. TL to 321 mm. Isla de Guanaja,
Honduras, sea level to 50 m elevation in wet
meadow and wet forest.
Enulius fiavitorques (COPE 1869), Proc. Acad.
Nat. Sci. Philadelphia 20: 307; type locality:
Rio Magdalena, Colombia. TL to 495 mm.
Southern Jalisco, Mexico, as far as Colombia,
sea level to approximately 1000 m elevation
in rain forest, wet forest and dry forest. In
Central America, the subspecies Enulius f.
fiavitorques (Guatemala to Colombia) and
Enulius f. sumichrasti BocoURT 1883 (south-
eastern Oaxaca and southern Chiapas,
Mexico) occur.
Enulius roatanensis MCCRANIE & KOHLER
1999b, Carib. J. Sci. 35: 17; type locality: near
Mud Hole Bay, l6°20.88'N, 86°32.05'W, less
than 10 m elevation, Isla de Roatan, Islas de
la Bahia, Honduras. TL to 346 mm. Isla de
Roatan, Honduras, sea level to 50 m elevation
in wet meadow and wet forest .
.& Enulius bifoveatus
"" Enulius roatanensis
Enulius flavitorques
225
 Colubridae

Key to Enulius
1 a Dorsal scales with two apical pits; pale
neck band interrupted or fragmented (Fig.
609b) ......................... Enulius bifoveatus
b Dorsal scales with one apical scale pit; pale
neck band complete or entirely absent .... 2
2 a Pale neck band present (Fig. 610); ventrals
166-190 in males, 184-216 in females; sub-
caudals 100-117 in males ................. ..
. . . . . . . . ..... . . . . . . . ......... Enulius f7.avitorques
b Without pale neck band (Fig. 609a); ventrals
165 in only known male, 176 in only known
female; subcaudals 121 in male ................. ..
. . . .. .................. .......... Enulius roatanensis
Fig. 611. Enulius bifoveatus (Isla de Guanaja,
Honduras). Photo: G. Kohler
Further Reading
DUNN 1938a, SMITH et al. 1967, MCCRANIE &
KOHLER 1999b

a. E. roatanensis b. E. bifoveatus
Fig. 612. Enulius roatanensis (Isla de Roatan,
Fig. 609. Dorsal head view. Photos: G. Kohler Honduras). Photo: G. Kohler

Fig. 610. Enulius fiavitorques (Ometepe, Fig. 613. Enulius fiavitorques (Costa Rica).
Nicaragua). Photo: J. Sunyer Photo: R. W. Van Devender

226
 Colubridae

Erythrolamprus

The false coral snakes of the genus

Erythrolamprus (altogether 7-8 species)
are conspicuous for their remarkable color-
ation. They are diurnal ground-dwellers
that feed on snakes, frogs, lizards, and
small mammals (SAZIMA & ABE 1991).
Their clutches of 6-9 eggs are hidden under
rotten logs, decomposing plant material, or
in abandoned rodent dens (J. HARDY &
Boos 1995).

Erythrolamprus bizona JAN 1863a, Arch. Fig. 614. Erythrolamprus mimus (Montana La
Zool. Anat. Fis. 2: 314; type locality: "Bahia, Galia, Jinotega, Nicaragua). Photo: G. Kohler
Messico, Popayan, Cayenne, Brasile,
Montevideo, Colombia". TL to 1000 mm.
Costa Rica to Colombia and Venezuela, sea
level to approximately 1500 m elevation (in
Colombia also to 2600 m elevation) in rain
forest.
Erythrolamprus mimus (COPE 1869), Proc.
Acad. Nat. Sci. Philadelphia 20: 307; type
locality: "High regions of Ecuador or New
Grenada." TL to 1000 mm. Honduras to Peru,
500-1300 m elevation (in Colombia also to
2000 m elevation) in rain forest. In Central
America, the subspecies Erythrolamprus m.
impar SCHMIDT 1936a (Honduras to Costa
Rica) and Erythrolamprus m. micrurus DUNN
& BAILEY 1939 (Panama to western Ecuador)
occur.

Key to Erythrolamprus
Fig. 615. Erythrolamprus bizona (Valle de
1 a A solid broad pale (orange, yellow or white) Anton, Panama). Photo: G. Kohler
band across parietal region; one black neck
band, in some specimens partly divided by
one pale band; black body rings single, may
be divided laterally by a pale band and
have a tendency to become offset along ver-
tebral line; infralabials and scales in thro-
at area without black pigment; 42-51 sub-
caudals ............. Erythrolamprus mimus
b Pale parietal band narrow or broken into
blotches; two black neck bands, completely
divided by one pale neck band; paired
black body rings not offset along vertebral
line; infralabials and scales in throat regi-
on with black pigment; 51-60 subcaudals
........................... Erythrolamprus bizona

Erythrolamprus mimus
Further Reading =: Erythrolamprus bizona
J . HARDY & Boos 1995
227
 Colubridae
Fig. 616. Ficimia publia (near La Ceiba,
Atlantida, Honduras). Photo: G. Kohler
Ffoimia
The genus Ficimia contains six species,
two of which occur in Central America, and
the remainder further north (L. HARDY
1975). The species of this genus are
distinguished by a rostral shield that is
upturned and pointed. They are completely
harmless, nocturnal ground-dwellers that
feed primarily on spiders (LEE 1996).
Ficimia publia is oviparous, as demonstrat-
ed in the report by GREER (1966) of a fe-
male that laid two eggs.
228
Ficimia publia COPE 1866, Proc. Acad. Nat.
Sci. Philadelphia 18: 126; type locality:
Yucatan, Mexico. TL to 461 mm. Northern
Veracruz, Mexico, to Honduras on the
Caribbean side and Guerrero, Mexico, to
Guatemala on the Pacific side, sea level to
approximately 1000 m elevation in dry forest
and wet forest.
Ficimia ramirezi SMITH & LANGEBARTEL 1950,
J. WashingtonAcad. Sci. 39: 411; type locality:
1 league [approx. 3.8-7.4 km] north of
Niltepec, Oaxaca, Mexico. TL to 313 mm.
Sierra Madre de Chiapas (Chiapas and
southeastern Oaxaca, Mexico), 500-1000 m
elevation in dry forest and pine-oak forest.
Key to Ficimia
1 a Intervals between dark spots less than
twice as wide as the diame~e~ o~ the spo.ts
..... ....... ...... ...... .. .. ......... Ficimia publia
b Intervals between dark spots at least twice
as wide as the diameter of t~e ;;pots ···:······:
.... .. .......... ... .... ........... Ficimia ramirezi
Further Reading
H. SMITH & TAYLOR 1941, L. HARDY 1975, 1979,
1980, 1990
,.. Ficimia ramirezi
Ficimia publia
Fig. 617. Ficimia publia (near La Ceiba,
Atlantida, Honduras). Photo: G. Kohler
 Colubridae

Geophis

The species of the genus Geophis are small,

harmless colubrids that are distributed
across Mexico, Central America, and north-
western South America. In Central
America, the genus is well represented by
19 species. The diet of these secretive
snakes consists of earthworms, slugs, and
small arthropods (CAMPBELL & MURPHY
1977, CAMPBELL et al. 1983). At the
Biosphere Preserve Bosawas, Nicaragua, I
Fig. 618. Geophis bellus (holotype).
found two Geophis hoffmanni under a Photo: C. W. Myers
large rotting log in the rain forest. It is
otherwise reported that most specimens of
this genus are found under rocks, roots, or
tree branches, and rarely in the open
(CAMPBELL & MURPHY 1977, CAMPBELL et
al.1983). All Geophis species are believed
to be egg-layers, with a reported clutch size
of 3-6 eggs documented for G. brachyceph-
alus (SASA 1993). The eggs of G. brachy-
cephalus incubated at 25-28 °C took 109
days to hatch; the hatchlings had a total
length of 136-143 mm (SASA 1993).

Geophis bellus MYERS 2003, Amer. Mus.

Novit. 3391: 30; type locality: near community
of Altos de Pacora (east of Cerro Jefe), 700 m
elevation, Panama, Panama. TL to 201 mm.
Known only from type locality. Fig. 619. Geophis brachycephalus (Monteverde,
Geophis brachycephalus (COPE 1871), Proc. Costa Rica). Photo: R. W. Van Devender
Acad. Nat. Sci. Philadelphia 23: 211; type
locality: near San Jose, Costa Rica. TL to 460
mm. Costa Rica to east-central Panama, 250-
2000 m elevation.
Geophis cancellatus SMITH 194lg, Smithsonian
Misc. Coll. 99 (19): 1; type locality:
Chicharras, Chiapas, Mexico. TL to 410 mm.
Southeastern Chiapas, Mexico, and south-
western Guatemala, 500-2000 m elevation in
montane rain forest.
Geophis carinosus STUART 1941c, Occ. Pap.
Mus. Zool. Univ. Michigan 452: 3; type locali-
ty: Finca San Francisco, about 1175 m eleva-
tion, 27 km NE Nebaj, El Quiche, Guatemala.
TL to 276 mm. Veracruz, Mexico, to western
Guatemala, 200-2000 m elevation in wet
forest, pine-oak forest and cloud forest.
Geophis championi BOULENGER 1894, Cat.
Snakes Brit. Mus. 2: 321; type locality: Fig. 620. Geophis brachycephalus (Tapanti,
Costa Rica). Photo: R. W. Van Devender
229
 Chriqui, Panama. TL to 255 mm. Chiriqui
Province, Panama, 1300-1400 m elevation.
Geophis damiani WILSON, MCCRANIE &
WILLIAMS 1998, Proc. Biol. Soc. Washington
111: 410. type locality: 2.5 airline km NNE of
La Fortuna, 1750 m elevation, 15°26'N,
87°18'W, Yoro, Honduras. TL to 327 mm.
Known only from the type locality.
Geophis downsi SAVAGE 1981, Copeia 1981:
549; type locality: Las Cruces Field Station
and Botanical Garden, 4 km S San Vito de
Jaba, 1200 m elevation, Canton Corredores,
Puntarenas, Costa Rica. TL to 248 mm.
Fig. 621. Geophis carinosus (Huehuetenango, Southern part of the Cordillera Costeiia in
Guatemala). Photo: J.A. Campbell southwestern Costa Rica, 1100-1200 m eleva-
tion in montane rain forest.
Geophis dunni SCHMIDT 1932b, Copeia 1932:
8; type locality: Matagalpa, Nicaragua. TL to
367 mm. Known only from the type locality.
Geophis fulvoguttatus MERTENS 1952b, Zool.
Anz. 149: 134; type locality: Hacienda
Montecristo, 2200 m elevation, Cordillera
Metapan, Santa Ana, El Salvador. TL to 398
mm. Northwestern El Salvador and western
Honduras, 1700-2200 m elevation.
Geophis godmani BOULENGER 1894, Cat.
Snakes Brit. Mus. 2: 322; type locality: Irazu,
Costa Rica. TL to 401 mm. Central Costa
Rica to western Panama, 1300-2100 m eleva-
tion.
Geophis hoffmanni (PETERS 1859),
Monatsber. Kong. Akad. Wiss. Berlin 1859:
Fig. 622. Geophis damiani (holotype). 276; type locality: Costa Rica and Puerto
Photo: J. R. McCranie Caballo, Costa Rica; fixed as Costa Rica by
lectotype designation in DOWNS (1967). TL to
300 mm. Eastern Honduras to east-central
Panama, possibly to northern Colombia (see
MYERS 2003), 300-500 m elevation.
Geophis immaculatus DOWNS 1967, Misc.
Puhl. Mus. Zool. Univ. Michigan 131: 90; type
locality: Finca Lorena, ca. 1700 m elevation,
Quetzaltenango, Guatemala. TL to 305 mm.
Chiapas, Mexico, and southwestern
Guatemala, 1500-2500 m elevation in wet
forest and cloud forest.
Geophis laticinctus SMITH & WILLIAMS 1963,
Herpetologica 19: 24; type locality: Colonia
Francisco I. Madero, municipality of Jitotol,
Chiapas, Mexico. TL to 384 mm. Mesa
Central in Chiapas, Mexico, 500-2000 m ele-
vation in wet forest, pine-oak forest and cloud
forest.
Geophis nasalis (COPE 1868), Proc. Acad. Nat.
Sci. Philadelphia 20: 160; type locality: near
Fig. 623. Geophis dunni (holotype). Guatemala City, Guatemala. TL to 350 mm.
Photo: J. H. Townsend
230
Fig. 624. Geophis fulvogutatus (Honduras). Fig. 627. Geophis immaculatus (Quetzal-
Photo: J. R. McCranie tenango, Guatemala). Photo: J. A. Campbell

Fig. 625. Geophis godmani (Jurutungo, Fig. 628. Geophis nasalis (El Rincon,
Chiriqui, Panama). Photo: S. Lotzkat Guatemala). Photo: T. Bille

Fig. 626. Geophis hoffmanni (near Pueblo Fig. 629. Geophis nasalis (Finca Irlanda,
Wiso, Jinotega, Nicaragua). Photo: G. Kohler Chiape.s). Photo: G. Kohler
231
 Colubridae

Pacific side of Guatemala and Chiapas,

Mexico, 400-1830 m elevation in rain forest,
wet forest and dry forest.
Geophis nephodrymus TOWNSEND & WILSON
2006, Proc. Biol. Soc. Washington 119: 151;
type locality: "Sendero Las Minas (l5°29.525'N,
88°12. 705'W), 1570 m elevation, Parque
Nacional El Cusuco, Sierra de Omoa,
Departamento de Cortes, Honduras". TL to
253 mm. Northwestern Honduras, 1545-1780
m elevation in mesic mixed pine-hardwood
forest.
Geophis rhodogaster (COPE 1868), Proc. Acad. Fig. 631. Geophis nephodrymus (Cusuco,
Nat. Sci. Philadelphia 20: 130; type locality:
Honduras). Photo: B. L. Talley
"the elevated country in the neighborhood of
the city of Guatemala". TL to 377 mm.
Eastern Chiapas, Mexico, to northwestern El
Salvador and southwestern Honduras, 1500-
2500 m elevation.
Geophis ruthveni WERNER 1925, Sitz. Akad.
Wiss. Wien 134: 60; type locality: Sarapigui,
Brazil (in error for Sarapiqui, Heredia, Costa
Rica, accoding to DOWNS 1967). TL to 260
mm. Northern and central Costa Rica, 550-
1600 m elevation.
Geophis talamancae LIPS & SAVAGE 1994,
Proc. Biol. Soc. Washington 107: 410; type
locality: Finca Jaguar, 1800 m elevation, Zona
Protectora Las Tablas, Canton Coto Brus,
Puntarenas, Costa Rica. TL to 218 mm.
Cordillera de Talamanca, Costa Rica, near
the border of Panama, 1800 m elevation.
Geophis zeledoni TAYLOR 1954, Univ. Kansas Fig. 632. Geophis nephodrymus (Cusuco,
Sci. Bull. 36: 693; type locality: Finca Zeledon, Honduras). Photo: B. L. Talley
between Volcan Barba and Volcan Poas, 6000
ft, Costa Rica. TL to 397 mm. Vulkan Poas,
Costa Rica, 1600-2000 m elevation.

Fig. 630. Geophis rhodogaster (Guatemala Fig. 633. Geophis nephodrymus (Cusuco,
City, Guatemala). Photo: L. Melendez Honduras). Photo: P. Lewis
232
 Colubridae

Key to Geophis
Geophis brachycephalus
= Geophis godmani
T Geophis downsi
1 a 17 dorsal scale rows .. .......... ...... ............... 2
b 15 dorsal scale rows ............. ... ................ 8
) . .... • Geophis talamancae
2 a Dorsal scales prominently keeled, at least
( .~~· ..
in the posterior halfofbody ...... .............. 3
b Dorsal scales smooth or only slightly

2~ -?:;"J~
keeled only over the cloaca} area .............. 5
3 a Dorsum yellow with brown bands .......... .
........... ........ . ...... . . . . . . . . . .... Geophis dunni

~j ~ b Dorsum unicolor brown or black .............. 4

4 a 41-48 subcaudals; loreal same length or
shorter than pre- and postnasal together;
length of internasal shield approximately
• Geophis ruthveni 3/4 that of prefrontal suture (Fig. 634a);
.&Geophis championi venter banded ........... Geophis carinosus
• Geophis zeledoni b 23-37 subcaudals; loreal longer than pre-
T Geophis bellus
and postnasal together; length of inter-
nasal shield less than 1/2 that of prefrontal
suture (Fig. 634b); venter not banded
...... .... ..... ...................... Geophis nasalis
5 a Dorsum dark with pale brown or orange-
red blotches; blotches not reaching below
2nd dorsal scale row; segmental count
(ventrals and subcaudals combined) 171-
181 ..... ................. Geophis fulvoguttatus
b Dorsum unicolor or, if pale brown or oran-
ge-red spots or bands are present, then
blotches and bands regularly entering 1st
scale row and segmental count smaller
than 161 ................................................... . 6
6 a Without supraocular shields, frontal shield
reaches to eye ... .... Geophis rhodogaster
b Supraocular shields present, frontal shield
does not reach to eye .. ...................... .. ....... 7
• Geophis nasa/is
• Geophis hoffmanni 7 a Chin and lateral edges of ventrals with dark
= Geophis cancellatus
T Geophis dunni
pigmentation ...... Geophis nephodrymus

• Geophis damiani internasal suture

prefrontal suture
Geophis carinosus
:> • Geophis immaculatus
= Geophis laticinctus
111 Geophis rhodogaster
T Geophis fulvoguttatus
......~ Geophis nephodrymus

j a. G. immaculatus

Fig. 634. Dorsal head view.

b. G. hoffmanni

233
 Colubridae
b Chin and ventral scales immaculate
... ... .. .. ................. Geophis immaculatus
8 a Without supraocular shields, parietal
shield reaches to eye ..... .. ......................... 9
b Supraocular shields present, parietal
shield does not reach to eye .... .. ............ 10
9 a Scales on dorsal side of tail and posterior
third of body markedly keeled; ventrals
122-133; subcaudals 41-46; rostral and
prenasals same color as adjacent scales .....
...................... ................ Geophis downsi
b Scales on dorsal side of tail and posterior
third of body smooth; ventrals 132-145;
subcaudals 26-36; rostral and prenasals
white or yellow, in sharp contrast to the
darker coloration of adjacent scales ... ....... .
.... ..... ... ........ .......... .... Geophis godmani
lOa 5 Supralabials; venter predominantly pale
in coloration ............ ...... ......................... 11
b 6 supralabials; venter pale or dark colored
···························································· ···· 12
Ua Dorsal scales smooth or slightly keeled just
over cloacal area ...... Geophis hoffmanni
b Dorsal scales markedly keeled ............ .. ... .
. . . ..... Geophis brachycephalus (in part)
12a Internasals fused with prefrontals; venter
uniformly pale in color ... .. .. ........................
... ... ...................... .... Geophis cancellatus
b Internasals not fused with prefrontals;
venter with dark pigmentation ............. 13
13a Dorsum black with narrow pale bands
throughout length of body; venter black
with a few pale spots ... ........................... 14
b Dorsal and/or ventral coloration different
than above; if pale bands are present,
these are restricted to the posterior portion
ofbody ............ ....... .... ..... ... ............ ....... 15
14a More than 160 ventrals; fewer than 37 sub-
caudals ... ... ... ... .. ..... Geophis laticinctus
b Fewer than 150 ventrals; more than 33
subcaudals .. ...... .. ....... Geophis damiani
15a Dorsal scales markedly keeled at least on
the posterior half of body .... ................. 16
b Dorsal scales smooth or slightly keeled
just over cloacal area .... .... .. ...... ........... 19
16a Loreal longer than pre- and postnasal
together; length of internasal shield less
than 1/2 that of prefrontal suture; frontal
shield one third longer than parietal
suture length ......................... . . . . . . . . . ... . . . 17
b Loreal same length or shorter than pre-
and postnasal together; length of interna-
sal shield corresponds to prefrontal suture
length; frontal shield twice as long as
234
parietal suture length ...... ... ... ..... .......... 18
17a Eye very small, ratio eye length I loreal
length <0.6; head and body uniform glossy
black above and below, with a conspicuous
white band across rear of head ...........
........................................... Geophis bellus
b Eye size moderate, ratio eye length I loreal
length >0.7; dorsum of body uniform black
or with pale bands on posterior part; no
white band across rear of head in adults
(can be present in individuals <150 mm
SVL) Geophis brachycephalus (in part)
18a Snout pointed; rostral markedly produced
posteriorly between internasals; mental
pointed ......... ........... ..... Geophis ruthveni
b Snout rounded; rostral barely produced
posteriory between internasals; mental
rounded ................. Geophis talamancae
19a 125-130 ventrals; 26-33 subcaudals; dorsal
scales smooth over cloacal area .................. .
.. .. .. .. .. .. . . .. .. . . .. ........ Geophis championi
b 141-150 ventrals; 37-46 subcaudals; dorsal
scales slightly keeled over cloacal area .....
...... ... .. ..... .. .. .. .. ........... Geophis zeledoni
Further Reading
DOWNS 1967, SAVAGE 1981, MCCRANIE & WILSON
199la, SASA 1993, LIPS & SAVAGE 1994, E. SMITH
1995, WILSON et al. 1998, MYERS 2003,
MCCRANIE & CASTANEDA 2004b, TOWNSEND &
WILSON 2006, MCCRANIE & CASTANEDA 2007
Hydromorphus
The water colubrids of the genus
Hydromorphus are mid-sized, harmless
snakes, of which one species (H. concolor)
is distributed widely in Central America,
whereas the other (H. dunni), is known
only from one location in western Panama.
In western Honduras, I found a H. conco-
lor in a small stream during a nightly
rain. Otherwise, these colubrids prefer to
remain in the area of piles of detritus in
streams and ponds (SAVAGE & DONNELLY
1988, personal observation). These ani-
mals only leave their aquatic habitat
during nightly rainfalls, as also supported
by the observations of VENCES et al. (1998).
Nothing is known about the diet of H. con-

color, however, they most likely feed on fish
and frogs. This species is oviparous, as
demonstrated by a captive-held female
who laid a clutch of four eggs on the 22nd
/23rd of June 1975, from which two young
hatched on the 11th of December 1975
(SAVAGE & DONNELLY 1988). Further repro-
ductive data have been supplied by
SoL6RZANO et al. (1989); a female laid
seven eggs, which were incubated at 19-24
°C (during the day) and 17-21 °C (at night)
and from which a single young hatched
after 94 days, with a total length of 165
mm.
Hydromorphus concolor PETERS 1859,
Monatsber. Kong. Akad. Wiss. Berlin 1859:
277; type locality: Costa Rica. TL to 709 mm.
Eastern Guatemala to central Panama, 100-
1400 m elevation in rain and wet forest.
Hydromorphus clarki DUNN 1942 is regarded
to be a synonym of H. concolor according to
SAVAGE & DONNELLY (1988).
Hydromorphus dunni SLEVIN 1942, Proc.
Calif. Acad. Sci. 23: 474; type locality: vicinity
north of Boquete, Chiriqui, Panama. TL to
482 mm. Western Panama (in the area of
Boquete, Chiriqui), 1250 m elevation.
Hydromorphus conco/or
• Hydromorphus dunni
Key to Hydromorphus
1 a One prefrontal shield; one internasal
shield; dorsal scales in 17 rows at midbody
... .. ........ .. ..... ... Hydromorphus concolor
b Three prefrontal shields; two internasal
shields; dorsal scales in 15 rows at mid-
body ...................... Hydromorphus dunni
Colubridae
Fig. 635. Hydromorphus concolor (western
Honduras). Photo: G. Kohler
Further Reading
NELSON 1966, VILLA 1970, SAVAGE & DONNELLY
1988, VILLA 1990
Imantodes
The species of the genus Imantodes are
distinguished by a very slender body, a
large head clearly distinct from the neck,
and large, protruding eyes. They are cre-
puscular and nocturnal shrub dwellers,
which are entirely harmless and rarely
attempt to bite. During the day, they hide
in bromeliads, hollow trees, under loose
bark and under rotten logs (HENDERSON &
NICKERSON 1976). At night, they can be
seen, usually stretched out, gliding slowly
through the vegetation, particularly along
streams on the lookout for anoles (Norops),
which make up the bulk of their prey
(MYERS 1982). They will also consume
small frogs and reptile eggs (LANDY et al.
1966). All species of this genus are ovipa-
rous, producing clutches of 1-8 (usually 2-
3) eggs (FITCH 1985, CAMPBELL 1998a). A
female I. cenchoa captured by MYERS
(1982) laid a clutch of 3 eggs on the 30th of
May, which, after 69 days of incubation,
contained embryos that were nearly ready
to hatch. It is otherwise reported that the
egg-laying period for this species in the
northern portion of Central America is
from April until June (STUART 1948).
235
 Colubridae

Imantodes cenchoa (LINNAEUS 1758), Systema

Naturae, ed. 10: 215; type locality: America.
TL to 1106 mm. Southern Tamaulipas and
Oaxaca, Mexico, to Argentina, sea level to
approximately 1500 m elevation in rain
forest, wet forest and dry forest. According to
MYERS (1982) and WILSON & MEYER (1985),
the division into subspecies suggested by H.
SMITH (1942b) and PETERS & OREJAS-MIRANDA
(1970) is insufficiently supported.
Imantodes gemmistratus (COPE 1861e), Proc.
Acad. Nat. Sci. Philadelphia 13: 296; type
locality: near Izalco, San Salvador, El
Salvador. TL to 880 mm. Northern Veracruz
Fig. 636. Imantodes cenchoa (Palo Seco, and southern Sonora, Mexico, to Colombia,
Panama). Photo: G. Kohler sea level to approximately 1500 m elevation
in dry forest and wet forest. According to
MYERS (1982) and WILSON & MEYER (1985)
the division into subspecies suggested by
ZWEIFEL (1959) and PETERS & OREJAS-
MIRANDA (1970) is insufficiently supported, at
least in Central America, and I prefer not to
recognize any subspecies.

Fig. 637. Imantodes cenchoa (La Selva,

Heredia, Costa Rica). Photo: G. Kohler

Fig. 639. Imantodes phantasma (holotype).

Photo: C. W. Myers

lmantodes

:= /. tenuissimus
I. cenchoa
Fig. 638 Imantodes cenchoa (Bartola, Rio San • I. phantasma
Juan, Nicaragua). Photo: G. Kohler
236
 Colubridae

Imantodes inornatus (BOULENGER 1896a),

Cat. Snakes Brit. Mus. 3: 88; type locality:
Hacienda Rosa de Jericho [= Jerico],
Nicaragua, 3250 ft. TL to 1035 mm.
Northwestern Honduras to northwestern
Ecuador, sea level to approximately 1000 m
elevation in rain and wet forest.
Imantodes phantasma MYERS 1982, Amer.
Mus. Novit. 2738: 3; type locality: SE slope of
Cerro Cituro, 1030 m elevation, a peak on
north end of the Serrania de Pirre, Darien,
Panama. TL to 1082 mm. Darien Region of
Panama, 1000-1100 m elevation in rain forest
and cloud forest. Fig. 641. lmantodes gemmistratus (Monteverde,
Imantodes tenuissimus COPE 1866, Proc. Costa Rica). Photo: R. W. Van Devender
Acad. Nat. Sci. Philadelphia 18: 317; type
locality: Yucatan, Mexico. TL to 600 mm.
Northern Yucatan Peninsula, sea level to 100
m elevation in dry forest.

Fig. 642 lmantodes inornatus (Los Guatuzos,

Fig. 640. Imantodes tenuissimus (Puerto Nicaragua). Photo: J. Sunyer
Morelos, Quintana Roo, Mexico).
Photo: R. Cedeno V.

lmantodes gemmistratus
= lmantodes inornatus Fig. 643. lmantodes inornatus (Tortuguero,
Limon, Costa Rica). Photo: G. Kohler
237
 Colubridae

Key to Imantodes
1 a Dorsum with large, dark, saddle-like spots
(Fig. 644a) ..................... ............... .. .. .. .. ... 2
b Dorsum unicolor pale brown or with irreg-
ularly distributed dark spots, indistinct
dark lines or pale saddle-like spots (Fig.
644b) ...... ..... ............ ... .. .... ..... .. ..... ..... 4
a. I. cenchoa
2 a Scales of the medial dorsal rows distinctly
enlarged (3-4 times as wide as the adjacent
scales; Fig. 645a) .. .... Imantodes cenchoa
b Scales of the medial dorsal rows not or only
slightly enlarged (1.5-2.0 times as wide as
the adjacent scales; Fig. 645b) ................ .. 3
3 a Dark dorsal spots not interrupted at sides;
more than 240 ventrals ....... ..................
......... ........ ........ Imantodes tenuissimus b. /. gemmistratus
b Dark dorsal spots often interrupted at
sides, especially posteriorly; usually fewer Fig. 645. Dorsal scales in Imantodes .
than 240 ventrals ..................... .................. .
.. . .. .. .. . . . . ...... ... Imantodes gemmistratus
4 a Fewer than 220 ventrals; fewer than 140
subcaudals; dorsum unicolor pale brown or
with irregularly distributed dark spots
.......... ... ... ..... ....... Imantodes inornatus
b More than 220 ventrals; more than 140
subcaudals; dorsum with pale saddle-
shaped spots ...... Imantodes phantasma Lampropeltis
Kingsnakes, as the species of the genus
Lampropeltis are known, are attractive, for
the most part beautifully colored snakes
that enjoy tremendous popularity as
,...,, ',.·, terrarium pets. For this reason, they are
probably the best known snakes of the
t)f Neotropics. Only a single species occurs in
,,, "'
'Ut
Central America. Lampropeltis triangulum

.... ..,, ,,,

\It is, however, extremely variable in colora-
I tion and markings throughout its range, as
,,,
,...,, '"·, reflected by the proliferation of subspecies
that have been described.
t)f ., ,,
,,. "' ,,, In Central America, the kingsnake is pre-
dominantly a nocturnal forest-dweller.
Along with J .R. MCCRANIE and K.
.... ". NICHOLSON in the Patuca National Park, I
a. I. gemmistratus b. /. inornatus came across a subadult specimen that was
Fig. 644. Dorsal pattern in Imantodes. active during the day. These strong colu-
brids feed on lizards, frogs, snakes, birds,
and rodents. Lampropeltis triangulum
reproduces oviparously, producing clutches
Further Reading of 4-24 eggs, which hatch after about two
H. SMITH 1942b, J. PETERS 1954, ZWEIFEL 1959, months (48-65 days depending on incuba-
MYERS 1982, SAVAGE & SCOTT 1985 tion temperature) (KOHLER 1997).
238
 Colubridae

Fig. 646. Lampropeltis triangulum hondurensis (Rio Patuca, Olancho, Honduras).Photo: G. Kohler

Lampropeltis triangulum (LACEPEDE 1789),

Hist. Nat. Quadr. Ovip. Serp. 2: 86; type local-
ity: America. TL to 2000 mm. Distributed all
across the central and eastern USA as well as
across Mexico, Central America and north-
western South America. In Central America,
the subspecies Lampropeltis t. abnorma (Bo-
COURT 1886) (central and northern Guate-
mala, southern Belize), Lampropeltis t. blan-
chardi STUART 1935b (Yucatan Peninsula),
Lampropeltis t. gaigeae DUNN 1937b (south-
eastern Costa Rica and western Panama),
Lampropeltis t. hondurensis WILLIAMS 1978
(Caribbean side of Honduras, Nicaragua and
northeastern Costa Rica), Lampropeltis t.
micropholis COPE 1861d (central Panama to
northwestern South America), Lampropeltis
t. oligozona BOCOURT 1886 (Pacific side of
Chiapas, Mexico, to southeastern Guate-
mala), Lampropeltis t. polyzona COPE 1860d
(Veracruz, Mexico, to western Guatemala)
and Lampropeltis t. stuarti WILLIAMS 1978
(Pacific side of southeastern Guatemala to
central Costa Rica) occur. Fig. 64 7. Lampropeltis triangulum (near La
Ceiba, Atlantida, Honduras). Photo: G. Kohler
239
 Colubridae

Fig. 648. Lampropeltis triangulum gaigeae Fig. 650. Leptodeira frenata (Pichucalco,
(Jurutungo, Chiriqui, Panama). Photo: G. Kohler Chiapas, Mexico). Photo: A. Ramirez V.

Fig. 649. Lampropeltis triangulum micropholis. Fig. 651. Leptodeira nigrofasciata (Liberia,
Photo: U. Kuch Guanacaste, Costa Rica). Photo: R. D. Bartlett

• Lampropeltis triangulum

Fig. 652. Juvenile Leptodeira septentrionalis

(Golfito, Costa Rica). Photo: D. M. Dehling
Further Reading
BLANEY 1973, K. WILLIAMS 1978, 1988, 1994

240
Leptodeira
The species of the genus Leptodeira
have the common name cat-eyed
snakes, due to their vertical pupils.
They are nocturnal colubrids that
are harmless as a rule, and rarely
attempt to bite. They are found in a broad
range of habitats, even in secondary vege-
tation close to human habitation. At night,
these semi-arboreal snakes are on the look-
out for their preferred prey of frogs, small
snakes, and lizards. Leptodeira annulata is
known in particular for consuming frog
spawn that has been deposited on leaves
(especially that of red-eyed tree-frogs
[Agalychnis callidryas] and glass frogs
[family Centrolenidae]) (CAMPBELL 1998a).
All Leptodeira species reproduce ovipar-
ously, with clutch sizes between 2 and 12
eggs (DUELLMAN 1958, FITCH 1985). The
incubation period, depending on tempera-
ture, is 56-85 days (KOHLER 1997,
CAMPBELL 1998a).
Leptodeira annulata
=Leptodeira frenata
111 Leptodeira rubricata
= Leptodeira nigrofasciata
Leptodeira septentrionalis
Fig. 653. Leptodeira
frenata (Isla Cozumel,
Quintana Roo, Mexico).
Photo: J.C. Lee
Leptodeira annulata (LINNAEUS 1758),
Systema Naturae, ed. 10: 215; type locality:
Amazon Basin. TL to 870 mm. Tamaulipas
and Guerrero, Mexico, to Brazil and
Argentina (excluding the Yucatan Peninsula),
sea level to approximately 1000 m elevation
in rain forest, wet forest and dry forest. In
Central America, the subspecies Leptodeira
a. cussiliris DUELLMAN 1958 (Tamaulipas and
Guerrero to Chiapas, Mexico) and Leptodeira
a. rhombifera GUNTHER 1872 (Guatemala to
Panama) occur.
Leptodeira frenata (COPE 1886b), Proc. U.S.
Natl. Mus. 9: 184; Jalapa [= Xalapa],
Veracruz, Mexico. TL to 715 mm. Veracruz,
Mexico, to northern Guatemala, as well as
the entire Yucatan Peninsula, sea level to
1000 m elevation in rain forest, wet forest
and dry forest. In Central America, the sub-
species Leptodeira f malleisi DUNN & STUART
1935 (southern half of the Yucatan Peninsula,
as well as along the east coast of the Yucatan
Peninsula) and Leptodeira f yucatanensis
COPE 1887 (the northern portion of the
Yucatan Peninsula) occur.
Fig. 654. Leptodeira rubricata (Province Punta-
renas, Costa Rica). Photo: R. W. Van Devender
241
 Colubridae
Leptodeira nigrofasciata GUNTHER 1868, Ann.
Mag. Nat. Hist. (4) 1: 425; type locality:
Nicaragua. TL to 550 mm. Guerrero, Mexico,
along the Pacific coast to Costa Rica; in
Honduras also on the Caribbean side, sea
level to approximately 1300 m elevation in
dry forest and wet forest.
Leptodeira rubricata (COPE 1893), Proc. Amer.
Philos. Soc. 31: 347; type locality: Boca Mala,
Costa Rica. TL to 700 mm. Costa Rica, in
mangroves.
Leptodeira septentrionalis (KENNICOTT 1859),
in BAIRD: Reptiles of the Boundary 2: 16; type
locality: Matamoras, Tamaulipas, Mexico. TL
to 1055 mm. Texas, USA, and Sinaloa,
Mexico, to Peru, sea level to approximately
1500 m elevation in rain forest, wet forest
and dry forest. In Central America, the sub-
species Leptodeira s. ornata (BOCOURT 1884)
(Panama and northwestern South America)
and Leptodeira s. polysticta GUNTHER 1895
(Veracruz and Guerrero, Mexico, to Costa
Rica) occur.
a. Leptodeira annulata
b. Leptodeira septentrionalis c. L. frenata
Fig. 655. Dorsal head pattern in three species of
Leptodeira (after DUELLMAN 1958).
242
Fig. 656. Leptodeira annulata (Bartola, Rio
San Juan, Nicaragua). Photo: G. Kohler
Key to Leptodeira
1 a 19 dorsal rows at midbody ........................ 2
b 21 or more dorsal rows at midbody .......... 3
2 a Dorsum with 10-21 dark bands that extend
to the ventrals; 54-76 subcaudals
.. ........ ........... Leptodeira nigrofasciata
b Dorsum with 30-70 dark bands that do not
extend to the ventrals; 72-107 subcaudals
..... Leptodeira septentrionalis (in part)
3 a One dark postorbital stripe present, in
broad contact with first body spot; a single
dark spot in the center of each parietal
(Fig. 655c) ................. Leptodeira frenata
b Dark postorbital stripe, if present, not in
broad contact with first body spot; mark-
ings on the parietals different than above
.......... ................... ... ..... ... .... ..... .. ......... ... 4
4 a Ventrals usually more than 185; 72-107
subcaudals; medial neck stripe, if present,
extends only 3-6 scales behind the
parietals and has no contact with first
body spot (Fig. 655b); dark dorsal spot
without pale border (Fig. 657a) ............. .... .
.... . Leptodeira septentrionalis (in part)
b Ventrals fewer than 185; 54-89 subcaudals;
medial neck stripe usually extends from
the parietals to first body spot, with which
it is confluent (Fig. 655a); dark dorsal spot
with pale border (Fig. 657b) ... .. ....... ...... 5
5 a Dorsum with 23-51 dark spots (usually
with pale borders; Fig. 657b) on pale brown
ground color or spots blend into a zigzag
pattern; venter cream colored in life ...........
.............. .. .... ......... Leptodeira annulata
b Dorsum with q7-67 dark, regular cross
bands (without pale borders) on red
ground color; venter red in life .................. .
.. .... ........ .. .. ......... Leptodeira rubricata
 Colubridae

. •'•.
,• '·.
Leptodrymus

As the species name pulcherrimus implies,

4'. -·
the only species of the genus Leptodrymus is
a particularly graceful, beautifully striped
snake, remarkable for the vivid green

•'
coloration of the dorsal surface of its head.
L. pulcherrimus is a fast moving colubrid,
at home in dry areas. It is a nocturnal
a. Leptodeira septentrionalis ground dweller, whose main diet consists of
lizards.

Leptodrymus pulcherrimus (COPE 1874),

Proc. Acad. Nat. Sci. Philadelphia 27: 65; type
locality: Western slope of Central America. TL
to 975 mm. Along the Pacific side of
Guatemala to Costa Rica; in Guatemala and
Honduras also on the Caribbean side, sea
level to approximately 1300 m elevation in
savanna and dry forest.

b. Leptodeira annulata Further Reading

BOGERT 1939, 1947
Fig. 657. Dorsal body pattern in Leptodeira
(after DUELLMAN 1958).

Further Reading
DUELLMAN 1958

Leptodrymus pulcherrimus

Fig. 658. Leptodeira septentrionalis (Finca

Irlanda, Chiapas, Mexico). Photo: G. Kohler

Fig. 659. Leptodrymus pulcherrimus.

Photo: R. W. Van Devender
243
 Colubridae

Leptophis

The genus Leptophis (parrot snakes) in-

cludes eight species that populate the
forests, mangroves, and savannas of Mexico,
Central, and South America. Leptophis
depressirostris is a magnificent leaf green
snake, as are some of the subspecies of L.
ahaetulla. As their large, round-pupilled
eyes suggest, the Leptophis species are pre-
dominantly diurnal. These agile colubrids
are tree-dwellers or, at least, semi-arboreal
and have an irritable temperament. When Fig. 661. Leptophis nebulosus (Monteverde,
handled, they will respond with a threat- Costa Rica). Photo: M. Franzen
ening gape, substantially expanding the
lower jaw, hence their Latin name. They
will often remain in this pose for an
extended period of time, but rarely make
use of their numerous pointed teeth.

Parrot snakes consume primarily frogs, as

well as the occasional salamander, lizard,
snake or bird egg (OLIVER 1948, STUART
1948, SEXTON & HEATWOLE 1965). All
Leptophis species are oviparous and pro-
duce clutches of 2-11 eggs (STUART 1948,
SEXTON & HEATWOLE 1965, CENSKY &
McCOY 1988, CAMPBELL 1998a). DUELLMAN
(1978) discovered a combined clutch of
Leptodeira and Leptophis ahaetulla eggs in
a bamboo internode about 12 meters off Fig. 662. Leptophis nebulosus (Krahkra,
Gracias a Dios, Honduras).
the ground. In northern Central America, Photo: J. R. McCranie
the egg-laying period occurs from March to
October (CAMPBELL 1998a).

Fig. 660. Leptophis ahaetulla (Selva Negra, Fig. 663. Leptophis ahaetulla (Rio Patuca,
Matagalpa, Nicaragua). Photo: G. Kohler Olancho, Honduras). Photo: G. Kohler

244
 Colubridae

Leptophis ahaetulla (LINNAEUS 1758),

Systema Naturae, ed. 10: 225; type locality:
"Asia, America". TL to 2250 mm. Veracruz
and Oaxaca, Mexico, to northwestern
Ecuador west of the Andes, as well as
Argentina and Brazil east of the Andes, sea
level to approximately 1300 m elevation in
rain forest, wet forest and dry forest. In
Central America, the subspecies Leptophis a.
occidentalis (GONTHER 1859) (Nicaragua to
Panama as well as western Venezuela to
Ecuador) and Leptophis a. praestans (COPE
1869) (Veracruz, Mexico, to Honduras) occur.
Fig. 664. Leptophis depressirostris (near Salto
Labu, Bosawas, Nicaragua). Photo: G. Kohler

Leptophis depressirostris (COPE 1861b), Proc.

Acad. Nat. Sci. Philadelphia 12: 557; type
locality: Cocuyas de Veraguas, New Grenada
[= Cocuyas, Panama). TL to 1400 mm.
Nicaragua to western Ecuador.
Leptophis diplotropis (GONTHER 1872), Ann.
Mag. Nat. Hist. (4) 9: 25; type locality:
Tehuantepec, Mexico. TL to 1410 mm.
Southwestern Chihuahua and southern
Sonora to southeastern Oaxaca, Mexico, sea
.a. Leptophis riveti level to approximately 1000 m elevation in
• Leptophis depressirostris dry forest and wet forest.
Leptophis mexicanus DUMERIL, BIBRON &
DuMERIL 1854, Erp. Gen., 7: 536; type locality:
Mexico. TL to 1270 mm. Tamaulipas and
Oaxaca, Mexico, to Costa Rica, sea level to
approximately 1700 m elevation in dry forest
and wet forest. In Central America, as well as
the nominate subspecies, the subspecies
Leptophis m . hoeversi HENDERSON 1976
(endemic to the Turneffe Islands, Belize)
occur.
Leptophis modestus (GONTHER 1872), Ann.
Mag. Nat. Hist. (4) 9: 26; type locality: Rio
Chisoy [= Chixoy or Negro], below the town of
::::: Leptophis nebulosus Cubulco, Baja Verapaz, Guatemala [erron-
Leptophis ahaetulla
eous according to MCCRANIE & WILSON 1993).
TL to 1720 mm. Northeastern Chiapas and
central Guatemala to northwestern El
Salvador and southwestern Honduras, 1500-
2000 in cloud forest.
Leptophis nebulosus OLIVER 1942, Occ. Pap.
Mus. Zool. Univ. Michigan 462: 12; type lo-
cality: Cariblanca, Costa Rica. TL to 854 mm.
Northeastern Honduras to Costa Rica.
Leptophis riveti DESPAX 1910, Bull. Mus. Hist.
Nat. Paris 7: 368; type locality: Gualaquiza,
730 m elevation, Ecuador. TL to 900 mm.
Southeastern Costa Rica to Ecuador west of
1111 Leptophis modestus
the Andes, as well as Peru east of the Andes,
• Leptophis mexicanus
.& Leptophis diplotropis
at elevations of 1000-1800 m .

245
 Colubridae

Key to Leptophis
1 a Loreal present ....... ...... .. .... .. ............. ..... 2
b No loreal .............. ...... ... .. .... ................. .. 5
2 a Keels present only on the paravertebral
scale rows ....... ... ..... ... ... .......................... 3
b Keels on all the dorsal scale rows, except
for the first row ............. ... .. ... .. .. .... .. ....... 4
3 a Fewer than 160 ventrals; 9 supralabials
........ .............. Leptophis depressirostris
b More than 160 ventrals; 8 supralabials ......
...... ... ....... .. ...... ..... Leptophis diplotropis
4 a Unicolor green; ventrals without lateral
keel; 108-183 ventrals .. ...... ............. ....... .
. .......... ...... Leptophis modestus (in part)
b Mid-dorsum bronze colored, bordered by a
dark longitudinal stripe; ventrals always Fig. 665. Leptophis ahaetulla (Selva Negra,
with a weakly defined lateral keel; 145-174 Matagalpa, Nicaragua). Photo: G. Kohler
ventrals ................ Leptophis mexicanus
5 a Fewer than 150 ventrals; all dorsal scales
keeled ... ... ... ... .. .. ..... .. ... Leptophis rive ti
b More than 150 ventrals; scales of the first
dorsal row without keels ... .... .. ... ... ......... 6
6 a A blue-green or blue longitudinal stripe
running along the second and third or
third and fourth dorsal rows .......................
.. . . . . . . . . . . . . . . . . . . ....... .. Leptophis nebulosus
b Unicolor green or, if a blue-green or blue
longitudinal stripe is present, it runs along
the third to fifth dorsal rows .................... 7
7 a Unicolor green; 168-183 ventrals; in the
highlands (1500-2000 m elevation) .............
. . . . . . . ... ...... Leptophis modestus (in part)
b Unicolor green or with a blue-green or blue
longitudinal stripe along the third to fifth
dorsal rows; 150-183 ventrals; in the low-
land and premontane areas (sea level to Fig. 666. Leptophis mexicanus (Isla de Utila,
about 1300 m elevation) ............................. . Honduras). Photo: G. Kohler
................ ............... Leptophis ahaetulla

Further Reading
OLIVER 1948, MERTENS 1971, 1973, MCCRANIE &
WILSON 1993

Fig. 667. Leptophis diplotropis (Canon de

Sumidero, Chiapas, Mexico).
Photo: A. Ramirez V.
246
 Colubridae

Liophis
The genus Liophis includes around 35 spe-
cies, only two of which (L. epinephelus and
L. lineatus) reach Central America (DIXON
1989). In the past, Liophis epinephelus was
placed in the genus Leimadophis (J.
PETERS & 0REJAS-MIRANDA 1970, SAVAGE &
VILLA 1986), whereas L. lineatus was
included in the genus Lygophis (J. PETERS
& 0REJAS-MIRANDA 1970). Both these spe-
cies are ground-dwellers that feed on Fig. 668. Liophis lineatus (Cordoba, Colombia).
lizards and frogs, and also on invertebrates Photo: M. Lundberg
as hatchlings (TAYLOR 1951, LANCINI &
KoRNACKER 1989). They reproduce ovipar-
ously, producing clutches of 1-8 eggs.

Liophis epinephelus COPE 1862a, Proc. Acad.

Nat. Sci. Philadelphia 14: 78; type locality:
Truando, Colombia. TL to 775 mm. Costa
Rica and Panama, as well as northwestern
South America, sea level to 2200 m elevation
in multiple habitats. In Central America, the
subspecies Liophis e. epinephelus (Panama
and northwestern South America) and
Liophis e. juuenalis DUNN 1937a (Costa Rica)
occur.
Liophis lineatus (LINNAEUS 1758), Systema
Naturae, ed. 10: 221; type locality: Asia [in Fig. 669. Liophis epinephelus (Costa Rica).
error] . TL to 737 mm. Panama and northern Photo: R. W. Van Devender
South America, in savanna and dry forest.

Key to Liophis
1 a 17 dorsal rows at midbody; fewer than 155
ventrals ................ Liophis epinephelus
b 19 dorsal rows at midbody; more than 155
ventrals ............ ........... Liophis lineatus

Liophis epinephelus
= Liophis lineatus

Fig. 670. Liophis epinephelus (Reserva

Foresta! Fortuna, Panama). Photo: G. Kohler

Further Reading
DIXON 1980, 1983, MICHAUD & DIXON 1987,
DIXON 1989.
247
 Colubridae
Manolepis
The genus Manolepis is monotypic, contain-
ing the species M. putnami. Interestingly,
this species displays a pronounced sexual
dichromatism: while both genders have the
dark dorsal stripe typical of the species, in
the male it is uniformly dark, and in the
female, it has a pale center (WERLER &
SMITH 1952, JOHNSON 1978). Little is
known about the habits of this colubrid.
They live in dry areas and reproduce by
laying eggs (lIARTWEG & OLIVER 1940).
Manolepis putnami (JAN 1863), Elenco
sistematico degli Ofidi: 67; type locality: San
Blas, Nayarit, Mexico. TL to 717 mm.
Nayarit, Mexico, along Pacific coast to
eastern Chiapas, 300-1000 m elevation in dry
forest and pine-oak forest.
Further Reading
WERLER & SMITH 1952, JOHNSON 1978
Fig. 671. Manolepis putnami (Monte Bonito,
Chiapas, Mexico). Photo: A. Ramirez V.
248
Masticophis
Masticophis, which is closely related to the
genus Coluber, includes eight mid-size to
large, very agile colubrid species that are
distributed all across the southern half of
the United States, as well as across
Mexico, and one species that is distributed
in Central America. Isolated populations of
this species also exist in Colombia and
Venezuela. Masticophis mentouarius is a
diurnal ground-dwelling colubrid that will
dart off at great speed if disturbed. If
handled, they will gape with wide-open
mouths and not hesitate to make warning
strikes and bite. Their diet consists mainly
of lizards (Aspidoscelis and Sceloporus),
but they will also hunt rodents and birds
(HENDERSON & HOEVERS 1977b, ALVAREZ
DEL TORO 1983, CAMPBELL 1998a).
Egglaying has so far been documented in
March, with clutch sizes between 7 and 30
eggs (WERLER 1951, MINTON & MINTON
1991, CAMPBELL 1998a).
Masticophis mentovarius (DUMERIL, BIBRON &
DuME:RrL 1854), Erp. Gen. 7: 187; type locali-
ty: Mexico. TL to 2500 mm. Sonora and San
Luis Potosi, Mexico, to Costa Rica; also dis-
junct in Panama, Colombia and Venezuela,
sea level to approximately 2000 m elevation
in savanna, pine forest and dry forest. In
Central America, the subspecies Masticophis
m . mentovarius (Jalisco and San Luis Potosi,
Mexico, to Costa Rica) and Masticophis m.
centralis (ROZE 1953) (disjunct in Panama,
Colombia and Venezuela) occur.
1111 Manolepis putnami
Masticophis mentovarius
 Colubridae

Mastigodryas
The nomenclatural history of the genera
Dryadophis and Mastigodryas is some-
what complicated (AMARAL 1964, H. SMITH
& LARSEN, DIXON & TIPTON 2004). I follow
DIXON & TIPTON (2004) in relegating
Mastigodryas and Dryadophis to the
single genus, Mastigodryas.

While adult specimens of the three

Central American species of the genus
Mastigodryas are all one color or have lon-
Fig. 672. Masticophis mentouarius (Valle de gitudinal stripes, the juveniles display
Otoro, Honduras). Photo: G. Kohler
pale cross bands (see Figs. 676-677). All
Mastigodryas species are agile and very
irritable. These diurnal colubrids live in
savannas, dry forest and rain forest, where
they hunt predominantly lizards (in parti-
cular, Basiliscus, Norops, Sceloporus, and
Aspidoscelis), but also frogs, snakes, and
rodents (SEXTON & HEATWOLE 1965,
HIMMELSTEIN 1980, SEIB 1984). All
Mastigodryas species lay eggs (2-5 per
clutch), with larger females producing lar-
ger clutches. Female M. melanolomus
reach sexual maturity at approximately
580 mm SVL, at the age of at least 18
months (CENSKY & McCoy 1988).

Fig. 673. Juvenile Masticophis mentouarius

(Moss, Atlantico Norte, Nicaragua).
Photo: J. Sunyer

Fig. 674. Masticophis mentouarius (Palo Verde,

Costa Rica). Photo: D. M. Dehling

Further Reading
0RTENBURGER 1928, WILSON 1973, JOHNSON Fig. 675. Mastigodryas melanolomus (Isla
1977' 1982, NAGY et al. 2004 Utila, Honduras). Photo: G. Kohler
249
 Colubridae

Mastigodryas dorsalis (BOCOURT 1890), Misc.

Sci. Mex., Rept. 1890: 724; type locality:
Guatemala. TL to 1140 mm. Guatemala to
central Nicaragua, 750-1900 m elevation in
montane rain forest and cloud forest.
Mastigodryas melanolomus (COPE 1868), J.
Acad. Nat. Sci. Philadelphia 20: 134; type
locality: Yucatan, Mexico. TL to 1360 mm.
Tamaulipas and Sinaloa, Mexico, to central
Panama, sea level to 1500 m elevation in dry
forest, wet forest, and rain forest. In Central
America, the subspecies Mastigodryas m .
melanolomus (Yucatan Peninsula), Mastigo-
dryas m. alternatus (BocoURT 1884)
(Honduras to Panama), Mastigodryas m. lae-
vis (FISCHER 1881) (Alta Verapaz, Guatemala, Fig. 678. Mastigodryas melanolomus (Volcan
and western Honduras), and Mastigodryas Mombacho, Granada, Nicaragua).
m. tehuanae (SMITH 1943) (Nayarit, Mexico, Photo: G. Kohler
along the Pacific side to western Guatemala)
occur.
Mastigodryas pleei (DUMERIL, BIBRON &
DuMERIL 1854), Erp. Gen. 7: 661; type locali-
ty: Venezuela. TL to 1400 mm. Eastern
Panama and northern South America
(Colombia and Venezuela).

Fig. 679. Mastigodryas melanolomus (Hitoy

Cerere, Costa Rica). Photo: D. M. Dehling

Fig. 676. Juvenile Mastigodryas melanolomus

(western Honduras). Photo: G. Kohler

Fig. 677. Juvenile Mastigodryas dorsalis Fig. 680. Mastigodryas pleei (Cordoba,
(Datanli, Nicaragua). Photo: J. Sunyer Colombia). Photo: M. Lundberg
250

Fig. 681. Mastigodryas dorsalis (Selva Negra,
Matagalpa, Nicaragua). Photo: G. Kohler
Key to Mastigodryas
1 a No vertebral longitudinal stripe ............... .
.... .. .. .......... Mastigodryas melanolomus
b A dark medial longitudinal stripe present
at least on the anterior end of the body
(additional longitudinal stripes may be
present posteriorly) ..................... .. ......... 2
2 a Only one dark medial longitudinal stripe
is formed; dark lateral stripes present or
not ....... .. .. ......... Mastigodryas dorsalis
b Three dark longitudinal stripes that run
close to one another anteriorly and join
into a single stripe posteriorly; in addition,
a dark lateral stripe present on each side .. .
..... .. .. .. . .. .. .. .. .. .. ....... Mastigodryas pleei
Further Reading
STUART 194la, LYNCH & H. SMITH 1966, SMITH &
LARSEN 197 4b, K6HLER 2005
:= Mastigodryas dorsa/is
• Mastigodryas pleei
Mastigodryas melanolomus
Colubridae
Nerodia
Only one species of the genus Nerodia , N
rhombifera, reaches as far as the northern
portion of Central America, whereas the
other nine species are distributed across
North America. As with all species of this
genus, it is semiaquatic, living in rivers,
lakes, ponds, and flooded meadows. Its diet
consists primarily of fish, but includes
frogs and freshwater shrimp as well
(WRIGHT & WRIGHT 1957, MANJARREZ &
MAciAs-GARCfA 1991). Nerodia rhombifera
is predominantly nocturnal, but is
occasionally seen basking during the day
on branches jutting out of the water. They
are very irritable snakes that will not
hesitate to bite when caught. This species
is a livebearer, with the tropical
populations bearing fewer babies than
their northern conspecifics (ALDRIDGE et
al. 1995). According to these authors, the
litter size of N rhombifera in Veracruz,
Mexico, is 8-35 (mean 17). Females reach
sexual maturity with a snout-vent length
of670 mm, males at 475 mm (ALDRIDGE et
al. 1995).
Fig. 682. Nerodia rhombifera (Veracruz,
Mexico). Photo: P. Heimes
251
 Colubridae
Nerodia rhombifera (HALLOWELL 1852), Proc.
Acad. Nat. Sci. Philadelphia 6: 177; type lo-
cality: the Arkansas river and its tributaries
near the northern boundary of the Creek
Nation. TL to 1200 mm. Iowa, Illinois and
Indiana, USA, to Tabasco and northern
Chiapas, Mexico, sea level to 500 m elevation.
The subspecies Nerodia r. werleri (CONANT
1953) occurs in Central America.
• Nerodia rhombifera
Further Reading
CONANT 1969, McALLISTER 1985,ALDRIDGE et al.
1995
Fig. 683. Ninia sebae (El lmposible National Park, Ahuachapan, El Salvador).
252
Ninia
The genus Ninia includes small, secretive,
harmless snakes that, in many places,
make up an important part of the leaf-lit-
ter herpetofauna. Very often, these snakes
are found under logs and in rotten roots on
coffee plantations, which is why they are
known as "coffee snakes". All Ninia species
are predominantly nocturnal and eat
earthworms, leeches, slugs, and snails
(TAYLOR 1951, GREENE 1975). If threatened,
the Ninia will flatten their bodies, hide
their heads under their body coils and lift
their tails in a spiral (GREENE 1975).
Occasionally, they will also lift the front of
their bodies "cobra style" in order to dis-
play to potential predator (HENDERSON &
HOEVERS 1977a, compare Figs. 687 and
690). All Ninia species are oviparous and
produce clutches of 1-4 eggs between July
and December (GAIGE 1936, GREENE 1975,
ALVAREZ DEL TORO 1983). The juveniles of
N. sebae hatch after an incubation period of
75-79 days (GREENE 1975).
Photo: G. Kohler
 Colubridae

Ninia atrata (HALLOWELL 1852), Proc. Acad.

Nat. Sci. Philadelphia 6: 125; type locality:
within 200 miles of Caracas, Venezuela. TL to
450 mm. Eastern Panama and northern
South America (Ecuador, Colombia and
Venezuela), 830-1360 m elevation in rain
forest and cloud forest.
Ninia celata MCCRANIE & WILSON 1995, J.
Herpetol. 29 (2): 225; type locality: Cinchona
(Isla Bonita), Atlantic side of Volcan Poas,
1200 m elevation, Alajuela, Costa Rica. TL to
360 mm. North-central Costa Rica and Fig. 684. Ninia espinali (Cerro El Pital, 2270 m,
western Panama, 830-1600 m elevation in Chalatenango, El Salvador). Photo: 0. Komar
rain forest and cloud forest.
Ninia diademata BAIRD & GIRARD 1853, Cat.
N. Amer. Rept.: 49; type locality: Orizaba,
Mexico. TL to 421 mm. San Luis Potosi and
Oaxaca, Mexico, to central Honduras, sea
level to 2200 m elevation in rain forest and
cloud forest. According to WILSON & MEYER
(1985), the division into subspecies suggested
by BURGER & WERLER (1954) is insufficiently
supported, and I prefer not to recognize any
subspecies.
Ninia espinali MCCRANIE & WILSON 1995, J.
Herpetol. 29 (2): 228; type locality: El Portillo
de Ocotepeque (14°28'N, 89°04'W), 1910 m
• Ninia maculata
elevation, Ocotepeque, Honduras. TL to 510 1111 Ninia pavimentata
mm. Western Honduras and northern El
Salvador, 1590-2270 m elevation in cloud
forest.
Ninia maculata (PETERS 1861), Monatsber.
Kong. Akad. Wiss. Berlin 1861: 924; type lo-
cality: Costa Rica. TL to 352 mm. Eastern
Honduras to eastern Panama, sea level to
1750 m elevation in rain forest and cloud
forest.
Ninia pavimentata (BocoURT 1883), Miss.
Sci. Mex., Rept.: 549; type locality: Alta Vera-
paz, Guatemala. TL to 380 mm. Fringe of high- • Ninia atrata
lands of Guatemala, 1300-1500 m elevation. • Ninia espinali
.a. Ninia celata
Ninia psephota (COPE 1876), J. Acad. Nat. Sci. := Ninia psephota
Philadelphia (2) 8: 146; type locality: higher
points on Pico Blanco, 5000-7000 ft, Costa
Rica. TL to 494 mm. Costa Rica and western
Panama, 1200-2770 m elevation in rain forest
and cloud forest.
Ninia sebae (DUMERIL, BIBRON & DUMERIL
1854), Erp. Gen., 7: 515; type locality: Mexico.
TL to 386 mm. Veracruz and Oaxaca, Mexico,
to central Costa Rica, sea level to 2200 m
elevation in wet forest, rain forest, and cloud
forest. According to WILSON & MEYER (1985),
the division into subspecies suggested by
SCHMIDT & RAND (1957) is insufficiently
supported, and I prefer not to recognize any Ninia sebae
subspecies. = Ninia diademata
253
 Colubridae

Fig. 685. Ninia diademata (near Quebrada Fig. 688. Ninia maculata (Selva Negra,
Grande, Copan, Honduras). Photo: J. R. McCranie Matagalpa, Nicaragua). Photo: G. Kohler

Fig. 686. Ninia psephota (Guanacaste, Costa Fig. 689. Ninia psephota (Monteverde, Costa
Rica). Photo: R. W. Van Devender Rica). Photo: R. W. Van Devender

Fig. 687. Display behavior in Ninia hudsoni Fig. 690. Display behavior in Ninia pavimentata
(Ecuador). Photo: G. Kohler (from DUMERIL et al. 1870-1909).
254
 Colubridae

Fig. 691. Ninia sebae (near Guanagasapa,
Escuintla, Guatemala).
Fig. 692. Ninia atrata (Cordoba, Colombia).
Photo: M. Lundberg
Key toNinia
1 a 1 7 dorsal rows at mid body ..... ..... ..... ..... .. .
. . . . . . . . . . . ...... ... ... ... .. ....... .. Ninia psephota
b 19 dorsal rows at midbody ............. ........... 2
2 a Venter with dark, often mosaic-patterned
spots ......................... ....... .. ................... .... 3
b Venter largely unpigmented ......... ........ .. 5
3 a Fewer than 62 subcaudals
........... ....... .... ... ....... ....... Ninia maculata
b More than 62 subcaudals ................ ......... 4
4 a Dorsum reddish-brown with narrow, dark
cross bands .... ......... Ninia pavimentata
b Dorsum dark brown or black, but without
Photo: G. Kohler dark cross bands .. ....... Ninia diademata
5 a Dorsum red or reddish brown in life (pale
to dark brown in preservative); pale neck
band (yellow to pale brown in life) with
posterior black border present, that does not
cover parietals (Fig. 693a) ..... Ninia sebae
b Dorsum black or dark; if a pale neck band
is formed, it also covers the posterior half
of the parietals (Fig. 693b) ........ ................ 6
6 a Without pale neck band ... Ninia espinali
b Pale neck band distinct ............................. 7
7 a More than 130 ventrals; 39-70 subcaudals
....... ....................................... Ninia atrata
b Fewer than 130 ventrals; 33-45 subcaudals
................ ...... ..... .... ...... .. .. .. Ninia celata
Further Reading
BURGER & WERLER 1954, K. SCHMIDT & RAND
1957, SAVAGE & LAHANAS 1991, MCCRANIE &
WILSON 1995, E . SMITH & CAMPBELL 1996,
MCCRANIE et al. 2001

a. N. sebae b. N. atrata c. N. diademata

Fig. 693. Dorsal head pattern in three species of N inia.
255
 Colubridae
Nothopsis
The genus Nothopsis is monotypic contain-
ing the species N. rugosus. Both the shape
of the head and the presence of small irreg-
ular scales on the dorsal surface of the
head give this colubrid a rather boid-like
appearance. Even today, our knowledge of
the biology of N. rugosus is very sketchy
and the species is scarcely represented in
museum collections. F. SCHMIDT and I dis-
covered a specimen of N. rugosus lying
motionless on the ground at 820 m eleva-
tion at Cerro Saslaya (northeastern
Nicaragua) during a rain shower at night
(KOHLER & SCHMIDT 2001). When we picked
it up, it made no attempt to bite. The few
other existing notes on the habits of this
species indicate that it is a nocturnal ground-
dweller (DUNN & DOWLING 1957).
Nothopsis rugosus COPE 1871, Proc. Acad.
Nat. Sci. Philadelphia 23: 201; type locality:
Darien, Panama. TL to 433 mm. Eastern
Honduras to Colombia, 250-900 m elevation
in rain forest.
Further Reading
DUNN & DOWLING 1957, O'SHEA 1986, KOHLER &
SCHMIDT 2001
Fig. 694. Nothopsis rugosus (Cerro Saslaya,
Atlantico Norte, Nicaragua). Photo: G. Kohler
256
Omoadiphas
The genus Omoadiphas (KOHLER et al.
2001b), containing the species 0. aurula,
was described very recently. MCCRANIE &
CASTANEDA (2004a) described a second spe-
cies, 0. texiguatensis, based on another
single specimen, also thought to be a fema-
le. MCCRANIE (2006c) reported the first
known male of 0. aurula and described its
hemipenis and coloration in life. So far, the
genus Omoadiphas is known from two
apparently subadult females and one male,
making it one of the most poorly known
snake genera in the Neotropics. The holo-
type of 0 . texiguatensis was crawling in
leaf litter next to a rotton log with only the
snake's tail exposed when first sighted; its
body was under the leaves (MCCRANIE &
CASTANEDA 2004a). The holotype of 0 .
aurula was found on a coffee plantation
near the village of Buenos Aires below
Cusuco National Park. It is a very delicate
colubrid that presumably digs in the
ground and leads a highly secretive exi-
stence.
Omoadiphas aurula KOHLER, WILSON &
MCCRANIE 2001, Senckenbergiana biologica
81: 271; type locality: near Buenos Aires, 1250
m elevation, Cortes, Honduras. TL to 289
mm. Cusuco region in northwestern
Honduras, 1250-1800 m elevation.
Omoadiphas texiguatensis MCCRANIE &
CASTANEDA 2004a, Proc. Biol. Soc. Washington
117: 312; type locality: "approximately 2.5
airline km NNE of La Fortuna, 15°25'49"N,
87°18'32"W, 1690 m elev., Cerro Texiguat
Wildlife Refuge, Departamento de Yoro,
Honduras". TL to 169 mm. Known only from
the type locality.
Key to Omoadiphas
1 a 24-39 subcaudals; 7 supralabials; 8
infralabials; two postoculars; posterior
nasal separated from prefrontal by loreal;
ventral surfaces pale brown to yellow in
preservative .......... Omoadiphas aurula
b 47 subcaudals; 6 supralabials; 7 infralabi-
als; one postocular; posterior nasal contac-
ting prefrontal; ventral surfaces dark
brown to nearly black in preservative
.................... Omoadiphas texiguatensis

Fig. 695. Omoadiphas texiguatensis (holotype).
Photo: J . R. McCranie
Fig. 696. Omoadiphas aurula (holotype). From
KOHLER et al. (2001b)
• Omoadiphas aurula
..,. Omoadiphas texiguatensis
Colubridae
Fig. 697. Omoadiphas aurula (Cusuco,
Honduras). Photo: J. R. McCranie
Further Reading
KOHLER et al. 200lb, MCCRANIE & CASTANEDA
2004a, MCCRANIE 2006c
Oxybelis
Vine snakes are characterized by their
very slender bodies and conspicuously
narrow and pointed heads. All species also
lack a loreal. They are rear-fanged snakes,
with mild venom that does not pose a
serious threat to humans, but can lead to
localized swelling and itching (CRIMMINS
1937). Large specimens need to be handled
with particular caution. When a vine
snake is handled, it will react with a
threatening gape, exposing the blue-black
interior of its mouth; it will also bite with-
out any warning. As a defensive reaction,
this colubrid will continuously hold its
outstretched tongue rigid with the two tips
of the tongue pressed together (compare
Fig. 701). The precise adaptive advantage
conferred by this unusual behavior
remains a mystery to scientists. All
Oxybelis species are diurnal and spend the
night coiled up asleep in shrubs or trees
up to 3 meters above the ground. They
257
 Colubridae

Fig. 698. Oxybelis brevirostris (near Santa Fe, Fig. 701. Oxybelis brevirostris (Cerro Saslaya,
Panama). Photo: G. Kohler Atlantico Norte, Nicaragua). Photo: G. Kohler

inhabit savanna, dry forest and wet forest,

Fig. 699. Oxybelis fulgidus (Gamboa Forest
Resort, Panama, Panama). Photo: M. Lundberg
Fig. 700. Oxybelis aeneus (Bartola, Rio San
Juan, Nicaragua). Photo: G. Kohler
258
where they spend most of their time in
shrubs, high grass, and trees anywhere
from 50 to 300 cm high. They can also be
seen on the ground (FRANZEN 1996,
personal observation). In vegetation, they
are well-camouflaged and very difficult to
spot. A large part of their diet is made up
of anoles (Norops), but other lizards are
also eaten along with frogs, rodents, birds
and insects (STUART 1948, H. SMITH &
GRANT 1958, HENDERSON 1982, CAMPBELL
1998a). On Utila, Honduras, I observed a
green vine snake (0. fulgidus) as it caught
and swallowed a mouse. Vine snakes repro-
duce oviparously, laying clutches of 3-5 (0.
aeneus), 8-10 (0. fulgidus) and 8 eggs (0.
wilsoni), respectively, during the egg-lay-
ing period in the northern portion of
Central America, which falls between
March and the beginning of September
(SEXTON & HEATWOLE 1965, GoLDER 1983,
CENSKY & McCOY 1988, GROVES 1995,
CAMPBELL 1998a). A clutch of 0. fulgidus
hatched after an incubation period of
slightly more than three months (CONNERS
1989), as compared to an incubation period
of92-94 days for eggs ofO. wilsoni (GROVES
1995).

Oxybelis aeneus (WAGLER 1824), Sp. Nov.
Serp. Bras.: 12; type locality: forest adjacent
to the Solimoens River, near Ega [Tefe),
Brazil. TL to 1455 mm. Southern Tamaulipas,
Mexico, and southern Arizona, USA, as far as
southern Brazil, sea level to approximately
1500 m elevation in savanna, wet forest, and
dry forest. No subspecies are recognized
(KEISER 1974).
Oxybelis breuirostris (COPE 1861b), Proc.
Acad. Nat. Sci. Philadelphia 12: 555; type
locality: "Veraguas, New Grenada" [Panama) .
TL to 1200 mm. Eastern Honduras to
western Ecuador, sea level to approximately
900 m elevation in rain forest.
Oxybelis fulgidus (DAUDIN 1803), Hist. Nat.
Rept. 6: 352; type locality: neighborhood of
Port-au-Prince, Santo Domingo [in error]. TL
to 2000 mm. Isthmus ofTehuantepec, Mexico,
to Argentina, sea level to approximately 1000
m elevation in savanna, dry forest and wet
forest.
Oxybelis wilsoni VILLA & MCCRANIE 1995,
Rev. Biol. Trop. 43 (1-3): 297; type locality:
Rocky Point, Santa Elena on the eastern end
of Isla de Roatan, Islas de la Bahia,
Honduras. TL to 1982 mm. Endemic on Isla
de Roatan, Honduras, sea level to 200 m ele-
vation in dry forest and wet forest.
Fig. 702. Oxybelis wilsoni (Isla de Roatan,
Honduras). Photo: J. R. McCranie
Colubridae
Key to Oxybelis
1 a 15 dorsal rows at midbody; cloaca! scute
undivided; usually 6 supralabials .............. .
.............................. Oxybelis brevirostris
b 17 dorsal rows at midbody; cloaca! scute
divided; usually 8-10 supralabials ........... 2
2 a Dorsal surface of head, body, and tail green
in life (blue in preservative) ...................... ..
.. ............................... Oxybelis fulgidus
b Dorsal surface of head, body and tail yel-
low, brown, or gray ................................ 3
3 a Dorsal surface of head, body, and tail uni-
formly yellow or yellow-brown; usually 10
supralabials ................... Oxybelis wilsoni
b Dorsal surface of head, body, and tail
brown or gray with small dark flecks;
usually 8-9 supralabials Oxybelis aeneus
Further Reading
BOGERT & OLIVER 1945, KEISER 1974, 1982,
1991, VILLA & MCCRANIE 1995, GROVES 1995,
FRANZEN 1996
:= Oxybelis brevirostris
Oxybelis aeneus
.a. Oxybelis wilsoni
Oxybelis fulgidus
259
 Colubridae

Oxyrhopus

The Neotropical genus Oxyrhopus consists

of about a dozen species, which are distrib-
uted predominantly in South America.
Only one species, 0. petola, occurs in
Central America. In the past, there was
some discussion as to whether "petola" or
"petolarius" should be used as the species
name for the Central American Oxyrhopus
species (BAILEY 1986, H. SMITH et al.
1986b). BAILEY (1986) was able to show
that "petola" was correct. This colubrid is a
strong, crepuscular and nocturnal ground-
dweller. Its diet is made up primarily of Fig. 703. Adult Oxyrhopus petola (Cerro
Saslaya, Nicaragua). Photo: G. Kohler
lizards, snakes, and rodents. Oxyrhopus
petola is oviparous and produces clutches
of 5-10 eggs, which hatch after a period of
approximately three months into young
that are about 20 cm long (FITCH 1970,
CAMPBELL 1998a).

Oxyrhopus petola (LINNAEUS 1758), Systema

Naturae, ed. 10: 225; type locality: "Africa" [in
error). TL to 2200 mm. Veracruz, Mexico, to
Bolivia and Northern Brazil, sea level to
approximately 800 m elevation in rain forest.
In Central America, the subspecies
Oxyrhopus p. sebae DUMERIL, BIBRON &
DUMERIL 1854 occur.

Further Reading
BAILEY 1970b, 1986, H. SMITH et al. 1986b

• Phimophis guianensis
Oxyrhopus petola Fig. 704. Juvenile Oxyrhopus petola (4 km
south of Ayapal, Nicaragua). Photo: G. Kohler
260
 Colubridae

Phimophis

The genus Phimophis includes four mid-

size rear-fanged snakes that are character-
ized by their protruding, pointed rostral
shield. Only one species, P. guianensis,
occurs in southern Central America. It is a
secretive ground-dweller about which little
is known. There are reports that this snake
is predominantly nocturnal, though it can
also be seen during the day, and that it
feeds on lizards and small mammals
(LANCINI & KORNACKER 1989). According to Fig 706. Pituophis lineaticollis (San martin,
these authors, P. guianensis reproduces by Guatemala). Photo: L. Melendez
laying eggs.

Phimophis guianensis (TROSCHEL 1848),

Reise nach Britisch-Guiana: 653; type locali-
ty: Savannah near Pirara, Guyana. TL to 960
mm. Panama, Colombia, Venezuela and
Surinam; sea level to approximately 500 m
elevation in open dry forest, as well as in
savanna, pampas and semi-desert.
Further Reading
MARTINS & OLIVEIRA 1998, ZAHER 1999
Pituophis
Only one species of the genus Pituophis, P.
lineaticollis, reaches to northern Central
America. The others are distributed in
North America.
Pituophis lineaticollis (COPE 1861e), Proc.
Acad. Nat. Sci. Philadelphia 13: 300; type
locality: Mexico. TL to 2100 mm. Pacific side
of Mexico from Michoacan and Morelos to
central Oaxaca, as well as in the highlands of
Chiapas, Mexico, and Guatemala, 800-2500 m
elevation in wet forest and cloud forest. In
Central America, the subspecies Pituophis l.
gibsoni STUART 1954 occurs.

Further Reading
STULL 1940, DUELLMAN 1960

Pituophis lineaticollis

Fig. 705. Phimophis guianensis (Colombia).

Photo: H. W. Greene
261
 Colubridae

Fig. 707. Pituophis lineaticollis (Jaltenango,
2000 m, Chiapas, Mexico). Photo: A. Ramirez V.
Pliocercus
The false coral snakes of the genus
Pliocercus were placed by SAVAGE &
CROTHER (1989), together with the species
of the Rhadinaea lateristriga group (sensu
MYERS 1974) in the genus Urotheca. While
the partitioning of the lateristriga group
into a genus separate from Rhadinaea is
justified, it also seems meaningful to sepa-
rate the Pliocercus species into their own
genus (which would presumably form a
sister genus to Urotheca). Aside from mor-
phological differences, this decision is justi-
fied by the evolutionary history of
Pliocercus as a mimic of the true coral
snakes (genus Micrurus), which is not the
case with the Urotheca species (MYERS &
CADLE 1994, H. SMITH et al. 1995a). H.
SMITH & CHISZAR (1996) have presented
the most extensive and recent revision of
the genus Pliocercus, although it has been
strongly criticized (WILSON & MCCRANIE
1997).

The long tail, which will break off easily if

it is gripped tightly, is characteristic of
these colubrids. The fact that most speci-
mens of Pliocercus that have been found
have incomplete tails suggests that the
breakable tail is a survival strategy to pro-
tect these species from predators (SAVAGE
& CROTHER 1989). STUART (1948) found the
Fig. 708. Pliocercus elapoides (Maya Mountains,
Belize). Photo: P. Stafford tail of a Pliocercus elapoides in the stomach
of an Indigo Snake (Drymarchon corais ).
Although these colubrids are generally
harmless, they will bite if they feel
threatened. In humans, the bite of a
Pliocercus will, at most, cause a painful
localized reaction and swelling of the
lymph nodes of the affected extremity
(SEIB 1980). All Pliocercus species are noc-
turnal ground-dwellers that feed on sala-
manders, frogs, and amphibian spawn
(STUART 1948, DUELLMAN 1963, LEE 1996).
During the rainy season and also during
the dry season, these colubrids will pro-
duce clutches of 4-8 eggs (CAMPBELL 1998a,
STAFFORD 1999b).
Fig. 709. Pliocercus euryzonus (La Selva,
Heredia, Costa Rica). Photo: G. Kohler
262
 Colubridae

Fig. 710. Pliocercus euryzonus (Rio Chilagres, Fig. 711. Juvenile Pseudoboa neuwiedi (road
Panama). Photo: S. Lotzkat David-Gualaca, Chiriqui, Panama).
Photo: M. Lundberg

Pliocercus elapoides COPE 1860d, Proc. Acad.

Nat. Sci. Philadelphia 12: 253; type locality:
Near Jalapa, Veracruz, Mexico. TL to 657
mm. Oaxaca and central Veracruz, Mexico, to
central Honduras and El Salvador, sea level
to 2000 m elevation in wet forest, cloud
forest, and dry forest. In Central America, the
subspecies P. e. aequalis (SALVIN 1861b)
(eastern Chiapas to central Honduras as well
as all across the Yucatan Peninsula), P. e.
andrewsi SMITH 1942d (the north of the
Yucatan Peninsula) and P. e. diastema .t.. Pseudoboa neuwiedii
(BocoURT 1886) (Oaxaca, Mexico, along the Pliocercus euryzonus
Pacific side to eastern El Salvador) occur.
Pliocercus euryzonus COPE 1862a, Proc. Acad.
Nat. Sci. Philadelphia 14: 72; type locality: Pliocercus elapoides
Region of Truando, New Grenada, Colombia.
TL to 500 mm. Northeastern Honduras to
Colombia and Ecuador, sea level to 1300 m
elevation in rain forest and cloud forest. In
Central America, the subspecies P. e. dimid-
iatus COPE 1865 (northern Nicaragua to
eastern Costa Rica) and P. e. burghardti
SMITH & CHISZAR 1996 (Panama) occur.

Key to Pliocercus
1 a Body with three colors (black-red-yellow
or black-red-white rings) ........................
... ........ ... ............ ..... Pliocercus elapoides Further Reading
b Body with only two colors (black-red or SAVAGE & CROTHER 1989, SMITH & CHISZAR 1996,
black-white rings) .. Pliocercus euryzonus WILSON et al. 1996, WILSON & MCCRANIE 1997
263
 Colubridae
Fig. 712. Adult Pseudoboa neuwiedi (Gamboa
Forest Resort, Panama, Panama).
Photo: M. Lundberg
Pseudoboa
The genus Pseudoboa, which is distributed
predominantly in South America, includes
four species, only one of which, P. neu-
wiedii, reaches to the southern extreme of
Central America. This rear-fanged snake is
nocturnal and lives as a ground-dweller in
dry forest and rain forest, generally in the
vicinity of bodies of water. Aside from
lizards, their diet also includes small mam-
mals (LANCINI & KORNACKER 1989).
Pseudoboa neuwiedii (DUMERIL, BIBRON &
DUMERIL 1854), Erp. Gen. 7: 1001; type locali-
ty: Cote Ferme and Brazil. TL to 1200 mm.
Panama, as well as all across tropical South
America, sea level to approximately 1000 m
elevation in dry forest and rain forest.
Further Reading
MARTINS & OLIVEIRA 1998, ZAHER 1999
Fig. 713. Pseustes poecilonotus (Bartola, Rio
San Juan, Nicaragua). Photo: G. Kohler
264
Pseustes
Pseustes is a Neotropical genus with four
recognized species, one of which is distri-
buted throughout Central America.
Pseustes poecilonotus lives primarily in old
growth rain forest, where it can be found
both on the forest floor, as well as in
shrubs and trees. If it feels threatened, it
will inflate its upper body, open its mouth
wide and bite without hesitation, which, in
view of the size of some specimens, can
inspire considerable respect. Pseustes poe-
cilonotus is diurnal and feeds primarily on
birds, small tree-dwelling mammals, and
bird eggs (SEXTON & HEATWOLE 1965,
ALVAREZ DEL TORO 1983, CAMPBELL 1998a).
The species of the genus Pseustes repro-
duce oviparously, with females laying 7-14
eggs (GOODE 1989).
Pseustes poecilonotus (GONTHER 1858), Cat.
Snakes Brit. Mus. 1858: 36; type locality:
Honduras and Mexico; restricted to
Honduras through lectotype designation by
BOULENGER (1894a). TL to 2100 mm. San Luis
Potosi and Oaxaca, Mexico, as far as Brazil,
sea level to approximately 1000 m elevation
in dry forest, wet forest, and rain forest. The
division into subspecies according to AMARAL
(1929a) seems problematic (WILSON & MEYER
1985). Phrynonax shropshirei BARBOUR &
AMARAL 1924 is regarded to be a synonym of
P. poecilonotus according to SAVAGE (2002).

Fig. 714. Juvenile Pseustes poecilonotus
(Manuel Antonio, Costa Rica). Photo: G. Kohler
Fig. 715. Pseustes poecilonotus (Cerro Saslaya,
Atlantico Norte, Nicaragua). Photo: G. Kohler
Pseustes poeci/onotus
Further Reading
AMARAL 1929a
Colubridae
Rhadinaea
The genus Rhadinaea includes just over 40
species, 21 of which occur in Central
America. Next to Tantilla, it is the genus
with the most species in this faunal region.
These are harmless, small to mid-size colu-
brids that live as diurnal ground-dwellers,
predominantly in forest areas. They are
most commonly encountered crawling over
the leaf-covered ground or crossing a forest
path. They lead a highly secretive exist-
ence, and can also be found by turning over
logs, roots, or large rocks. Their diet con-
sists mostly of salamanders, frogs
(Eleutherodactylus), and lizards, as well as
the terrestrial eggs of these animals
(MYERS 1974). Some species will also con-
sume invertebrates such as earthworms
(TAYLOR 1949a). All species of this genus -
even those that occur in the highlands -
are believed to be oviparous and produce
clutches of 2-13 eggs (MYERS 1974).
Fig. 716. Rhadinaea stadelmani (near
Chiantla, Cordillera de los Cuchumatanes,
Huehuetenango, Guatemala). Photo: T. Bille
265
 Colubridae

Rhadinaea anachoreta SMITH & CAMPBELL

Fig. 717. Rhadinaea anachoreta (El Paraiso
Valley Reserve, Cortes, Honduras).
Photo: J . R. Mccranie
Fig. 718. Rhadinaea calligaster (Jurutungo,
Chiriqui, Panama).
Fig. 719. Rhadinaea decorata (Bartola, Rio
San Juan, Nicaragua).
1994, Occ. Pap. Mus. Nat. Hist. Univ. Kansas
167: 2; type locality: N slope of Cerro de
Aguacate, Aldea Negro Norte, Sierra de
Caral, Municipio de Morales, Izabal,
Guatemala. TL to 251 mm. Eastern
Guatemala and western Honduras, 500-1180
m elevation in rain forest.
Rhadinaea calligaster (COPE 1876), J. Acad.
Nat. Sci. Philadelphia (2) 8: 146; type locality:
Pico Blanco, Costa Rica. TL to 513 mm.
Central Costa Rica to western Panama, 1220-
2440 m elevation in rain forest and cloud
forest.
Rhadinaea decorata (GONTHER 1858), Cat.
Snakes Brit. Mus. 1858: 35; type locality:
Mexico. TL to 470 mm. Caribbean side of San
Luis Potosi, Mexico, to northwestern Ecuador,
sea level to 1200 m elevation in rain forest.
Rhadinaea godmani GONTHER 1865, Ann.
Mag. Nat. Hist. 3 (15): 94; type locality:
Duenas, Guatemala. TL to 568 mm.
Discontinuous from southeastern Oaxaca,
Mexico, into western Panama, 1000-2650 m
elevation in pine-oak forest and cloud forest.
Rhadinaea hannsteini (STUART 1949), Proc.
Biol. Soc. Washington 62: 165; type locality:
Finca La Paz, 1450 m elevation, 18 km N
Coatepeque, San Marcos, Guatemala. TL to
393 mm. Southeastern Chiapas, Mexico, and
southwestern Guatemala, 500-2000 m eleva-
tion on coffee plantations and in rain forest.
Rhadinaea hempsteadae STUART & BAILEY
1941, Occ. Pap. Mus. Zool. Univ. Michigan
442: 2; type locality: above Finca Chichen,
about 5700 ft elevation, Alta Verapaz,
Guatemala. TL to 593 mm. Alta Verapaz and
Photo: S. Lotzkat Sierra de las Minas in eastern Guatemala,
1200-3000 elevation in pine-oak forest and in
cloud forest.
Rhadinaea kanalchutchan MENDELSON &
KlzIRIAN 1995, Herpetologica 51: 302; type
locality: 8 km S San Cristobal de las Casas,
Chiapas, Mexico. TL to 490 mm. Western por-
tion of the Meseta Central de Chiapas,
Mexico, 2300-2700 m elevation in pine-oak
forest.
Rhadinaea kinkelini BOETTGER 1898, Katalog
der Reptilien-Sammlung im Museum der
Senckenbergischen naturforschenden Gesell-
schaft 2: 68; type locality: Matagalpa, Nicara-
gua. TL to 376 mm. Central Guatemala to
Central Nicaragua, 1300-2200 m elevation in
cloud forest. Rhadinaea pinicola MERTENS
1952b is regarded to be a synonym of R. kin-
kelini according to KOHLER & MCCRANIE
Photo: G. Kohler (1999b).

266
 Colubridae

Fig. 720. Rhadinaea decorata (Nusagandi, Fig. 723. Rhadinaea kanalchutchan (8.6 mi SE
Comarca de San Blas, Panama). San Cristobal de las Casas, Chiapas, Mexico).
Photo: G. Kohler Photo: C. W. Myers

Fig. 721. Rhadinaea godmani (Alto Chiquero, Fig. 724. Rhadinaea kinkelini (Miraflores,
Chiriqui, Panama). Photo: G. Kohler Nicaragua). Photo: J . Sunyer

Fig. 722. Rhadinaea hempsteadae (baja Fig. 725. Rhadinaea lachrymans (Pico La Pi-
Verapaz, Guatemala). Photo: J. A Campbell cucha, Sierra de Agalta, Olancho, Honduras).
Photo: J . R. McCranie
267
 Colubridae
Rhadinaea lachrymans (COPE 1870), Proc.
Amer. Phil. Soc. 11: 154; type locality: un-
known. TL to 493 mm. Southeastern Chiapas,
Mexico, and southwestern Guatemala as well
as in the Sierra de Agalta in east-central
Honduras (see MCCRANIE & WILSON 2001a),
500-3000 m elevation in pine-oak forest.
Rhadinaea macdougalli SMITH & LANGE-
BARTEL 1950, J . Washington Acad. Sci. 39: 413;
type locality: Buena Vista, at crest of Sierra
Madre, 4000-4500 ft, directly north of Rio
Grande, Oaxaca, Mexico. TL to 292 mm.
Pacific side of the Isthmus of Tehuantepec
(eastern Oaxaca, Mexico), 1200-2000 m eleva-
tion in rain forest.
Rhadinaea montecristi MERTENS 1952b, Zool.
Anz. 149: 136; type locality: Hacienda
Montecristo, 2200 m elevation, mountains of
Metapan, Santa Ana, El Salvador. TL to 555
mm. Northwestern El Salvador, eastern
Guatemala, and western Honduras, 1300-
2600 m elevation in cloud forest.
Rhadinaea pegosalyta MCCRANIE 2006b, Proc.
Biol. Soc. Washington 119: 528; type locality:
"near visitor's center of Parque Nacional El
Cusuco, l5°3l'N, 88°12'W, 1550 m elev.,
Departamento de Cortes, Honduras." TL to
355 mm. Known only from type locality.
R. decorata
• R. kinkelini
.., R. serperaster
.a. R. anachoreta
Rhadinaea
• R. cal/igaster
=: R. pulveriventris
.a. R. sargenti
.., R. vermiculaticeps
Rhadinaea
268
Rhadinaea pilonaorum (STUART 1954), Proc.
Biol. Soc. Washington 67: 176; type locality:
Finca La Gloria, about 950 m elevation, about
12 km northeast of Chiquimulilla, Santa
Rosa, Guatemala. TL to 310 mm.
Southeastern Guatemala and western El
Salvador, 670-950 m elevation in wet forest.
Rhadinaea posadasi (SLEVIN 1936), Proc.
Calif. Acad. Sci. 23: 79; type locality: southern
slope of Volcan Zunil, Suchitepequez,
Guatemala. TL to 288 mm. Southwestern
Guatemala, 500-1800 m elevation in wet
forest.
Rhadinaea pulueriventris BoULENGER 1896a,
Cat. Snakes Brit. Mus. 3: 635; type locality:
Azahar de Cartago, Costa Rica. TL to 502
mm. Central Costa Rica to western Panama,
1370-1600 m elevation in rain forest and
cloud forest.
Rhadinaea rogerromani KOHLER & MCCRANIE
1999a, Senckenbergiana biol. 79: 244; type
locality: S slope of Cerro Saslaya (13°46.0'N,
85°02.3'W), 1450 m elevation, Atlantico
Norte, Nicaragua. TL to 286 mm. Known only
from the type locality.
Rhadinaea sargenti DUNN & BAILEY 1939,
Bull. Mus. Comp. Zool. 86: 10; type locality:
. R. godmani
.a. R. montecristi
.., R. rogerromani
Rhadinaea
./
i-; R. hannsteini
.a. R. stade/mani
..- R. hempsteadae
• R. kanalchutchan
• R. pilonaorum
~ ~~'~
,. . / ~:.. - :_ _ ~~1
- ~..... ,, / " (
~ J (
Rhadinaea " ~- , (j
 Colubridae

Pequeni-Esperanza Ridge, 1800 ft, near head

of Rio Pequeni, Panama. TL to 310 mm. East
of the Canal Zone in central Panama, 300-770
m elevation in rain forest.
Rhadinaea serperaster COPE 1871, Proc.
Acad. Nat. Sci. Philadelphia 23: 212; type
locality: near San Jose, Costa Rica. TL to 445
mm. Central Costa Rica, 1220-1450 m eleva-
tion in rain forest and cloud forest.
Rhadinaea stadelmani STUART & BAILEY
1941, Occ. Pap. Mus. Zool. Univ. Michigan
442: 4; type locality: Todos Santos, 8000 ft ele-
vation, Huehuetenango, Guatemala. TL to
450 mm. Sierra de los Cuchumatanes and
Montafias del Cuilco im western Guatemala.
Rhadinaea tolpanorum HOLM & CRUZ D.
1994, Herpetologica 50: 17; type locality: 0.8 Fig. 727. Rhadinaea lachrymans (near Finca
km N and 1.5 km E Cerro San Francisco in Irlanda, Chiapas, Mexico). Photo: G. Kohler
the Cordillera Nombre de Dios, 1900 m eleva-
tion, Atlantida, Honduras. TL to 381 mm.
Known only from the type locality (cloud
forest).
Rhadinaea vermiculaticeps (COPE 1860d),
Proc. Acad. Nat. Sci. Philadelphia 12: 249;
type locality: Veragua, Panama. TL to 374
mm. Western Central Panama, 700-900 m
elevation in rain forest.

Fig. 728. Rhadinaea lachrymans (near Finca

Irlanda, Chiapas, Mexico). Photo: G. Kohler

Fig. 726. Rhadinaea pilonaorum (Finca La

Giralda, La Libertad, El Salvador).
Photo: E. Greenbaum

.. ) R. /achrymans
(:7 .t.R. macdougalli
"" R. tolpanorum
• R. posadasi
'- • R. pegosalyta

~ , "{ \ -;-~-,,
• / :_\t r 'i
Rhadinaea ~. - ~ ~ Fig. 729. Rhadinaea posadasi (Quetzal-
tenango, Guatemala). Photo: J. A. Campbell
269
 Colubridae
Key to Rhadinaea
1 a 19 or 21 dorsal rows at midbody .............. 2
b 17 dorsal rows at midbody ............. ....... ... 9
2 a 21 dorsal rows at midbody ................ .... .. . 3
b 19 dorsal rows at midbody .... ............. ... .. . 5
3 a Dorsal surfaces usually some shade of
brown and ventral surfaces always yellow
or white .. ........ ..... Rhadinaea godmani
b Red dorsal and ventral surfaces in life ... 4
4 a Middorsal ground color the same color as
the lateral ground color; stripes on adja-
cent edges of scale rows eight and nine
poorly developed and broken ......... ..... .... .
.. . . . . . . . . . . . . ....... .... Rhadinaea pegosalyta
b Middorsal ground color much darker than
lateral ground color; stripes on adjacent
edges of scale rows eight and nine well
developed, continuous ................. ... ... .. ..... .
............. ......... ... Rhadinaea rogerromani
5 a Very wide and contrasting dark longitud~
nal stripes present, lateral stripes (on 3r
dorsal row and adjacent scales) and ven-
trolateral stripes (on the ends of the ven-
trals and 1st dorsal row) ............... .. .... .... . .
..... .. .... ..... ... .. .. Rhadinaea montecristi
b Longitudinal stripes narrower than above
and not as obviously contrasting as above
....... .. .......... .. .................... ...... ..... ............. 6
6 a A conspicuous, narrow, dark longitudinal
stripe 'J1nning along the borders of the 1st
and 2n dorsal rows .............. .......................
..... ...... ..... ......... Rhadinaea serperaster
b Without a longitudinal stripe along the
borders of the 1st and 2nd dorsal rows, al-
though other longitudinal stripes may be
present ....... .... ... ........ ..................... .......... 7
7 a A dark loncitudinal stripe running along
the 6t1i, 7tfi or gth dorsal rows; 169-191
ventrals ...... .... . Rhadinaea stadelmani
b Without s.uch a longitudinal stripe on the 6 th,
7th or gth dorsal rows; 156-175 ventrals 8
8 a 13-15 maxillary teeth; striped or unicolor
... ..... ............... Rhadinaea hempsteadae
b 16-18 maxillary teeth; without longitudi-
nal stripes Rhadinaea kanalchutchan
9 a A postocular pale spot or longitudinal
stripe beginning at the upper posterior cor-
ner of the eye (Fig. 730a) .. .............. .. ..... 10
b If a postocular pale spot or longitudinal
stripe is present, it begins at the lower
posterior corner of the eye (Fig. 730b) .... 14
lOa Broad dark vertebral stripe intermittent
along the middle of a pale line, such that
270
the markings could be mistaken for a pair
of paravertebral stripes; conspicuous pale
reticulum on dorsal surface of head ........... .
... ....... .. .. ... Rhadinaea vermiculaticeps
b Dark vertebral stripe, if present, not inter-
mittent along the middle of a pale line;
pale reticulum on dorsal surface of head
present or not .................. ...................... 11
lla Lower (lSt to 3rd) dorsal scale rows uni-
color black, sharply contrasting with pale
mediodorsal area; conspicuous pale reticu-
lar markings on top of head .............. .
.. .... ............... ... .. ... Rhadinaea sargenti
b Dorsal and head markings different than
those above ...... ... ........ ...... .. ...... .. ... .... 12
12a Dorsum either with a dark dorsol~teral
line along the border of both the 4t and
5th dorsal rows or predominantly unicolor
brown ......... ......................... .................. ...... .
...... Rhadinaea pulveriventris (in part)
b Dorsal and head markings different ...... 13
13a Two pale spots, one of which is directly
behind the eye, the other on the neck; these
spots may also blend into a pale stripe that
runs horizontal posteriorly ......................... .
. ...... ............ ... .... . Rhadinaea decorata
b From the eye, a pale stripe runs obliquely
across temporal region .... ........... ............. .
.. ........ .... ... .. .. Rhadinaea macdougalli
14a A series of dark spots, bars or a continuous
stripe along the ventromedial line ........... . .
.... ...... ........ ..... . Rhadinaea calligaster
b Ventral markings different ............. ....... 15
15a At least one obvious dark longitudinal
stripe present on side of body ...... .. .... .. 16
b Body without obvious dark longitudinal
stripes or only with dark vertebral stripes
... .... ... .............. ... .. .... ... ... .. .. ....... .. ... 21
16a Dark vertebral stripe short and restricted
mostly to neck area; a black stripe present
on dorsal surface of head ............. ..... ... ..... .
..... Rhadinaea pulveriventris (in part)
b Dorsal and head markings different than
those above .. ............... ... .... .. .. ..... ....... 17
17a Venter red to red-orange (in life); dorsum
with three dark longitudinal stripes (verte-
br;il stripe plus lateral stripes along the
4th dorsal row and adjacent areas of the
3rd row) .... ........ Rhadinaea tolpanorum
b Venter white or cream colored; dorsal
markings different than those above ..... 18
18a Without a large, pale spot on each frontal
shield, dorsal surface of head uniformly
dark (Fig. 731a); fewer than 153 ventrals;
one postocular .. ....... ........... ........ ..... .... 19
 Colubridae

b A large, dark spot on each frontal shield

usually clearly formed (Fig. 731b); if with-
out pale frontal spot, then more than 153
ventrals; usually two postoculars .......... 20
19a One posterior temporal; dark vertebral
stripe covers the vertebral and parts of the
paravertebral row ..... .. ......... .... ...... .... ... . .
. . . . . . .. . . . . . . ........... Rhadinaea anachoreta
b Two posterior temporals; dark vertebral
stripe restricted to medial third of the ver-
tebral row ....... ... Rhadinaea hannsteini
20a Dorsal pattern dominated by pairs of la-
teral and ventrolateral longitudinal stripes
... ......... .. .. ..... ... Rhadinaea lachrymans Fig. 732. Rhadinaea pulueriventris (Fortuna,
b Dorsal pattern more complex and not Chiriqui, Panama). Photo: C. W. Myers
dominated by pairs of lateral and ventro-
lateral longitudinal stripes ........................ .
. . . . . . .... . . . . . . . . . . . . ...... Rhadinaea kinkelini
21a More than 150 ventrals ....................... ... . .
. . . . . . . . . . . . . . . .... .. ... Rhadinaea pilonaorum
b Fewer than 147 ventrals .......................... .
. . . . . . . . . . . . . . . .............. Rhadinaea posadasi

a. R. decorata

Fig. 733. Rhadinaea rogerromani (holotype).

Photo: G. Kohler

Fig. 730. Lateral head pattern in Rhadinaea.

a. R. montecristi b. R. kinkelini Fig. 734. Rhadinaea rogerromani (holotype).

Photo: G. Kohler
Fig. 731. Dorsal head pattern in Rhadinaea.
271
 Colubridae

Salvadora

The species of the genus Salvadora all

have a conspicuously large rostral shield,
that extends further upward on the snout
than in other colubrids. These are agile,
diurnal snakes that inhabit the dry areas
of the southwestern US and Mexico. Only
one species, S. lemniscata, ranges as far as
the northern portion of Central America. It
feeds primarily on lizards (BOGERT 1939)
and will lay clutches of 5-12 eggs, which
hatch after 3-4 months of incubation (OBST
et al. 1984, D. SCHMIDT 1990).
Fig. 739. Rhinobothryum bovallii.
Photo: R. W. Van Devender Salvadora lemniscata (COPE 1895), Trans.
Amer. Philos. Soc. 18: 203; type locality:
western Mexico. TL to 1300 mm. Along Pacific
coast of Mexico from Guerrero to Chiapas;
also in Grijalva Valley of Chiapas and adja-
cent Guatemala; sea level to 1000 m elevati-
on on savannas and in dry forest.

Further Reading
BOGERT 1939, WERLER & SMITH 1952

Fig. 740. Hatchling of Rhinobothryum bovallii

(Reptilandia, Costa Rica). Photo: G. Kohler

Rhinobothryum bovallii ANDERSSON 1916,

Medd. Goteborgs Mus. Zool. Afd. 9: 32; type
locality: Siquirres, Costa Rica. TL to 1762 mm
(see SOLORZANO 2003). Costa Rica to Ecuador
and Venezuela, also recorded in eastern
Honduras, sea level to 600 m elevation in rain
forest.

Further Reading
HOHMEISTER 2001, HAMMACK & BRINKER 2008
Fig. 741. Salvadora lemniscata (near Mix-
tequilla, Oaxaca, Mexico). Photos: G. Kohler

273
 Colubridae
Scaphiodontophis
Contrary to the earlier view of the genus
8caphiodontophis as monotypic (SMITH et
al. 1986a, SAVAGE & SLOWINSKI 1996), the
current view is that there are two different
species, one of which is distributed in
northern Central America (8. annulatus),
the other in the south (8. uenustissimus)
(MCCRANIE 2006d). 8caphiodontophis
annulatus lives on the leaf strewn ground
of the rain forest and is an agile, diurnal
hunter, that feeds primarily on lizards
(SAVITZKY 1981, HENDERSON 1984). Their
hinged and shovel-shaped teeth are parti-
cularly well suited to holding onto their
primary prey, the smooth, slender skink
8phenomorphus cherriei. 8caphiodont-
ophis annulatus reproduces oviparously,
laying clutches of 1-10 (usually 2-4) eggs
under rotting logs or in other suitable
hiding places during the rainy season
(ALVAREZ DEL TORO & SMITH 1958, ALVAREZ
DEL TORO 1983, HENDERSON 1984,
CAMPBELL 1998a). Similar to the species of
the genera Pliocercus and Urotheca, 8 .
annulatus also has a very long tail that
will easily break off if the animal is seized.
Most of the specimens of this species found
have incomplete tails.
Scaphiodontophis
• S. annulatus
S. venustissimus
274
Scaphiodontophis annulatus (DUMERIL,
BIBRON & DuMERIL 1854), Erp. Gen. 7: 335;
type locality: Cohan, Alta Verapaz, Guate-
mala. TL to 900 mm. Oaxaca and southern
Tamaulipas, Mexico, to eastern Honduras,
sea level to 1300 m elevation in dry forest and
wet forest.
Scaphiodontophis venustissimus (Gl.JNTHER
1894), Biologia Centrali-Americana. Reptilia
and Batrachia: 144; type locality: Hacienda
Santa Rosa de Jerico, Matagalpa, Nicaragua.
TL to 900 mm. Eastern Honduras to
Colombia, sea level to 1300 m elevation in dry
forest and wet forest.
Key to Scaphiodontophis
1 a Subcaudals 105-121 in males, 92-105 in
females; combined ventrals plus subcau-
dals 234-262; usually with a pattern consi-
sting of black bands bordered by yellow
bands ......................... ................... ... ........... ..
..... .. .. Scaphiodontophis venustissimus
b Subcaudals 123-149 in males, 110-126 in
females; combined ventrals plus subcau-
dals 255-294; usually with a pattern consi-
sting of yellow bands bordered by black
bands ..... Scaphiodontophis annulatus
Further Reading
HENDERSON 1984, H . SMITH et al. 1986a, SAVAGE
& SLOWINSKI 1996, MCCRANIE 2006d
Fig 742. Scaphiodontophis annulatus (Tuxtla
Gutierrez, Chiapas, Mexico).
Photo: A. Ramirez V.
 Colubridae

Scolecophis

The genus Scolecophis is monotypic.

Scolecophis atrocinctus is a conspicuous
ground-dweller in the dry areas along the
Pacific coast of Central America. The orange-
red spot on its back is so bright that it
seems to have been painted on with fluor-
escent paint. The species is diurnal and
feeds primarily on insects and arachnids.
The species reproduces oviparously
(MARINEROS 2000).

Fig. 906. Scaphiodontophis venustissimus Scolecophis atrocinctus (SCHLEGEL 1837),

(near Limon, Costa Rica). Photo: A Hohmeister
Essai physion. Serpens 2: 48; type locality:
Chile [in error] . TL to 410 mm. Along the
Pacific side of eastern Guatemala into north-
western Costa Rica, sea level to 700 m eleva-
tion in dry forest and wet forest.

Further Reading
WILSON & WILLIAMS 2002

Fig. 744. Scaphiodontophis annulatus

(Veracruz, Mexico). Photo: P. Heimes

\ \_ ''-~ 0
) ,-~
'-, , ... ,
''{ \ /
-·.~· ~

~x
Scolecophis atrocinctus 1~~
r Fig. 745. Scolecophis atrocinctus.
Photo: R. W. Van Devender
275
 Senticolis

The genus Senticolis was established by

DOWLING & FRIES (1987) for the species
triaspis, which was formerly included in
the genus Elaphe. The young of this spe-
cies are conspicuously spotted, as opposed
to the unicolor yellowish brown or olive
green adult specimens. Senticolis triaspis
is a nocturnal, terrestrial to semi-arboreal
colubrid that eats primarily small mam-
mals and the occasional lizard or bird
(DOWLING 1960, CAMPBELL 1998a). Females
reach sexual maturity around the age of
two, with a snout-vent length of at least
625 mm (CAMPBELL 1998a). This oviparous
species produces clutches of 3-9 eggs,
which hatch after 75-100 days at an incu- Fig. 747. Juvenile Senticolis triaspis (El
bation temperature of 25-29 °C (SCHlJLZ lmposible National Park, Ahuachapan, El
Salvador). Photo: G. Kohler
1996).

Senticolis triaspis (COPE 1866), Proc. Acad.

Nat. Sci. Philadelphia 18: 128; type locality:
Belize. TL to 1220 mm . Southeastern
Arizona, USA, and Tamaulipas, Mexico, to
Costa Rica, sea level to 1500 m elevation in
dry forest and wet forest.

Further Reading
DOWLING 1960, DOWLING & FRIES 1987, PRICE
1991, SCHULZ 1996

Fig. 748. Senticolis triaspis (Rancho Alegre,

Chiapas, Mexico). Photo: G. Kohler

Fig. 746. Adult Senticolis triaspis (El lmposible

National Park, Ahuachapan, El Salvador). • Senticolis triaspis
Photo: G. Kohler
276
Sibon

The genus Sibon includes about a dozen

species that are distributed in Mexico,
Central America, and South America.
SMITH (1982) pointed out that the gender of
the genus name Sibon is masculine, thus
the species should be called annulatus,
dimidiatus, and nebulatus. Similar to the
species of the genera Dipsas and
Tropidodipsas, the Sibon species specialize
in snails as a food source. The morphologi-
cal adaptations that enable these colubrids
to pull their slimy prey out of their shells Fig. 749. Sibon annulatus (Cerro Saslaya,
include particularly long teeth in the lower Atlantico Norte, Nicaragua). Photo: G. Kohler
jaw and the conspicuously short snout of
most of the Sibon species. All species of
this genus are nocturnal and live predomi-
nantly in trees and shrubs. They reproduce
by laying 2-6 eggs per clutch (KOFRON
1983, 1987).

Sibon annulatus (GUNTHER 1872), Ann. Mag.

Nat. Hist. (4) 9: 30; type locality: Near
Cartago, Costa Rica. TL to 557 mm.
Northeastern Honduras to eastern Panama,
sea level to about 1200 m elevation in rain
forest.
Sibon anthracops (COPE 1868), Proc. Acad.
Nat. Sci. Philadelphia 20: 136; type locality:
Central America. TL to 664 mm. Pacific ver-
sant from El Salvador to Costa Rica; in Guate-
mala (Rio Motagua Valley) and Honduras Fig. 750. Sibon anthracops (El Salvador).
(Rio Aguan valley) also on Caribbean versant, Photo: G. Kohler
sea level to about 500 m elevation in dry
forest.
Sibon argus (COPE 1876), J. Acad. Nat. Sci.
Philadelphia (2) 8: 130; type locality: Costa
Rica. TL to 690 mm. Southeastern Costa Rica
to eastern Panama, sea level to about 600 m
elevation in rain forest.
Sibon carri (SHREVE 1951), Copeia 1951: 52;
type locality: Escuela Agricola Panamericana,
near Tegucigalpa, Honduras. TL to 408 mm.
Central El Salvador to southern Honduras,
sea level to about 1000 m elevation in dry
forest.
Sibon dimidiatus (GUNTHER 1872), Ann. Mag.
Nat. Hist. (4) 9: 31; type locality: Mexico. TL
to 680 mm. Veracruz and Oaxaca, Mexico, to
southeastern Costa Rica, sea level to about
1600 m elevation in rain forest and cloud
forest. Fig. 751. Sibon dimidiatus (Selva Negra,
Matagalpa, Nicaragua). Photo: G. Kohler
277
 Colubridae

Fig. 752. Sibon dimidiatus (Palo Seco, Fig. 755. Sibon dimidiatus (near Quebrada
Panama). Photo: G. Kohler Grande, Copan, Honduras). Photo: G. Kohler

Fig. 753. Sibon longifrenis (Costa Rica). Fig. 756. Sibon manzanaresi (San San Hi.I,
Photo: T. Leenders Gracias a Dios, Honduras).
Photo: J. R. McCranie

Fig. 754. Sibon longifrenis (Bosawas, Nicaragua). Fig. 757. Sibon miskitus (Gracias a Dios,
Photo: J. Sunyer Honduras). Photo: J. R. McCranie
278
 Colubridae

Sibon lamari SoL6RZANO 2002, Rev. Biol. • Sibon annulatus

Trop. 49: 1111; type locality: Guayacan de Sibon anthracops
Siquirres, 83°32'30"N, l0°02'10"W, Limon .& Sibon miskitus
Province, Costa Rica. TL to 589 mm. Known Y Sibon /amari
only from type locality, 550-650 m elevation.
Sibon longifrenis (STEJNEGER 1909), Proc.
U.S. Natl. Mus. 36: 457; type locality: Bocas
del Toro, Panama. TL to 624 mm. Caribbean
versant of eastern Honduras, Costa Rica, and
western Panama, 400-600 m elevation in rain
forest.
Sibon manzanaresi MCCRANIE 2007,
Herpetologica 63: 213; type locality: "San San
Hil, 14°56.809'N, 84°30.715'W, 300 m elevati-
on, Departamento de Gracias a Dios,
Honduras. TL to 407 mm. Known only from
type locality.
Sibon miskitus MCCRANIE 2006a, Journal of
Herpetology 40: 16; type locality: ''Warunta
Tingni Kiamp, 14°55.336'N, 84°41.471'W, 150
m elevation, Departamento de Gracias a
Dios, Honduras. TL to 499 mm. Montafias de
Colon in northeastern Honduras.
Sibon nebulatus (LINNAEUS 1758), Systema .& Sibon manzanaresi
Naturae, ed. 10: 222; type locality: America. Sibon argus
TL to 830 mm. Nayarit and Veracruz, Mexico, := Sibon longifrenis
to Ecuador and Brasil, sea level to about 1500
m elevation in rain and dry forest. The sub-
species Sibon n. nebulatus (Mexico to central
Panama) and Sibon n. leucomelas (BOULEN- Sibon dimidiatus
GER 1896b) (eastern Panama to northwestern 111 Sibon carri
Ecuador) occur in Central America. := Sibon sanniolus
Sibon sanniolus (COPE 1867), Proc. Acad. Nat.
Sci. Philadelphia 18: 318; type locality:
Yucatan, Mexico. TL to 430 mm. Northern
and Eastern portions of the Yucatan
Peninsula, sea level to about 300 m elevation
in dry forest. The subspecies Sibon s. sannio-
lus (northern portion of Yucatan Peninsula)
and Sibon s. neilli HENDERSON, HOEVERS &
WILSON 1977 (Belize and northeastern
Guatemala) occur in Central America.

Sibon nebulatus

Fig. 758. Sibon miskitus (Hiltara Kiamp, Gracias

a Dios, Honduras). Photo: J. R. McCranie
279
 Colubridae

Sibon lamari SOL6RZANO 2002, Rev. Biol. • Sibon annulatus

Trop. 49: 1111; type locality: Guayacan de Sibon anthracops
Siquirres, 83°32'30"N, l0°02'10"W, Limon • Sibon miskitus
Province, Costa Rica. TL to 589 mm. Known "' Sibon lamari
only from type locality, 550-650 m elevation.
Sibon longifrenis (STEJNEGER 1909), Proc.
U.S. Natl. Mus. 36: 457; type locality: Bocas
del Toro, Panama. TL to 624 mm. Caribbean
versant of eastern Honduras, Costa Rica, and
western Panama, 400-600 m elevation in rain
forest.
Sibon manzanaresi MCCRANIE 2007,
Herpetologica 63: 213; type locality: "San San
Hil, l4°56.809'N, 84°30.715'W, 300 m elevati-
on, Departamento de Gracias a Dios,
Honduras. TL to 407 mm. Known only from
type locality.
Sibon miskitus MCCRANIE 2006a, Journal of
Herpetology 40: 16; type locality: "Warunta
Tingni Kiamp, 14°55.336'N, 84°41.471'W, 150
m elevation, Departamento de Gracias a
Dios, Honduras. TL to 499 mm. Montaftas de
Colon in northeastern Honduras.
Sibon nebulatus (LINNAEUS 1758), Systema • Sibon manzanaresi '
Naturae, ed. 10: 222; type locality: America. Sibon argus '
TL to 830 mm. Nayarit and Veracruz, Mexico, = Sibon longifrenis
\
to Ecuador and Brasil, sea level to about 1500
m elevation in rain and dry forest. The sub-
species Sibon n. nebulatus (Mexico to central
Panama) and Sibon n. leucomelas (BoULEN- Sibon dimidiatus
GER 1896b) (eastern Panama to northwestern 111 Sibon carri
Ecuador) occur in Central America. = Sibon sanniolus
Sibon sanniolus (COPE 1867), Proc. Acad. Nat.
Sci. Philadelphia 18: 318; type locality:
Yucatan, Mexico. TL to 430 mm. Northern
and Eastern portions of the Yucatan
Peninsula, sea level to about 300 m elevation
in dry forest. The subspecies S ibon s. sannio-
lus (northern portion of Yucatan Peninsula)
and Sibon s. neilli HENDERSON, HOEVERS &
WILSON 1977 (Belize and northeastern
Guatemala) occur in Central America.

Sibon nebulatus

Fig. 758. Sibon miskitus (Hiltara Kiamp, Gracias

a Dios, Honduras). Photo: J. R. McCranie
279
 Colubridae

Key toSibon
1 a 13 dorsal rows at midbody ............... ........ 2
b 15 dorsal rows at midbody ......... .............. 3
2 a Without anterior temporal, first postocular
supralabial in contact with parietal (Fig.
760a) .. ..... .............................. Sibon carri
b One anterior temporal, no supralabial in
contact with parietal .. Sibon anthracops
3 a Enlarged penultimate supralabial in
contact with eye; dark dorsal bands or
spots reach at most to outer edges of ven-
trals, although dark shading may be pre-
sent on venter ...................................... ... 4
b Enlarged penultimate supralabial not in Fig. 759. Sibon sanniolus (near Coba,
contact with eye; dark dorsal bands or Quintana Roo, Mexico). Photo: J. C. Lee
spots reach well onto venter or even across
venter, or venter completely immaculate .. 5
4 a 181-201 ventrals; 112-121 subcaudals .... ..
.... .. .. .. . .. .. .. .. .. . . . .. .. .. .. . . .. ..... Sibon argus
b 151-173 ventrals; 82-106 subcaudals .........
.. .. .. .. .. .. .. .... .. .. .. .. .. .. ... Sibon longifrenis
5 a First infralabials in contact with each
other behind the mental Sibon nebulatus
b First infralabials separated from each
other by one or two postmentals behind the
mental (Fig. 761) ...... .. ............ ................ .. 6 a. S . carri
6 a 8-10 supralabials, three in contact with
eye; 143-162 ventrals; 57-88 subcaudals
.... ................ .... .. .. ........ Sibon sanniolus
b 7-8 supralabials, two in contact with eye;
161-200 ventrals; 95-133 subcaudals ....... 7
7 a Ventral surfaces immaculate .. ................... .
. . ......... .. .... ..... .. . ...... Sibon manzanaresi
b Dark dorsal bands or spots reach well onto
venter or even across venter .......... .......... 8 b. S. nebulatus

8 a Usually one postmental present (Fig. 761a);

dorsal bands cream to brown ........... ........ 9 Fig. 760. Lateral view of head in two species of
Sibon . Drawings: M. Vesely
b Usually two postmentals present (Fig.
761b); dorsal bands reddish brown to red
.................................................................. 10
9 a Ventrals 174-200; subcaudals 101-120 in
females, 113-126 in males; combined ven-
trals plus subcaudals 281-317 in females,
303-323 in males ......... Sibon dimidiatus
b Ventrals 168-172; subcaudals 95-100 in
females, 112 in single male; combined ven-
trals plus subcaudals 263-271 in females,
284 in single male ...... ..... Sibon miskitus
lOa Dorsal surface of head with green or yellow
as well as red and black markings; ground
color of body green .............. Sibon lamari a. S. dimidiatus b. S. annulatus
b Dorsal surface of head with pale and dark
brown markings; ground color of body usual- Fig. 761. Scalation of chin region (postmentals
ly olive to pale brown ...... Sibon annulatus orange).
280

Fig. 762. Sibon nebulatus (El Imposible
National Park, Ahuachapan, El Salvador).
Photo: G. Kohler
Fig. 763. Sibon nebulatus (Bartola, Rio San
Juan, Nicaragua). Photo: G. Kohler
Fig. 764. Sibon nebulatus (Volante, Panama).
Photo: S. Lotzkat
Further Reading
PETERS 1960, HENDERSON et al. 1977' KOFRON
1983, 1985, McCoy 1986, KOFRON 1990, SAVAGE
& MCDIARMID 1992, MCCRANIE et al. 2001
Colubridae
Siphlophis
The only species of the genus Siphlophis
that occurs in southern Central America is
S. cervinus. VILLA et al. (1988) erroneously
listed S. longicaudatus from Panama;
however, its distribution is restricted to
Brazil (J. PETERS & 0REJAS-MIRANDA 1970,
IBANEZ & SOLIS 1993). Siphlophis cervinus
is a tree-dweller in warm and wet lowland
rain forest and feeds on frogs, lizards, and
small mammals (COSTA PRUDENTE et al.
1998). The species reproduces oviparously
(PEREZ-SANTOS & MORENO 1988).
Siphlophis cervinus (LAURENTI 1768),
Synopsin Reptilium: 88; type locality:
America. TL to 645 mm. Panama to Bolivia as
well as on Trinidad.
Further Reading
HOGE 1964, COSTA PRUDENTE et al. 1998
Fig. 765. Siphlophis cervinus (Peru).
Photo: G. Kohler
281
 Colubridae

Fig. 766. Spilotes pullatus (Cerro Musfui, Fig. 767. Spilotes pullatus trying to swallow a
Nicaragua). Photo: J. Sunyer Mexican Porcupine (Volcan Mombacho,
Nicaragua). Photo: G. Kohler

Spilotes Spilotes pullatus (LINNAEUS 1758), Systema

Naturae, ed. 10: 225; type locality: "Asia" [in
error]. TL to 2600 mm. Tamaulipas and
The genus Spilotes consists of one widely- Oaxaca, Mexico, to Argentina, sea level to
distributed species, S. pullatus. This large 1500 m elevation in rain forest, wet forest
diurnal colubrid can be found both on the and dry forest. According to Wilson & MEYER
(1985), the division into subspecies suggested
ground and in the branches of large trees. by AMARAL ( 1929a) is insufficiently support-
This snake is extremely agile at moving ed, and I prefer not to recognize any subspe-
through vegetation, thus its Spanish cies, at least for Central America.
name, "voladora" (the flying one). It feeds
primarily on small mammals, birds, and
bird eggs (HENDERSON & HOEVERS 1977b,
LEE 1996). We saw a large S. pullatus at
the foot of the Mombacho Volcano
(Nicaragua), that was in the process of
swallowing a Mexican Porcupine (Sphig-
gerus mexicanus) (KOHLER & SEIPP 1999).
Spilotes pullatus reproduces oviparously,
producing clutches of 7-25 eggs, which
hatch in about two months (HIMMELSTEIN
1980, D. SCHMIDT 1990, CAMPBELL 1998a).

• Siphlophis cervinus
Spilotes pullatus

Further Reading
AMARAL 1929a
282
 Colubridae

Stenorrhina

Both species of this genus are distributed

widely across Mexico, Central America,
and northern South America. It is charac-
teristic of this genus to have the prenasal
scale fused with the internasal scale. These
are harmless, rear-fanged snakes, which
will not attempt to bite. As nocturnal
ground-dwellers, they live predominantly
in dry areas. They feed almost exclusively
on spiders and scorpions (DUELLMAN 1963,
SEXTON & HEATWOLE 1965, personal ob-
servation). Both species are oviparous and
produce clutches of 4-9 eggs during the dry
season (STUART 1935a, CENSKY & McCOY Fig. 768. Stenorrhina degenhardtii (Veracruz,
1988). Mexico). Photo: P. Heimes

Stenorrhina degenhardtii (BERTHOLD 1845),

Nachr. Georg-Augusts Univ. und K. Ges.
Wiss. Gottingen 3: 39; type locality: western
Colombia (see MYERS & BOHME 1996). TL to
800 mm. Southern Veracruz, Mexico, to
Ecuador and Venezuela, sea level to 1650 m
elevation in dry forest and wet forest.
According to STUART (1963) and WILSON &
MEYER (1985), the division into subspecies is
insufficiently supported, and I prefer not to
recognize any subspecies.
Stenorrhina freminvillei DUMERIL, BIBRON &
DUMERIL 1854, Erp. Gen. 7: 868; type locality:
Mexico. TL to 700 mm. Veracruz and
Guerrero, Mexico, to Costa Rica, sea level to
2200 m elevation in dry forest and wet forest.
According to STUART (1963) and WILSON &
MEYER (1985), the division into subspecies is
insufficiently supported, and I prefer not to
recognize any subspecies. Fig. 769. Stenorrhina degenhardtii (C6rdoba,
Colombia). Photo: M. Lundberg

Stenorrhina freminvillii Stenorrhina degenhardtii

283
 Colubridae

Storeria

The genus Storeria includes two species

that are distributed widely across North
America, with the species S. dekayi reach-
ing only to the northern portion of Central
America. These are small ground-dwelling
colubrids that feed on invertebrates such
as earthworms, slugs, and soft insects. At
the beginning of the rainy season, S. dekayi
gives birth to between 3 and 24 live young
(WRIGHT & WRIGHT 1957, CAMPBELL 1998a).

Storeria dekayi (HOLBROOK 1842), North

American Herpetology, Ed. 2, 4: 53; type lo-
cality: Massachusetts and New York. TL to
Fig. 770. Stenorrhina degenhardtii (Penas 450 mm. All across the eastern USA and
Blancas, Nicaragua). Photo: B. Aleman along the Caribbean side of Mexico, as well as
disjunct from Chiapas, Mexico, to Honduras,
100-1900 m elevation. In Central America,
the subspecies S. d. tropica COPE 1885 occurs.

Further Reading
TRAPrno 1944, ANDERSON 1961, CHRISTMAN 1982

Fig. 771. Stenorrhina freminvillei (Honduras).

Photo: G. Kohler

Key to Stenorrhina
1 a More than 159 ventrals; dorsum longitudi- Fig. 772. Storeria dekayi (Honduras, Dpto. de
nally striped or unicolor .................... ... .. . Intibuca). Photo: J. R. McCranie
..... ................. Stenorrhina freminvillei
b Fewer than 160 ventrals; dorsum with fine
spotting or unicolor .................................... .
. . . . . . . . . . . . ........ Stenorrhina degenhardtii

Further Reading
STUART 1963, WILSON & MEYER 1985

284
 Colubridae

Symphimus

Both species of this genus are diurnal and

semi-arboreal, inhabiting dry areas. The
species mayae was earlier included in the
genus Opheodrys (H. SMITH & TAYLOR
1945), until ROSSMAN & SCHAEFER (1974)
placed it in Symphimus . Symphimus
mayae is known to feed on insects (mostly
crickets and grasshoppers) (ROSSMAN &
SCHAEFER 1974). The species is oviparous
and produces clutches of 2-4 eggs during
the wet season (STAFFORD 2005). Fig. 773. Symphimus leucostomus (Oaxaca,
Mexico). Photo: W. E. Duellman
Symphimus leucostomus COPE 1870, Proc.
Arner. Philos. Soc. 11: 150; type locality:
Chihuitan, Oaxaca, Mexico. TL to 810 mm.
Pacific versant of Isthmus of Tehuantepec
(southeastern Oaxaca and southwestern
Chiapas, Mexico), 200-1000 rn elevation in
dry forest.
Symphimus mayae (GAIGE 1936), Carnegie
Inst. Washington Puhl. 457: 300; type locality:
Dzitas, Yucatan, Mexico. TL to 890 mm.
Northern portion of the Yucatan Peninsula,
Mexico, and central Belize, sea level to about
700 m elevation in dry forest.

Key to Symphimus
1 a Prenasal fused with adjacent internasal;
162-181 ventrals ... .................... .................. .
........... .............. Symphimus leucostomus
b Prenasal not fused with adjacent internasal;
150-165 ventrals ......... Symphimus mayae

Fig. 774. Symphimus mayae (Calakmul Bio-

sphere Reserve, Campeche, Mexico).
Photo: H. Bahena B.

Further Reading
JOHNSON 1973, ROSSMAN & SCHAEFER 1974

285
 Colubridae

Tantilla

This species contains 58 species, 23 of

which occur in Central America. These are
without exception small, secretive, harm-
less colubrids that will rarely attempt to
bite when they are handled. They are most
commonly found by turning over rotten
logs or rocks in the forest, as well as in
meadows and on coffee plantations. They
feed on invertebrates, such as beetle larvae
and centipedes (VAN DEVENDER & COLE
1977, LEE 1996, CAMPBELL 1998b). All spe-
cies are oviparous and produce clutches of
1-3 eggs (VAN DEVENDER & COLE 1977,
FITCH 1985).
Fig. 775. Tantilla alticola (Montana La Galia,
Jinotega, Nicaragua). Photo: G. Kohler
Tantilla albiceps BARBOUR 1925, Occ. Pap.
Boston Soc. Nat. Hist. 5: 156; type locality:
Barro Colorado Island, Gatun Lake, Panama.
TL to 212 mm. Barro Colorado Island,
Central Panama, around 100 m elevation.
Known only from the holotype.
Tantilla alticola (BOULENGER 1903), Ann.
Mag. Nat. Hist. (7) 12: 353; type locality:
Santa Rita, northern Medellin, Colombia. TL
to 327 mm. Central Nicaragua to north-
western Colombia, 90-2700 m elevation in
rain forest and cloud forest.
Tantilla armillata COPE 1876, J . Acad. Nat.
Sci. Philadelphia (2) 8: 143; type locality:
Middle Costa Rica. TL to 490 mm. On the
Pacific versant from Guatemala to the
Meseta Central Occidental of Costa Rica; also
on the Caribbean versant in Honduras; sea Fig. 776. Tantilla armillata (near Maritza,
level to 1430 m elevation in dry forest and Guanacaste, Costa Rica). Photo: M. Franzen
rain forest. Tantilla armillata was regarded
as a synonym of T. melanocephala by WILSON
& MENA (1980) but resurrected as a valid spe-
cies by SAVAGE (2002).
Tantilla bairdi STUART 1941c, Occ. Pap. Mus.
Zool. Univ. Michigan 452: 1; type locality: 2
km SE Finca Chichen, about 1550 m eleva-
tion, 10 airline km S Cohan on Chamelco
trail, Alta Verapaz, Guatemala. TL to 455.
Alta Verapaz and Baja Verapaz, Guatemala,
1500-1550 m elevation.
Tantilla breuicauda MERTENS 1952b, Zool.
Anz. 149: 137; type locality: El Grito, 1510 m
elevation, Los Angeles, La Libertad, El
Salvador. TL to 171 mm. South-central
Guatemala to northeastern El Salvador, 600-
1750 m elevation. Fig. 777. Tantilla bairdi (Baja Verapaz,
Guatemala). Photo: J. A. Campbell
286

Tantilla cuniculator SMITH 1939b, Field Mus.
Nat. Hist. Puhl. Zool. Ser. 24: 32; type locality:
Merida, Yucatan, Mexico. TL to 220 mm.
Northern Guatemala and the Yucatan
Peninsula, sea level to 50 m elevation.
Tantilla hendersoni STAFFORD 2004, Journal
of Herpetology 38: 44; type locality: 0.5 km
east of Las Cuevas on trail to Monkey Tail
River, Cayo District (GR 1643'95"N,
8859'17"W)". TL to 272 mm. Known only from
type locality.
Tantilla impensa CAMPBELL 1998b, Sci. Pap.
Nat. Hist. Mus. Univ. Kansas 7: 6; type locali-
ty: Aldea San Miguelito, 460 m elevation,
l5°22'N, 88°43'W, Sierra de Caral, Municipio
de Morales, Izabal, Guatemala. TL to 725
mm. Eastern Chiapas, Mexico, to western
Honduras, sea level to 1600 m elevation.
Fig. 778. Tantilla brevicauda (El Salvador).
Photo: T. Leenders
Fig. 779. Tantilla hendersoni (holotype).
Photo: P. Stafford
Colubridae
Tantilla Jani (GUNTHER 1895), Biologia
Centrali-Americana. Reptilia and Batrachia:
148; type locality: Guatemala. TL to 242 mm.
Pacific versant of Guatemala near the
Mexican border at around 1000 m elevation.
Tantilla cuesta WILSON 1982 is regarded to be
a synonym of T Jani according to CAMPBELL
(1998b).
Tantilla lempira WILSON & MENA 1980, Mem.
San Diego Soc. Nat. Hist. 11: 25; type locality:
41 km NW Tegucigalpa, Francisco Morazan,
Honduras. TL to 254 mm. South-central
Honduras, 1450 m elevation.
Tantilla moesta (GUNTHER 1863), Ann. Mag.
Nat. Hist. (3) 12: 352; type locality: El Peten,
Guatemala. TL to 592 mm. Northern
Guatemala, as well as the northern half of
the Yucatan Peninsula, sea level to 280 m.
Fig. 780. Tantilla impensa (near Quebrada
Grande, Copan, Honduras). Photo: G. Kohler
Fig. 781. Tantilla moesta (14 km SE Coba,
Quintana Roo, Mexico). Photo: J. C. Lee
287
 Tantilla reticulata COPE 1860c, Proc. Acad.
Nat. Sci. Philadelphia 12: 77; type locality:
Cocuyas de Veraguas, New Grenada
[= Cocuyas, Panama]. TL to 312 mm.
Southeastern Nicaragua to northwestern
Colombia, sea level to 1430 m elevation.
Tantilla rubra COPE 1876, J. Acad. Nat. Sci.
Philadelphia (2) 8: 144; type locality:
Tapanatepec, Oaxaca, Mexico. TL to 594 mm.
Central Nuevo Leon, Mexico, to western
Guatemala, 100-1000 m elevation.
Tantilla ruficeps (COPE 1894), Proc. Acad.
Nat. Sci. Philadelphia 46: 204; type locality:
Cocuyas de Veragua, "New Grenada". TL to
500 mm. Nicaragua to Panama and possibly
northwestern Colombia; sea level to 1600 m
Fig. 782. Tantilla reticulata (La Selva, Costa elevation in moist forest and rain forest.
Rica). Photo: D. M. Dehling Tantilla ruficeps was regarded as a synonym

Fig. 783. Tantilla reticulata (Costa Rica). Fig. 785. Tantilla supracincta (Comadre, (Limon,
Photo: T. Leenders Costa Rica). Photo: R. W. Van Devender

T. armillata
• T. ruficeps
• T. moesta
• T.jani
.a. T. impensa
"""~~~Y T. lempira

Fig. 784. Tantilla rubra (Tuxtla Gutierrez, Tantilla

Chiapas, Mexico). Photo: A. Ramirez V.
288
 • T. alticola of T. melanocephala by WILSON & MENA
• T. brevicauda (1980) but resurrected as a valid species by
T. cuniculator SAVAGE (2002).
... T. a/biceps
..a.. T. bairdi
Tantilla schistosa (BocoURT 1883), Miss. Sci .
Mex., Rept.: 584; type locality: Alta Verapaz,
Guatemala, and Mexico. TL to 293 mm.
Veracruz and Oaxaca, Mexico, to central
Panama, sea level to 1600 m elevation.
Tantilla supracincta (PETERS 1863c)
Monatsber. Kong. Akad. Wiss. Berlin 1863:
272; type locality: Guayaquil, Ecuador. TL to
Tantilla 562 mm. Southeastern Nicaragua to Ecuador,
sea level to 850 m elevation. Tantilla annula-
ta BOETTGER 1892 is regarded to be a syno-
nym of T. supracincta according to WILSON
T. supracincta (1987c).
• T. vulcani
... T. tecta Tantilla taeniata (BOCOURT 1883), Miss. Sci.
Mex., Rept.: 587; type locality: Guatemala. TL
to 415 mm. Antigua Basin and the south-
eastern highlands of Guatemala across El
Salvador and Honduras as far south as
northern Nicaragua, sea level to 1550 m
elevation.
Tantilla tayrae WILSON 1983, J. Herpetol. 17:
54; type locality: Finca San Jeronimo, 7.5 km
Tantilla N (by road) Cacaohatan [= Cacahoatan or
Cacahuatan], 760 m elevation, Volcan
Tacana, Municipio de Union Juarez, Chiapas,
Mexico. TL to 317 mm . Southeastern
Chiapas, Mexico, 760-960 m elevation.
• T. reticulata
• T. rubra Tantilla tecta CAMPBELL & SMITH 1997,
T. taeniata Herpetologica 54: 333; type locality: slope
.& T. tritaeniata flanking the NE side of Laguna Yaxha, 220 m
"f' T. hendersoni elevation l 7°03'43"N 89°23'12"W Peten·
Guatemaia. TL to 222 'mm. Northea'stern Ei
Peten, Guatemala, 220 m elevation in dry
forest.
Tantilla triseriata SMITH & SMITH 1951, Proc.
Biol. Soc. Washington 64: 97; type locality:
Coatlan, Oaxaca, Mexico. TL to 203. South-
Tantilla central Oaxaca, Mexico.
Tantilla tritaeniata SMITH & WILLIAMS 1966,
Southwest. Nat. 11: 483; type locality: Isla de
Guanaja, Islas de la Bahia, Honduras. TL to
T. schistosa 273 mm. Isla de Guanaja, Honduras.
• T. vermiformis Tantilla vermiformis (HALLOWELL 1860), Proc .
... T. tayrae
..a.. T. triseriata Acad. Nat. Sci. Philadelphia 12: 484; type
locality: Nicaragua. TL to 157 mm. Pacific
side of El Salvador to northwestern Costa
Rica, sea level to 520 m elevation.
Tantilla vulcani CAMPBELL 1998, Sci. Pap.
Nat. Hist. Mus. Univ. Kansas 7: 11; type lo-
cality: Finca El Carmen, 518 m elevation, km
197.5 on CA-2, Quetzaltenango, Guatemala.
TL to 246 mm. Pacific side of Guatemala.
Tantilla

289
 Colubridae

Key to Tantilla
1 a Dorsum with pale dark-bordered cross
bands (Fig. 785c) ... Tantilla supracincta
b Dorsal coloration different than that above
······································· ······· ··· ·· ·· ··· ······· · 2
2 a Dorsal surface of head and first one or two
dorsals white to cream-colored with indis-
tinct dark shading along the frontal-pre-
frontal sutures; dorsum unicolor dark olive
brown; one postocular .. Tantilla albiceps
b Head and/or dorsal coloration different
than above; usually two postoculars ........ 3
3 a Dorsum and venter uniformly dark brown
to black; conspicuous bright yellow or whit-
Fig. 786. Tantilla taeniata (Miraflor, Estell, ish head band present (covers parietals
Nicaragua). Photo: M. Jansen and the first 2-7 dorsal rows) ........ ............. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . ... ... .. Tantilla moesta
b Venter paler in color than dorsum; pale
head band present or not ....... ................. 4
4 a Dark vertebral stripe present ............... ... 5
b No dark vertebral stripe present .... ......... 8
5 a Fewer than 30 subcaudals ................... .. ......
. . . . . . . . . . . . . . . . . . . . . ...... Tantilla vermiformis
b More than 30 subcaudals .................. ....... 6
6 a Fewer than 155 ventrals; pale neck spot
indistinct and predominantly on the scales
behind the parietals; pale lateral stripes
with lower dark border Tantilla lempira
b More than 154 ventrals, or if fewer than
155 ventrals, pale neck spot distinct and
predominantly on the parietals; pale la-
Fig. 787. Tantilla tritaeniata (Isla de Guanaja, teral stripe, if present, without lower dark
Honduras). Photo: J . R. McCranie border ................................................ ........ 7
7 a Ventrals 155-174 (males), 158-177 (fe-
males); subcaudals 46-59 (males), 48-60
(females); dorsum of head usually dark
brown or black with a distinct small pale
spot on each parietal Tantilla armillata
b Ventrals 139-149 (males), 143-155 (fe-
males); subcaudals 64-83 (males), 59-75
(females); dorsum of head usually pale
brown with large indistinct pale spots on
parietals .... ................... Tantilla ruficeps
8 a Dorsum without pale longitudinal stripes
.... ................................................................ 9
b Dorsum with pale longitudinal stripes
.................................... ....................... ....... 13
9 a Snout with pale coloration; pale postocular
spot present ................. Tantilla alticola
b Snout color similar to rest of dorsal head
surface, or only slightly paler; without pale
postocular spot ......... .............................. 10
Fig. 788. Tantilla uermiformis (Guanacaste, lOa More than 52 subcaudals Tantilla rubra
Costa Rica). Photo: R. W. Van Devender
290

b Fewer than 52 subcaudals ...................... 11
Ila More than 155 ventrals .. Tantilla bairdi
b Fewer than 155 ventrals ..... .................... 12
12a More than 43 subcaudals; 140-154 ventrals
................... ...... ... .... ....... Tantilla tayrae
b Fewer than 43 subcaudals; 117-147 ven-
trals ......... ... ... ........... Tantilla schistosa
13a Fewer than 30 subcaudals ........ ............... .
.. ............................ .. Tantilla brevicauda
b More than 30 subcaudals .. ........ ........... .. 14
14a No pale vertebral stripe present .. .... .. .. .. ...
......... ..... Tantilla cuniculator (in part)
b Pale vertebral stripe present ........ .. ....... 15
15a Edges of ventral scales darkly pigmented
or with heavy dark spotting .... ............... 16
b Edges of ventral scales unpigmented or
only with isolated spots .......................... 20
16a More than 157 ventrals; 58-70 subcaudals
... .. ....... ......... ... ......... Tantilla reticulata
b Fewer than 157 ventrals; 37-64 subcaudals
........ .................................................... ...... 17
l 7a Pale neck band reduced to two neck spots
............ ... .. .. ..... ....... ..... .... Tantilla jani
b Pale neck band complete ........................ 18
18a More 60 subcaudals; no pale spot on snout
................................. Tantilla hendersoni
b Fewer than 60 subcaudals; a pale spot on
snout .............................................. .......... 19
a. T. reticulata b. T. taeniata
c. T. supracincta d. T. schistosa
Fig. 789. Dorsal coloration in Tantilla.
Colubridae
19a Pale vertebral stripes continuous at least
on anterior half of body; pale lateral stripes
reach as far as tail ............. Tantilla tee ta
b Vertebral stripe reduced to a series of pale
spots or almost completely absent; pale
lateral stripes fade on posterior half of
body ...... .... ....... .......... Tantilla vulcani
20a Pale longitudinal stripe only indistinct;
fewer than 59 subcaudals .......... .................
.......... ..... Tantilla cuniculator (in part)
b Pale longitudinal stripe distinct; more
than 59 subcaudals ....................... .......... 21
2la Ground color so dark that dark border of
pale longitudinal stripe not distinguish-
able; first dorsal row (paraventrals) free of
dark pigmentation, at least on anterior
halfof body ................ Tantilla triseriata
b Pale longitudinal stripe distinctly bordered
by a dark line; upper half of first dorsal
row (paraventrals) darkly pigmented .... 22
22a 162-171 ventrals; pale vertebral stripe
restricted to vertebral row; pale lateral
stripes (yellow in life) very distinct over
entire length of body; pale neck band com-
plete or interrupted dorsally ....... ............ .. .
... ... .... ........ ............... Tantilla impensa
b 143-161 ventrals; pale vertebral stripe
covers vertebral row and adjacent third of
paravertebral rows; pale lateral stripes
less distinct; pale neck band complete ... 23
23a Pale vertebral stripe begins three scales
behind the parietals and is not narrower at
the beginning than posteriorly; pale later;d
stripe begins three scales behind the sth
supralabial an9 covert the adjacent two
thirds of the 3r and 4t dorsal rows ......... .
.... .. . . . . . . .. . . . . ............ Tantilla tritaeniata
b Pale vertebral stripe begins five scales
behind the parietals and is narrower at the
beginning than posteriorly; pale lateral
stRpe begins five or six scales behind the
6t supralabial and covers Qnly the adja-
cent halves of the 3rd and 4th dorsal rows
..... ........ .... ...... ........ .. Tantilla taeniata
Further Reading
H . SMITH 1942c, HARDY & COLE 1967, WILSON &
MEYER 1971, WILSON & VILLA 1973, VAN
DEVENDER & COLE 1977, WILSON & MENA 1980,
WILSON 1982a, b, 1985a-e, 1986, 1987a-c, 1988a,
1990a-c, 1992, CAMPBELL & SMITH 1997,
CAMPBELL 1998b, H. SMITH et al. 1998, WILSON
1999, WILSON & MCCRANIE 1999, DIXON et al.
2000, SAVAGE 2002
291
 Colubridae

Tantillita
Three very small, secretive colubrid spe-
cies, which are very similar to the Tantilla
species, are included in the genus
Tantillita. They usually stay on the
ground, under rotten logs or in soft plant
material. In northeastern Nicaragua (in
the vicinity of Pueblo Wiso), I startled a
specimen of Tantillita lintoni in the wet
leaves on the forest floor during the day
(KOHLER 1999d). These small snakes are
probably nocturnal, feed on small inverte-
brates (termites?) and are oviparous (LEE Fig. 790. Tantillita canula (El Peten,
1996, CAMPBELL 1998a). Guatemala). Phot.o: J. A Campbell

Tantillita brevissima (TAYLOR 1937b), Trans.

Kansas Acad. Sci. 39: 344; type locality:
Tonala, Chiapas, Mexico. TL to 150 mm.
Pacific side of southeastern Oaxaca, Mexico,
to south central Guatemala, 200-1700 m ele-
vation. Tantilla excubitor WILSON 1982a is
regarded to be a synonym of Tantillita brevis-
sima according to WILSON' (1988e).
Tantillita canula (COPE 1876), J. Acad. Nat.
Sci. Philadelphia (2) 8: 144; type locality:
Yucatan, Mexico. TL to 175 mm. Yucatan
Peninsula, sea level to 200 m elevation.
Tantillita lintoni (SMITH 1940), Proc. Biol.
Soc. Wash. 53: 61; type locality: Piedras
Negras, El Peten, Guatemala. TL to 200 mm.
Veracruz, Mexico, to northern Nicaragua, sea
level to 400 m elevation. In Central America, Fig. 791. Tantillita lintoni (Las Cuevas,
only the nominate subspecies occurs. Belize). Phot.o: P. Stafford

Key to Tantillita
1 a 116-126 ventrals; 28-34 subcaudals; pale
neck spot present Tantillita brevissima Tantillita canula
b 103-115 ventrals; 32-56 subcaudals; with- ..,. Tantillita brevissima
out pale neck spot ............... .................. .. 2 • Tantillita /intoni
2 a 32-44 subcaudals; pale vertebral stripe
present; pale areas present on dorsal sur-
face of head .................. Tantillita canula
b 43-56 subcaudals; no pale vertebral stripe;
dorsal surface of head unicolor brown
......... ............ ... .......... Tantillita lintoni

Further Reading
WILSON 1988b, d

292
 Colubridae

Thamnophis

There are approximately 30 species of gar-

ter snakes distributed predominantly
across North America, with only four spe-
cies occurring in Central America. The
nominate form T. sumichrasti cerebrosus,
described by H . SMITH (1942b) on the basis
of three specimens from "Esquintla,
Guatemala", cannot be assigned to any of
the known Central American species. Since Fig. 792. Thamnophis marcianus (Quintana
no further specimens have been found, this Roo, Mexico). Photo: J. R. McCranie
is currently regarded as a nomen dubium
(name of dubious merit) (ROSSMAN 1991).

Garter snakes are semiaquatic, harmless

snakes, that will expel a vile-smelling
secretion from the cloaca as their first line
of defense before they will bite. They live in
ponds, lakes, swamps, streams, and rivers
and feed on tadpoles, frogs, fish, and earth-
worms (ROSSMAN et al. 1996). They are pre-
dominantly diurnal, but are occasionally
also seen at night (CAMPBELL 1998a). All Fig. 793. Thamnophis proximus (Palo Verde,
species are livebearers, producing litters of Costa Rica). Photo: D. M. Dehling
6-22 young (WRIGHT & WRIGHT 1957'
ROSSMAN et al. 1996).
Thamnophis proximus (SAY 1823), in JAMES:
Exped. Pittburgh to Rocky Mts. 1: 339; type
locality: approximately 3 mi ENE Fort
Thamnophis cyrtopsis CKENNICO'IT 1860), Calhoun, Washington County, Nebraska,
Proc. Acad. Nat. Sci. Philadelphia 12: 333;
USA. TL to 900 mm. Indiana and Kansas,
type locality: Rinconada, Coahuila, Mexico. USA, to Costa Rica, sea level to 2500 m ele-
TL to 1145 mm. Utah and Colorado, USA, to vation. In Central America, the subspecies
Guatemala, sea level to 2700 m elevation. In Thamnophis p. alpinus ROSSMAN 1963 (cen-
Central America, only the subspecies tral Chiapas, Mexico) and Thamnophis p.
Thamnophis c. collaris (JAN 1863) occurs. rutiloris (COPE 1885) (Guerrero and southern
Thamnophis fulvus (BOCOURT 1893), Miss. Tamaulipas, Mexico, to Costa Rica, except
Sci. Mex., Rept.: 777; type locality: Alta central Chiapas, Mexico) occur.
Verapaz, Guatemala. TL to 652 mm. Central
Chiapas, Mexico, to southwestern Honduras
and adjacent El Salvador, 1400-3350 m eleva-
tion.
Thamnophis marcianus (BAIRD & GIRARD
1853), Cat. North Amer. Rept. : 36; type lo-
cality: Red River, Arkansas. TL to 1090 mm.
Kansas and California, USA to northern
Costa Rica, sea level to 1640 m elevation. In
Central America, the subspecies Thamnophis
m. bovalli DUNN 1940b (Nicaragua and north-
ern Costa Rica) and Thamnophis m. prae-
ocularis BocOURT 1892 (populations in the
Yucatan Peninsula, Guatemala, and "" Thamnophis marcianus
Honduras) occur. Thamnophis cyrtopsis

293
 Colubridae

Key to Thamnophis
1 a Pale longitudinal stripe on the side covers
3rd and 4th dorsal rows, at least in anter-
ior body area; without dark spots on the
supralabials; no pale neck spots .............. .
... . . . . . . . . . . . . . . . ...... .. Thamnophis proximus
b Pale longitudinal stripe on si~, if present,
covers 2nd and 3rd or just 3r dorsal row,
at least in anterior body area; dark spots
on the supralabials present; with pale neck
spots ............. .............................. .............. . 2
2 a Pale longitudinal stripe on side, if present,
covers only 3rd dorsal row, at least in
anterior body area; posterior supralabials
distinctly darker colored (similar to tem-
porals) than anterior supralabials ............ .
. . . . . . . . . . . . . . .. ..... .. Thamnophis marcianus
b Pale longitudinal stripe on side, if present, Abb. 795. Thamnophis cyrtopsis (Santa Rosa,
covers 2nd and 3rd dorsal row, at least in Guatemala). Photo: L. Melendez
anterior body area; all supralabials simi-
larly colored ......................................... 3
3 a No dark dorsal neck spot present; pale ver-
tebral stripe indistinct or absent, at least
in posterior body area; tongue completely
black; 56-80 subcaudals ......................... .
. . . . . . ........... .... ....... .. ... Thamnophis fulvus
b One broad dark dorsal neck spot present;
pale vertebral stripes distinct as far as tail
tip; tongue red with black tips; 75-101
subcaudals .......... Thamnophis cyrtopsis

Further Reading
SHREVE & GANS 1958, ROSSMAN 1970, 1971,
WEBB 1980, 1982, ROSSMAN et al. 1996

Fig. 796. Thamnophis fuluus (La Tigra,

Honduras). Photo: M. Lundberg

.., Thamnophis fulvus

Fig. 794. Thamnophis proximus (Guanacaste Thamnophis proximus
National Park, Costa Rica). Photo: M. Franzen
294
 Colubridae

Tretanorhinus

Two of the three aquatic species belonging

to the genus Tretanorhinus occur in
Central America. On the Bay Islands
(Honduras), there are specimens of T.
nigroluteus with gray-brown underbellies
as well as some that have brilliant orange
coloration on their ventral shields. These
colubrids are rarely aggressive and live in
ponds, lakes, and swamps, as well as
muddy streams and rivers with slow flow-
ing water; only rarely will they leave their Fig. 797. Tretanorhinus nigroluteus (Isla de
aquatic homes. At dusk, they become active Guanaja, Honduras). Photo: G. Kohler
and can be observed eating small fishes
and tadpoles. They will not, however, eat
the freshwater shrimp, which are often
abundant in their aquatic habitat (WILSON
& HAHN 1973). All Tretanorhinus species
reproduce oviparously, producing clutches
of 6-9 eggs (CAMPBELL 1998a).

Tretanorhinus nigroluteus COPE 1861e, Proc.

Acad. Nat. Sci. Philadelphia 13: 298; type
locality: San Juan del Norte, Nicaragua [in
error, actually "Aspinwall, Panama," accord-
ing to PETERS & 0REJAS-MIRANDA (1970)]. TL
to 765 mm. Southern Veracruz, Mexico, to
Panama, sea level to 750 m elevation.
According to WILSON & MEYER (1985) the
division into subspecies promulgated by
VILLA (1969a) is insufficiently supported, and
I prefer not to recognize any subspecies. Fig. 798. Tretanorhinus nigroluteus (Bosawas,
Nicaragua). Photo: J. Sunyer
Tretanorhinus mocquardi BocOURT 1891, Le
Naturaliste (2) 101: 122; type locality: "a
Panama"[= Panama City according to DUNN
1939] . TL to 730 mm. Central and eastern
Panama.

Key to Tretanorhinus
1 a Dorsum with three longitudinal stripes;
more than 165 ventrals ... .............. .. ....... . .
.... . . . . .. . ......... Tretanorhinus mocquardi
b Dorsum with two longitudinal rows of dark
spots; fewer than 155 ventrals ............. .. .
.... . . . . . . ......... Tretanorhinus nigroluteus

Further Reading
DUNN 1939a, VILLA 1969a
 Colubridae

Trimetopon

The species of the genus Trimetopon are

small, gracile colubrids, about which little
is known. Certain species which had ear-
lier been included in this genus (e.g., hann-
steini, pilonaorum and posadasi; see J.
PETERS & OREJAS-MIRANDA 1970) have
been placed in Rhadinaea (MYERS 1974).
The species of Trimetopon are known to
feed on salamanders and lizards (SAVAGE
2002). At least Trimetopon slevini is
oviparous and produces clutches of 1-2
eggs (DUNN 1940a). Abb. 799. Trimetopon barbouri (holotype, MCZ
23877). Photo: Museum of Comparative
Trimetopon barbouri DUNN 1930a, Occ. Pap. Zoology, Harvard University. Scale bar = 5 mm
Boston Soc. Nat. Hist. 5: 156; type locality:
Pedro Miguel, Canal Zone, Panama. TL to 260
mm. Central Panama.
Trimetopon gracile (GONTHER 1872), Ann.
Mag. Nat. Hist. (4) 9: 18; type locality: near
Cartago, Costa Rica. TL to 300 mm. Costa
Rica.
Trimetopon pliolepis COPE 1894, Proc. Acad.
Nat. Sci. Philadelphia 46: 201; type locality:
San Jose, Costa Rica. TL to 287 mm. Costa
Rica and Panama.
Trimetopon simile DUNN 1930a, Occ. Pap.
Boston Soc. Nat. Hist. 5: 331; type locality:
Reventaz6n, Costa Rica. TL to 155 mm.
Northeastern Costa Rica.
Trimetopon sleuini DUNN 1940a, Proc. Acad. Abb. 800. Trimetopon simile (holotype, MCZ
Nat. Sci. Philadelphia 92: 117; type locality: 15263). Photo: Museum of Comparative
near Boquete, Chiriqui, Panama. TL to 275 Zoology, Harvard University. Scale bar= 5 mm
mm. Central-eastern Costa Rica and western
Panama.
Trimetopon uiquezi DUNN 1937a, Copeia
1937: 215; type locality: Siquirres, Costa Rica.
TL to 250 mm. Northeastern Costa Rica.

~
Trimetopon slevini
• Trimetopon barbouri
• Trimetopon gracile
~ & Trimetopon pliolepis

(
~ --- - '" ""' Trimetopon simile
1.·~ .(,,.. • Trimetopon viquezi

-:::J~h'it;f~ -~--
\'<Sl~~-
... _/\-1':? ~~~;
. Q '
Abb. 801. Trimetopon barbouri (holotype, MCZ
\j\ ) ' 23877). Photo: Museum of Comparative
Cl"'\. J
Zoology, Harvard University.
296
 Colubridae

Key to Trimetopon
1 a 15 dorsal rows at midbody ........................ 2
b 17 dorsal rows at midbody ........... ............. 4
2 a Two separate prefrontals present ............. .
............... ............ .... Trimetopon barbouri
b Prefrontals fused into a single shield ...... 3
3 a More than 135 ventrals; one postocular;
dorsum with indistinct dark vertebral
stripe; pale nuchal collar, if present, not
continuous with throat color .................... .
........................... ........ Trimetopon gracile
b Fewer than 130 ventrals; two postoculars;
dorsum uniform; pale nuchal collar pre-
sent and continuous with throat color ..... . . Fig. 803. Trimetopon gracile (Monteverde,
...... ..... .. ........ ........... Trimetopon simile Costa Rica). Photo: M. & P. Fogden
4 a Prefrontals fused into a single shield; a
pale nuchal collar present, sometimes bro-
ken by a vertebral dark stripe and contin-
uous with throat color ...... ....... ............ .
.. .............................. Trimetopon pliolepis
b Two separate prefrontals present; no pale
nuchal collar, although a pair of pale neck
spots can be present .......... ............... ..... .. 5
5 a More than 40 subcaudals; a pair of pale
neck spots; body with a vertebral dark
stripe and 2 or 3 lateral dark stripes ....... .
.. . . . . . . . . . . . . . . .. . . . . . . . . . .... Trimetopon slevini
b Fewer than 40 subcaudals; without pale
neck spots; body with a vertebral dark
stripe and a single late~al dark st~pes ··;
.......... ... ... ..... ......... Trimetopon viquezi

Further Reading
DUNN 1930a, 1940a, TAYLOR 1951, 1954 Fig. 804. Trimetopon pliolepis (Rio Sereno,
Chiriqui, Panama). Photos: G. Kohler

Abb. 802. Trimetopon simile (holotype, MCZ

15263). Photo: Museum of Comparative Fig. 805. Trimetopon slevini (Los Cusingos,
Zoology, Harvard University. Puntarenas, Costa Rica). Photos: G. Kohler
297
 Colubridae

Trimorphodon

According to the most recent review of the

Trimorphodon biscutatus species complex
by DEVI'IT et al. (2008), the former subspe-
cies of T. biscutatus have been elevated to
species level. Therfore, we recognize two
species of this genus in Central America, T.
biscutatus and T. quadruplex. Lyre snakes
are nocturnal, semi-arboreal colubrids that
are distributed all across the dry regions of
the southwestern U.S., as well as through
Mexico and Central America (excluding
the Yucatan Peninsula). They feed on
lizards, small mammals and birds, with
spiny-tailed iguanas (Ctenosaura) being
their preferred prey in some areas.
Swallowing a large spiny-tailed iguana,
however, can prove life threatening for
these snakes, as described by RAM1REZ-
BAUTISTA & URIBE (1992). They observed a
175 cm long lyre snake (weight 1183 g) in
Mexico as it seized a large male
Ctenosaura pectinata, strangled it, and
swallowed the lizard headfirst. After wrig-
gling around restlessly in the same spot for
about 20 minutes, the snake suddenly
stretched, flipped onto its back and died.
Necropsy revealed that the gullet and sto-
mach of the snake had been slit open by
the sharp tail spines of the iguana.
Trimorphodon biscutatus reproduces ovi-
parously during the dry season; clutches of
7-21 eggs will hatch after an incubation
period of79-106 days (FITCH 1970, KOHLER
1997).
Fig 807. Trimorphodon biscutatus (near
Mixtequilla, Oaxaca, Mexico). Photo: G. Kohler
Trimorphodon biscutatus (DUMERIL, BIBRON
& DuMERIL 1854), Erp. Gen. 7: 1153; type
locality: "Mexique". TL to 1750 mm. Jalisco,
Mexico to southwestern Guatemala, sea level
to 1590 m elevation.
Trimorphodon quadruplex (SMITH 194lb),
Proc. U.S. Natl. Mus. 91 (3130): 157; type loca-
lity: Estell, Nicaragua. TL to 1750 mm.
Southeastern Guatemala to Costa Rica, sea
level to 1595 m elevation.
Key to Trimorphodon
1 a Primary dark blotches divided by central
light band into undivided secondary blot-
ches ........ ..... . Trimorphodon biscutatus
b Primary dark blotches divided by central
light band into secondary blotches, most of
which are further subdivided by central
light markings .............................................
..................... Trimorphodon quadruplex
Further Reading
H. SMITH 194lb, GEHLBACH 1971, SCOTT &
MCDIARMID 1984, DEVITT et al. 2008

• Trimorphodon quadruplex
Abb. 806. Trimorphodon quadruplex (Morgans Trimorphodon biscutatus
Rock, Nicaragua). Photo: J. Sunyer
298
 Colubridae

Abb. 808. Trimorphodon quadruplex

(Apoyeque, Nicaragua). Photo: A. Z. Quintana

Tripanurgos

ZAHER & PRUDENTE (1999, 2003) synonym-

ized Tripanurgos with Siphlophis based on
ZAHER's unpublished PhD. However, until
the evidence for this action has not been
published, I prefer to recognize Tripanur-
gos as a valid genus. Tripanurgos com-
pressus is a nocturnal colubrid that feeds
predominantly on frogs and lizards. They
live primarily in the branches of trees
(GREENE 1997). This species is seldom seen,
therefore hardly anything is known about
its ecology. Shortly after nightfall, E. LEHR
(pers. comm. 2000) encountered an adult T.
compressus, crossing a path in the lowland
rain forest (230 m elevation) in central
eastern Peru. The animal seemed entirely Abb. 809. Tripanurgos compressus (Drake Bay,
harmless and made no attempt to bite. Costa Rica). Photo: J. Kohler
RILEY & WINCH (1985) reported on eggs of
T. compressus, which they had discovered
in a nest of leaf-cutter ants on Trinidad.
Because two distinct size classes were pre-
sent among the eggs, it was suggested that
two or more females oviposited in the same
nest.

Tripanurgos compressus (DAUDIN 1803), Hist.

Nat. Rept. 6: 247; type locality: Surinam. TL
to 750 mm. Southeastern Costa Rica to Peru
and Bolivia, sea level to 1000 m elevation (in
South America to 1600 m elevation). Further Reading
RILEY & WINCH 1985, ZAHER 1999
299
 Colubridae
Fig. 810. Tropidodipsas sartorii (Campeche, Mexico).
Tropidodipsas
The species of the genus Tropidodipsas
are closely related to the Sibon species and
were earlier included in the latter genus by
KOFRON 1985. WALLACH (1995) showed that
Tropidodipsas was distinguishable as a
separate genus from Sibon. Like the spe-
cies of the genera Dipsas and Sibon, the
Tropidodipsas species feed primarily on
snails and slugs (DUNDEE et al. 1986, LEE
1996). All Tropidodipsas species are noc-
turnal, mostly terrestrial. They reproduce
oviparously, laying 3-6 eggs per clutch
towards the end of the dry season or at the
beginning of the rainy season (KOFRON
1987, CAMPBELL 1998a).
300
Photo: J . C. Lee
Tropidodipsas fasciatus (GONTHER 1858),
Cat. Snakes Brit. Mus.: 181; type locality:
Mexico. TL to 717 mm. Disjunct from
Guerrero and Veracruz, Mexico, as far as
Chiapas, Mexico, and in the north of the
Yucatan Peninsula, sea level to 2000 m eleva-
tion in rain forest. In Central America, only
the nominate subspecies occurs.
Tropidodipsas fischeri BoULENGER 1894a,
Cat. Snakes Brit. Mus. 2: 296; type locality:
Guatemala. TL to 652 mm. Western Oaxaca,
Mexico, to western El Salvador and south-
western Honduras, 1350-3000 m elevation in
pine-oak forest and cloud forest. Two subspe-
cies are recognized: Tropidodipsas f fischeri
(same distribution as the species except for
Alta Verapaz, Guatemala) and Tropidodipsas
f kidderi STUART 1942c (Alta Verapaz,
Guatemala).
Tropidodipsas sartorii COPE 1863, Proc. Acad.
Nat. Sci. Philadelphia 15: 100; type locality:
Mirador, Veracruz, Mexico. TL to 857 mm.
Nuevo Leon, Mexico, to Nicaragua, sea level
to approximately 2000 m elevation in dry
forest and wet forest. In Central America,
only the nominate subspecies occurs.
 Colubridae

Fig. 811 Tropidodipsas sartorii (Huejutla, Fig. 813. Tropidodipsas fasciatus (Calakmul,
Hidalgo, Mexico). Photo: G. Kohler Campeche, Mexico). Photo: R. Cedeno V.

Key to Tropidodipsas
1 a Dorsal scales completely smooth .............. .
. . . .. . . ... .............. Tropidodipsas fasciatus
b Dorsal scales weakly keeled ..................... 2
2 a Dorsum with 12-30 black rings, that reach
as far as venter; spaces between the dark
bands cream colored or orange red in life;
white in preservative (scale tips may be
black) .... ... ... ...... Tropidodipsas sartorii
b Dorsum with 20-64 dark brown bands that
do not reach to venter; spaces between
dark bands pale brown to gray-brown ....... .
. . . . . . . . . . . . . . . ....... ... Tropidodipsas fischeri

Fig. 812. Tropidodipsas fischeri (Guatemala). Further Reading

Photo: L. Melendez KOFRON 1985, 1987, 1988b, WALLACH 1995

Tropidodipsas sartorii

Tropidodipsas fasciatus
111 Tropidodipsas fischeri

301
 Colubridae

Urotheca

The genus name Urotheca was revived at

the end of the 1980's for the species of the
Rhadinaea lateristriga group sensu MYERS
1974 (SAVAGE & CROTHER 1989). Char-
acteristically, these colubrids have a con-
spicuously long tail capable of autotomy,
unique hemipene morphology, and a white
line along the first dorsal rows that is
generally quite distinct. The tail will break
off easily if it is seized. In fact, most
specimens of this genus that have been
found have incomplete tails, which sup- Fig. 814. Urotheca guentheri (Quebrada
ports the assumption that the breakable Machin, 540 m elevation, Colon, Honduras).
Photo: J. M. McCranie
tail is a survival strategy to protect against
predators (MYERS 1974, SAVAGE & CROTHER
1989). Most of the species of this genus are
thought to be diurnal, although Urotheca
fuluiceps has been seen out in the open at
night (MYERS 1974). All Urotheca species Urotheca decipiens (GONTHER 1893), Biologia
Centrali-Americana. Reptilia and Batrachia:
are ground-dwellers that feed on salaman- 105; type locality: Irazu, Costa Rica. TL to 569
ders, frogs, and lizards. They reproduce mm. Northern Costa Rica to Colombia,
oviparously (MYERS 1974). recently recorded in northeastern Honduras
(MCCRANIE et al. 2003), sea level to about
1600 m elevation (in Colombia also at higher
elevations) in rain forest.

Urotheca decipiens Urotheca fulviceps COPE 1886a, Proc. Amer.

.a. Urotheca fulviceps Phil. Soc. 23: 279; type locality: Panama. TL
• Urotheca myersi to 489 mm. Costa Rica to northwestern
Ecuador, sea level to about 500 m elevation in
rain forest.
Urotheca guentheri (DUNN 1938b), Copeia
1938: 198; type locality: Irazu, Costa Rica. TL
to 540 mm. Northeastern Honduras to cen-
tral Panama, sea level to about 2100 m eleva-
tion in rain forest.
Urotheca myersi SAVAGE & LAHANAS 1989,
Copeia 1989: 892; type locality: near Finca de
Jardin, 2255 m, approx. 9.6 km S by road of
Villa Mills, just south of Division, on the
Carretera Interamericana (Ruta 2), Canton
= Urotheca guentheri
Urotheca pachyura Perez-Zeledon, Cordillera de Talamanca, San
Jose, Costa Rica. TL to 349 mm. Pacific ver-
sant of Cordillera de Talamanca in central
and southern Costa Rica, 1500-2255 m eleva-
tion in cloud forest.
Urotheca pachyura (COPE 1876), J . Acad. Nat.
Sci. Philadelphia (2) 8: 145; type locality:
Sipurio [near Suretka, Limon, according to
MYERS 1974], Costa Rica. TL to 725 mm.
Central Costa Rica to western Panama, sea
level to about 1600 m elevation in rain forest.

302
 Colubridae

Fig. 815. Urotheca decipiens (north slope of Cerro Pando, 1620 m elevation, Bocas del Toro,
Panama). Photo: C. W. Myers

Key to Urotheca
1 a Dorsal surface of head pale yellowish
brown, in sharp contrast to the darker
body coloration ................. ...... .. ............... 2
b Coloration of dorsal surface of head similar
to that of body ... .............. .......................... 4
2 a Pale head coloration reaches 3-4 dorsal
scales past the parietals; lip shields inten-
sively dark spotted ... Urotheca fulviceps
b Pale head coloration reaches 1-2 dorsal
scales past the parietals; lip shields, if at
all, only with indistinct spots .. ................. 3
3 a One narrow, dark neck band present
......... .. .......... .. ............... Urotheca myersi
b Without dark neck band ...... ............... ........ .
... . . .. . . .. ...................... Urotheca pachyura
Fig. 816. Urotheca guentheri (Rio San Juan,
4 a Without white ocelli at side of back of Nicaragua). Photo: D. M. Dehling
head; venter white in life; ventrals 132-137
(males) and 122-140 (females) ..... .. ... . .
. . ..... . .. . ..... .. .. ... . . . .... Urotheca decipiens
b White ocellus formed on each side of back
of head; venter reddish color in life; ven- Further Reading
trals 135-164 (males) and 147-176 (fe- MYERS 1974, SAVAGE & CROTHER 1989, SAVAGE &
males) ........ .... ..... .... Urotheca guentheri LAHANAS 1989, MCCRANIE et al. 2001

303
 Colubridae

Fig. 817. Xenodon rabdocephalus (upper Bladen River, Ek Xux, Toledo District, Belize).
Photo: P. Stafford

Xenodon Xenodon rabdocephalus (WIED 1824), Isis von

Oken 6: 668; type locality: Brazil. TL to 790
mm. Guerrero and Veracruz, Mexico, to
The species of the genus Xenodon are Ecuador, Bolivia and Brazil, sea level to 1300
strong, heavily-built rear-fanged snakes m elevation in dry forest, wet forest, and rain
that are similar in their external appear- forest. According to WILSON & MEYER (1985),
the division into subspecies is insufficiently
ance to the dangerous fer-de-lance, supported, and I prefer not to recognize any
Bothrops asper, and are known in Spanish subspecies.
as "falso terciopelo''. If the false fer-de-
lance feels threatened, it will flatten its
Further Reading
body dorsoventrally and gape with an open WILSON & MEYER 1985
mouth. Xenodon rabdocephalus feeds pre-
dominantly on frogs and toads (LEE 1996).
Toads in particular are known to inflate
themselves when threatened in order to
make it more difficult for predators to
swallow them. Xenodon rabdocephalus is
able to puncture its prey with the enlarged
rear teeth of the upper jaw, so that they
deflate and are easier to swallow. This spe-
cies reproduces by laying eggs in clutches
of9-10 during the rainy season (CAMPBELL
1998a). Freshly hatched young have been
seen during the months from June until
November in the northern portion of Xenodon rabdocephalus
Central America (CAMPBELL 1998a).
304

Elapidae
In addition to the true coral snakes (genus
Micrurus), which are distributed in
America, the family Elapidae also includes
the cobras, mambas, and kraits in Asia and
Africa, the Australian venomous snakes,
and the sea snakes. At various times in the
past, the latter have been split off from the
Elapidae into a separate family ("Hydro-
phiidae"), however, subjected to more
recent investigations, this division has not
been retained (GREENE 1997). Char-
acteristic of all members of the family
Elapidae is a pair of deeply grooved fangs,
located at the front of the upper jaw, which
are relatively immobile.
Fig. 818. Micrurus diastema (Vaca Plateau, Belize).
Elapidae
Micrurus
In view of the highly potent venom of the
Micrurus species, extreme caution is ad-
vised in dealing with them. Those rare
instances in which a Micrurus bite is
sustained tend to be fatal. The venom is
predominantly neurotoxic, acting as a par-
alytic agent. In Central America, there are
16 species of the genus Micrurus, all of
which have very conspicuous and strik-
ingly colorful coloration. Coral snakes feed
predominantly on other snakes, but will
also eat eels, caecilians, and lizards. The
species of the genus Micrurus reproduce
oviparously, and produce clutches of 1-13
eggs, which hatch after 70-80 days (ROZE
1996). The key to Micrurus below is based
largely on ROZE (1996). I have followed
CAMPBELL & LAMAR (2004) in treating mos-
quitensis as a subspecies of M. nigrocinc-
tus.
Photo: P. Stafford
305
 Elapidae

Micrurus alleni SCHMIDT 1936b, Field Mus. Micrurus elegans (JAN 1858), Rev. Mag. Zool.
Nat. Hist. Zool. Ser. 20: 209; type locality: Rio (2) 10: 524; type locality: Mexico. TL to 733
Mico, 7 miles above Rama, Atlantico Sur, mm. Central Veracruz, Mexico, to Alta
Nicaragua. TL to 1165 mm. Eastern Verapaz, Guatemala, 100-1700 m elevation in
Honduras to eastern Panama, sea level to dry forest and wet forest. In Central America,
1400 m elevation in rain forest. the subspecies Micrurus e. veraepacis
SCHMIDT 1933c occurs.
Micrurus ancoralis (JAN 1872), in JAN &
SoRDELLI, Icon. Gen. Ophid., Livr. 42: pl. 4, fig. Micrurus hippocrepis (PETERS 1862), Monats.
2; type locality: Ecuador. TL to 1480 mm. Akad. Wiss. Berlin 1861: 925; type locality:
Eastern Panama to southwestern Ecuador, Santo Tomas [= Puerto Matias de Galvez,
sea level to 1000 m elevation in rain forest. In according to PETERS & OREJAS-MIRANDA
Central America, the subspecies Micrurus a. 1970], Guatemala. TL to 710 mm. Belize and
Jani SCHMIDT 1936c occurs. eastern Guatemala, sea level to 600 m eleva-
tion in rain forest and pine forest.
Micrurus bogerti ROZE 1967, Amer. Mus.
Novit. 2287: 9; type locality: Tangola-Tangola Micrurus latifasciatus SCHMIDT 1933c, Field
(Tangolunda), east of Puerto Angel, Oaxaca, Mus. Nat. Hist. Puhl. Zool. Ser. 20: 35; type
Mexico. TL to 770 mm. Pacific side of the locality: Finca El Cipres, Volcan Zunil,
Isthmus of Tehuantepec, Mexico, sea level to Suchitepequez, Guatemala. TL to 1140 mm.
400 m elevation in dry forest. Oaxaca, Mexico, to western Guatemala, sea
level to 1200 m elevation in dry forest and
Micrurus browni SCHMIDT & SMITH 1943, wet forest.
Field Mus. Nat. Hist. Puhl. Zool. Ser. 29: 29;
type locality: Chilpancingo, Guerrero, Mexico. Micrurus mipartitus (DUMERIL, BIBRON &
TL to 830 mm. Guerrero, Mexico, to south- DUMERIL 1854), Erp. Gen. 7: 1220; type locali-
western Honduras, 500-2000 m elevation in ty: "Col. Rio-sucio ou senio" [ROZE 1967 sug-
dry forest, cloud forest and pine-oak forest. In gested that this might be the same as "Simi,
Central America, the subspecies Micrurus b.
browni (distribution same as species except
for the Antigua Basin, Guatemala) and
Micrurus b. importunus ROZE 1967 (Antigua
Basin, Guatemala) occur.
Micrurus clarki SCHMIDT 1936b, Field Mus.
Nat. Hist. Puhl. Zool. Ser. 20: 211; type locali- Micrurus al/eni
ty: Yavisa, Darien, Panama. TL to 832 mm. • Micrurus ancoralis
Southeastern Costa Rica to eastern Panama
(possibly to western Colombia), sea level to
900 m elevation in rain forest.
Micrurus diastema (DUMERIL, BIBRON &
DUMERIL 1854), Erp. Gen. 7: 1222; type locali-
ty: Mexico. TL to 895 mm. Central Veracruz
and northern Oaxaca, Mexico, to western
Honduras, 400-1500 m elevation in wet forest
and cloud forest. In Central America, the sub-
species Micrurus d. aglaeope (COPE 1859)
(northwestern Honduras), Micrurus d. alie-
nus (WERNER 1903b) (the north of the
Yucatan Peninsula), Micrurus d. apiatus (JAN
Micrurus browni
1858) (Alta Verapaz and Huehuetenango, • Micrurus bogerti
Guatemala), Micrurus d. macdougalli ROZE • Micrurus stuarti
1967 (eastern Oaxaca, Mexico) and Micrurus
d. sapperi (WERNER 1903b) (the southern por-
tion of the Yucatan Peninsula) occur.
Micrurus dissoleucus (COPE 1860a), Proc.
Acad. Nat. Sci. Philadelphia 11: 345; type
locality: Venezuela. TL to 376 mm. Central
Panama to Colombia and Venezuela, sea level
to 250 m elevation in dry forest and wet
forest. In Central America, the subspecies
Micrurus d. dunni BARBOUR 1923 occurs.

306

Colombia"). TL to 1000 mm. Eastern Panama
and northwestern South America, sea level to
850 m elevation in rain forest. In Central
America, the nominate subspecies occurs.
Micrurus multifasciatus (JAN 1858), Rev.
Mag. Zool. (2) 10: 521; type locality: Central
America. TL to 1130 mm. Nicaragua to
Central Panama, sea level to 1200 m eleva-
tion in rain forest. The subspecies Micrurus
m . multifasciatus (Central Panama) and
Micrurus m. hertwigii (WERNER 1897)
(Nicaragua to northwestern Panama) are
recognized.
Micrurus nigrocinctus (GIRARD 1854), Proc.
Acad. Nat. Sci. Philadelphia 6: 226; type lo-
cality: Taboga Island, Bay of Panama. TL to
1150 mm. Southern Chiapas, Mexico, and
northern Honduras to northwestern
Colombia (excluding the Yucatan Peninsula;
STAFFORD 2000), sea level to 1500 m elevation
in dry forest, wet forest, and rain forest . In
Central America six subspecies are recog-
nized: Micrurus n. nigrocinctus (Pacific side
of Nicaragua to northwestern Colombia),
Micrurus n. babaspul ROZE 1967 (Corn
Islands, Nicaragua), Micrurus n. coibensis
Micrurus clarki
""' Micrurus dissoleucus
• Micrurus stewarti
• Micrurus diastema
'(/ ""' Micrurus e/egans
• Micrurus hippocrepis
• Micrurus ruatanus
Elapidae
SCHMIDT 1936b (Coiba Island, Panama),
Micrurus n. divaricatus (HALLOWELL 1855)
(northern and central Honduras), Micrurus
n. mosquitensis SCHMIDT 1933c (Caribbean
side of Nicaragua to northwestern Panama)
and Micrurus n. zunilensis SCHMIDT 1932a
(southern Chiapas, Mexico, to El Salvador)
occur.
Micrurus ruatanus (GONTHER 1895), Biologia
Centrali-Americana. Reptilia and Batrachia:
185; type locality: Isla de Roatan, Islas de la
Bahia, Honduras. TL to 681 mm. Isla de
Roatan, Honduras.
Micrurus stewarti BARBOUR & AMARAL 1928,
Bull. Antivenin Inst. Amer. 1: 100; type lo-
cality: Nombre de Dios, Sierra de la Bruja,
Panama. TL to 833 mm. Central Panama,
500-1200 m elevation in rain forest.
Micrurus stuarti ROZE 1967, Amer. Mus.
Novit. 2287: 47; type locality: Finca La Paz
[= 1450 m elevation, 18 km N Coatepeque),
San Marcos, Guatemala. TL to 745 mm.
Southwestern Guatemala, 800-1600 m eleva-
tion in wet forest and cloud forest.
• Micrurus latifasciatus
Micrurus multifasciatus
""' Micrurus mipartitus
307
 Elapidae

Fig. 819. Micrurus alleni (Costa Rica). Fig. 822. Micrurus browni (Tuxtla Gutierrez,
Photo: R. W. Van Devender Chiapas, Mexico). Photo: G. Kohler

Fig. 820. Micrurus alleni (Bartolo, Rio San Fig. 823. Micrurus clarki (Comarca de San
Juan, Nicaragua). Photo: G. Kohler Blas, Panama). Photo: C. W. Myers

Fig. 821. Micrurus alleni (Bartolo, Rio San Fig. 824. Micrurus diastema (Estacion Juarez,
Juan, Nicaragua). Photo: G. Kohler Chiapas, Mexico). Photo: A. Ramirez V.
308
 Elapidae

Fig. 825. Micrurus diastema (near Quebrada Fig. 828. Micrurus latifasciatus (Quetzaltenan-
Grande, Copan, Honduras). Photo: G. Kohler go, Guatemala). Photo: J. A. Campbell

Fig. 826. Micrurus elegans (El Ocote region, Fig. 829. Micrurus multifasciatus (Reserva
Chiapas, Mexico). Photo: G. Kohler Foresta! Fortuna, Panama). Photo: M. Schroth

Fig. 827. Micrurus hippocrepis (Livingston, Fig. 830. Micrurus multifasciatus (Costa Rica).
Izabal, Guatemala). Photo: G. Kohler Photo: T. Leenders
309
 Fig. 831. Micrurus nigrocinctus (El Cusuco, Fig. 834. Micrurus nigrocinctus (Gamboa
Honduras). Photo: J. Kolby Forest Resort, Panama). Photo: M. Lundberg

Fig. 832. Micrurus nigrocinctus (Cerro Fig. 835. Micrurus ruatanus (Isla de Roatan,
Saslaya, Nicaragua). Photo: G. Kohler Honduras). Photo: J. R. McCranie

Fig. 833. Micrurus nigrocinctus (Rio San Juan, Fig. 836. Micrurus stewarti (Panama).
Nicaragua). Photo: J. Sunyer Photo: M. A. Guerra
310
311
 Elapidae
Key to Micrurus
1 a Black rings arranged in triads ................. 2
b Black rings not arranged in triads ........... 4
2 a Between pale head band and first red ring
only a single black ring .... ...... .... .. .. ...
.. .. .. ... ...... .... .. ...... Micrurus dissoleucus
b Between pale head band (which may be
interrupted in the middle of head) and first
red ring one pair or a triad of black rings 3
3 a Head largely red (Fig. 840a); between pale
head band and first red ring a triad of
black rings .... ..... .. .. Micrurus ancoralis
b Head largely black (Fig. 840g); between
pale head band and first red ring one pair
of black rings ... ......... . Micrurus elegans
4 a Body with only two colors (black and red or
black and white) ......................................... 5
b Body with three colors (black-red-yellow or
black-red-white) ... .... ................... ........... 10
5 a With alternating longer and shorter black
rings ....... ............... ..... .... .. ....................... 6
b All black rings of equal length ... .............. 7
6 a Ventrals 178-188 (males) and 193-203
(females) ....... .......... Micrurus ruatanus
b Ventrals 200-207 (males) and 224-228
(females) .... ..... ........ . Micrurus stewarti
7 a More than 40 black body rings; 8-15 black
tail rings ... Micrurus diastema (in part)
b Generally fewer than 8 black tail rings; if
8, then fewer than 30 black body rings .... 8
8 a Red body rings at least twice as long as
black rings ...... .................... ........ ...... ....... .
.. .......... Micrurus nigrocinctus (in part)
b Red or white body rings less than double
the length of black rings ..... .. ... ....... .. .... 9
9 a Body with black and white rings; black
neck band usually reaches to the tips of the
parietals (Fig. 840i); pale dorsal rings less
than two dorsals l?ng ·· ···· ·:·······; ·····
... ........ ... ......... ....... Micrurus mipartitus
b Body with black and red rings; black neck
band usually does not reach the tips of the
parietals (Fig. 840j); pale dorsal rings at
least two dorsals long ............................. .
... ...................... Micrurus multifasciatus
lOa Head without pale bands ................ .. .
........ .... ..... Micrurus diastema (in part)
b Head with at least one pale band ........... 11
Ila Black cephalic cap continues as a narrow
stripe along the parietal suture (Fig. 840b)
. . . . . . . . . . . . . .. .. ..... ..... .......... Micrurus alleni
b Head markings different than those above
················ ··· ····· ···· ················· ······ ·· ·· ·· · 12
312
12a Black cephalic cap completely or almost
completely covers the parietals including
the posterior tips ............ Micrurus clarki
b Black cephalic cap does not cover the
posterior tips of the parietals .. .............. 13
13a Fewer than 10 black body rings .............. 14
b Usually more than 9 (seldom 8 or 9) black
body rings ................ .. ... .... ... ................. 17
14a Black neck band reaches 8-13 dorsals
posteriorly ..................... ..... .. ..... .......... 15
b Black neck band reaches fewer than 8 dor-
sals posteriorly ...... .. ........ .. ................. .. 16
15a Males with supra-anal tubercles (Fig. 838);
females with fewer than 213 ventrals
.. .... ... .. ... ........ ... Micrurus latifasciatus
b Males without supra-anal tubercles; fe-
males with more than 212 ventrals .. ....... .
... ... ....... Micrurus hippocrepis (in part)
16a Black neck band reaches 2-3 dorsals
posteriorly; red body scales with or without
little black pigment .... ............... ........ .
... ... ....... Micrurus hippocrepis (in part)
b Black neck band reaches 4-5 dorsals
posteriorly; tips of red body scales with
black spot or these scales with irregularly
distributed black pigment ............... .... . .
... .... .......... Micrurus diastema (in part)
l 7a Black neck band reaches the parietals (Fig.
840e) ........................ ... ... ... ....... ............ 18
b Black neck band does not reach the
parietals (Figs. 840c, k) ......... .............. 19
18a Males with supra-anal tubercles (Fig. 838);
females with fewer than 40 subcaudals ...
............ Micrurus nigrocinctus (in part)
b Males without supra-anal tubercles; fe-
males with more than 40 subcaudals ..... .. ..
... ............. ..Micrurus diastema (in part)
19a Red body scales without conspicuous black
tips, at most with very small dots or
brownish shading .. ....... ..... .... ........... 20
b Red body scales with conspicuous black
tips ........ .... ...... .. .... ...... ........... .. ........... 23
supra-anal tubercles
Fig. 838.1\{icrurus
nigrocinctus (male;
lateral view of cloacal
region) .
Drawing by M. Vesely
313
 Elapidae

20a Ventrals usually more than 207 (males)

and more than 221 (females) .. .... ... ....... 21
b Ventrals fewer than 208 (males) and fewer
than 222 (females) .. .... ... ....... ............. .. 22
21a Males with supra-anal tubercles; females
with 17-27, usually more than 18, black
body rings ... . Micrurus browni (in part)
b Males without supra-anal tubercles; fema-
les with 13-18 black body rings ........... ....... .
... . . . . . . . .. . . . . . ............ ..... .. Micrurus bogerti
a. M. ancoralis b. M. a/Jeni c. M. bogerti
22a Black neck band reaches at least 3 dorsals
posteriorly; dorsal black body rings at least
3 dorsals long; first red ring usually less
than 13 dorsals long; males with supra-
anal tubercles .......... ................... ........... .
... ......... Micrurus nigrocinctus (in part)
b Black neck ring reaches 2-3 dorsals
posteriorly; dorsal black body rings usual-
ly less than 3 dorsals long; first red ring
usually more than 3 dorsals long; males
without supra-anal tubercles ........... ... .
. . ........... Micrurus hippocrepis (in part)
23a Not all red body scales with black tips, d. M. browni e. M. diastema f. M. dissoleucus
rather irregularly distributed, some with
black tips, others not ......... ............... ... .. .
. . ... . ........... Micrurus diastema (in part)
b All red body scales with regular black tips
........... ..... ... ... ... .... ... ........... ..... ............ 24
24a Ventrals usually fewer than 206 (males)
and fewer than 223 (females) ................. 25
b Ventrals more than 205 (males) and more
than 223 (females) .......... ..... ................ 26
25a 11-75, usually more than 15 black body
rings; males with supra-anal tubercles
............ Micrurus nigrocinctus (in part) g. M. elegans h. M. /atifasciatus i. M. mipartitus
b 8-17, usually fewer than 15 black body
rings; males without supra-anal tubercles .
..... ...... ....... Micrurus diastema (in part)
26a Black tips on red scales somewhat irregu-
lar, very large (cover about 3/4 of the red
scales); black neck band reaches 6-8
(usually 7) dorsals posteriorly .................... .
.................... .......... ...... ... Micrurus stuarti
b Black tips on red scales regular, small
(cover about 1/4 of the red scales); black
neck band reaches 4-6 (usually 5) dorsals
posteriorly .................... .......... .. ...... ...... 27 j. M. multifasciatus k. M. nigrocinctus I. M. stewarti
27a Snout tip black ... .. .. .... ........ .. .... .. ...... .
........... Micrurus nigrocinctus (in part) Fig. 840. Dorsal head coloration in Micrurus;
partially after ROZE (1996).
b Snout tip yellow or white ....... ... ... .. .
. . .. . . . . . . . . .. ...... Micrurus browni (in part)

Further Reading MCCRANIE &WILSON 1991a, WILSON et al. 1992,

AMARAL 1926, ROZE 1967, SAVAGE & VIAL 1974, MCCRANIE 1993b, WILSON et al. 1996, STAFFORD
GREENE & SEIB 1983, ROZE 1983, 1996, 2000
314

Pelamis
Only one of the approximately 55 species of
sea snakes occurs along the Pacific coast of
Central America, the pelagic sea snake
(Pelamis platura). The correct scientific
name of the pelagic sea snake is platura,
not platurus (see LANZA & BOSCHERINI
2000). Their absence along the Caribbean
Coast implies that this species reached the
Pacific fewer than four million years ago,
after the Panama land bridge was formed
and separated the two oceans. Another sea
snake, the banded sea snake (Laticauda
colubrina), was reported in Nicaragua by
VILLA (1962) and, as a result, included in
various checklists for the region (J. PETERS
& 0REJAS-MIRANDA 1970, VILLA 1984, VILLA
et al. 1988). However, the existence of this
Fig. 841. Pelamis platura (Playa El Coco, Costa Rica).
Elapidae
snake is considered highly unlikely
(KOHLER 1999c), particularly as the
sightings that have been reported so far
have not been supported by any specimens.
If any specimens of Laticauda colubrina
have reached Central America in the past,
then these were likely isolated stray speci-
mens that have not established self-sustai-
ning populations.
The pelagic sea snake (Pelamis platura)
has a striking appearance, with its yellow-
black coloration. It is remarkably well
adapted for a life at sea, with its flattened
rudder-like tail, nostrils located on the
upper side of the snout, and salt glands
under the tongue.
Pelamis platura usually is seen floating
on the surface, which is why some au-
Photo: R. D. Bartlett
315
 Elapidae

thors have described them as pelagic

(GREENE 1997). They particularly like to
dwell in calm waters with a lot of small
fish, their preferred food. They catch
their prey with quick sideways bites,
sometimes swimming rapidly back-
wards. This species has highly effective
venom at its disposal, which it uses to
catch prey and to protect itself from pre-
dators. Bites from these sea snakes
should be taken very seriously, even
though the few documented cases have
been accompanied by relatively mild
symptoms, such as localized pain and
swelling CKROPACH 1975, SOLORZANO
1995). Fig. 842. Pelamis platura (Playa El Coco,
Costa Rica). Photo: R. D. Bartlett
Pelagic sea snakes will mate floating on
the surface of the water and also bear
their young in the water. Sea snakes
shed at frequent intervals (approximate-
ly every 3-6 weeks), among other rea-
sons, to rid themselves of unwanted
external parasites (algae, as well as
barnacles and mussels). In order to shed
their old skin, the snakes will form
complex knots with their bodies, and
then crawl out through them. The shed
skin is usually left behind still tied in
knots (GREENE 1997). Apparently
Pelamis platura has few natural pre-
dators, perhaps due to its warning color-
ation and highly toxic venom. Sea birds
have been observed catching pelagic sea
snakes and then dropping them again
after a short time (GREENE 1997). The Fig. 843. Pelamis platura (Playa La Flor,
most deadly circumstance for a Pelamis Nicaragua). Photo: G. Kohler
platura is to be washed up on the beach.
The animal would have a very limited
chance of being able to return to the
water under its own power and would
die after a short time.

Pelamis platura (LINNAEUS 1766), Systema Further Reading

Naturae, ed. 12: 391; type locality: unknown. KROPACH 1975, PICKWELL & CULOTIA 1980,
TL to 750 mm (the exception to 1100 mm). VORIS 1983, SLOWINSKI et al. 1997, KEOGH 1998,
Indonesia, tropical Pacific Ocean, west coast SLOWINSKI & KEOGH 2000
of America from Mexico to Ecuador.

316

Viperidae
Except for the species of the genus
Atractaspis, all venomous snakes that have
enlarged, highly movable, tube-like hollow
fangs have been placed in the family
Viperidae. About 160 of these species are
characterized by the presence of pit organs
and have been combined in the subfamily
Crotalinae (pitvipers or lanceheads), to
which all Central American vipers belong.
Pit organs are heat-sensitive, tube-like
depressions found between the eye and the
nostril. They assist the snakes in tracking
down warm-blooded prey, such as small
mammals. All vipers have a highly evolved
venom delivery apparatus. When the
mouth of the snake is closed, the long fangs
are protected in a fold of skin and they
Fig. 844. Bothriechis nigroviridis (Costa Rica).
Viperidae
swing down and forward when the animal
is in the process of striking. The venom of
most vipers disrupts both the circulatory
system, as well as tissue and, in some cases
(e.g. , South American rattlesnakes,
Crotalus durissus terrificus) , also has
strong neurotoxic effects. With the excep-
tion of the bushmasters (genus Lachesis),
which reproduce oviparously, all other
Neotropical pitvipers are livebearers.
Further Reading
CAMPBELL & LAMAR 1989, 1992, CADLE 1992,
WERMAN 1992, 1999, SALOMAO et al. 1999
Photo: R. W. Van Devender
317
 Viperidae

Key to Viperidae postoculars preoculars canthal

1 a Tail end with rattle ............ ........ Crotalus
b Tail end without rattle ..... ....... .................. 2
2 a Underside of tail tip with 4-5 rows of nar-
row spiny scales (Fig. 846a) ... .... Lachesis
b Underside of tail with 1-2 rows ofrelative-
ly wide, flat scales (Figs. 846b,c) ... ........... 3
3 a Snout covered with large symmetrical
shields (Fig. 847a) .... ............. Agkistrodon
b Snout covered with small irregular scales
(Figs. 847b,c) .......... ...... .. .. ... ........ ....... 4
4 a All or most subcaudals paired ..............
.. ... .... ..... ...... .... ......... ...... ...... Bothrops ultimate mental
infralahi<il first infralabial
b All or most subcaudals single .. ............. .... 5
5 a Eyelid with projecting, elongated, pointed
scales ... ....... .. ...... Bothriechis (in part) Fig. 845. Scalation characters on the head of a
viperid snake (Cerrophidion godmani).
b Eyelid without projecting, elongated, Drawing by M. Vesely
pointed scales .. .. .......... .... .... ....... ............ 6
6 a Unicolor green in life (blue in preserva-
tive); arboreal with prehensile tail .............
... . . . .. . .. . . ... . . ..... ...... Bothriechis (in part)
b Ground color red-brown, gray, or brown;
terrestrial without prehensile tail ........... 7
7 a Dorsal pattern usually made up of opposite
or alternating dark paravertebral spots
and one pale vertebral stripe ...... .. .
... . . .. ... . . . . . . . . . .. . . . . . . .... . .. ......... Porthidium
b Dorsal pattern different from that above
(often a dark zigzag pattern present)
......................................... .............. ...... ...... 8
8 a 1-7 scales between the supraoculars; body
scales keeled, but not conspicuously rough;
19-23 dorsal rows at midbody .... ............. .
........... ........... ..... .. ... ..... ... Cerrophidion
a. Lachesis b. Bothrops c. Bothriechis
b 9-12 scales between the supraoculars (Fig.
847b); body scales very rough; 23-31 dorsal Fig. 846. Scalation of distal portion of tail
rows at midbody ............ .... .... Atropoides (ventral view) in some pitvipers.

rostral

supraoculars

Fig. 84 7. Scalation of
dorsal surface of head.
a. Agkistrodon b. Atropoides c. Bothrops

318
 Viperidae

Fig. 848. Agkistrodon bilineatus (Taxisco, Santa Rosa, Guatemala). Photo: G. Kohler

Agkistrodon tion in dry forest and savanna. In Central

America, the subspecies Agkistrodon b. bilin-
eatus (along the Pacific coast of Mexico to El
The four species of the genus Agkistrodon Salvador), Agkistrodon b. howardgloydi
are restricted in their distribution to North CONANT 1984 (along the Pacific coast from
and Central America. Agkistrodon bilinea- Honduras to Costa Rica) and Agkistrodon b.
tus is a mid-size powerful pitviper that pre- russeolus GLOYD 1972 (Yucatan Peninsula)
occur.
fers to live in dry areas, often near water.
At nightfall, it will become active and leave
its daytime hiding place.Agkistrodon bilin-
eatus feeds primarily on rodents, but will
also consume fish, frogs, lizards, and
snakes (GLOYD & CONANT 1990). The spe-
cies is a livebearer and produces 6-20
young per litter (ALVAREZ DEL TORO 1983,
GLOYD & CONANT 1990).

Agkistrodon bilineatus (GONTHER 1863), Ann.

Mag. Nat. Hist. (3) 12: 364; type locality:
Pacific coast of Guatemala. TL to 1380 mm. Agkistrodon bilineatus
Southern Sonora, Mexico, along the Pacific
coast as far northwestern Costa Rica, as well
as disjunct in Grijalva Basin (Chiapas,
Mexico) and in the northern portion of the
Yucatan Peninsula, sea level to 1500 m eleva- Further Reading
GLOYD & CONANT 1990, PARKINSON et al. 2000
319
 Viperidae
Fig. 849. Atropoides picadoi (Costa Rica).
Photo: G. Kohler
Atropoides
The three species that are included in the
genus Atropoides are stockily-built ground-
dwelling pitvipers that are distributed in
Mexico and Central America (WERMAN
1992). Two species of the genus occur in
Central America. They are usually found in
remote areas of extensive primary growth.
As juveniles, their diet consists of insects,
frogs, and lizards, whereas adults usually
hunt small mammals (ALVAREZ DEL TORO
1983). During the rainy season, A. nummi-
fer will give birth to 5-34 young with a total
length of 178-220 mm (SOLORZANO 1989,
LEE 1996). Neonate Costa Rican A nummi-
fer exhibit a marked sexual dichromatism
in head pattern (SOLORZANO 1990).
"' Atropoides picadoi
Atropoides nummifer
320
Fig. 850. Atropoides nummifer (near Quebrada
Grande, Copan, Honduras). Photo: G. Kohler
Atropoides nummifer (ROPPELL 1845), Verb.
Mus. Senckenberg 3: 313; type locality:
Mexico. TL to 800 mm. San Luis Potosi and
Oaxaca, Mexico, to Central Panama, 100-
1500 m elevation in dry forest and wet forest.
Atropoides picadoi (DUNN 1939b), Proc. Biol.
Soc. Washington 52: 165; type locality: La
Palma, 4500 ft, Costa Rica. TL to 1000 mm.
Costa Rica and western Panama, 50-1500 m
elevation in rain forest and cloud forest.
Key to Atropoides
1 a More than 138 ventrals .... ............. ........ .
........ .... .. .................... Atropoides picadoi
b Fewer than 138 ventrals ......................... .
..... .. .... ................. Atropoides nummifer
Further Reading
CAMPBELL & LAMAR 1989, SOLORZANO 1989,
WERMAN 1992, CASTOE et al. 2003
Abb. 851. Juvenile Atropoides nummifer
(Union Barrios, Baja Verapaz, Guatamala).
Photo: G. Kohler
 Viperidae

Bothriechis

The genus Bothriechis includes a series of

small (generally reaching maximum total
lengths between 700 and 850 mm, rarely to
1000 mm), arboreal, prehensile-tailed pit-
viper species, which occur predominantly
in Central America. Only one species, B.
schlegelii, also occurs north of the Isthmus
of Tehuantepec and south of Panama. This
species and B. supraciliaris are characteri-
Fig. 854. Bothriechis thalassinus (Montan.as de
la Uni6n, Zacapa, Guatemala).
Photo: G. Kohler

zed by spiky, enlarged, and protruding

eyelid scales. All other species of Bothri-
echis, which are mostly green colored in
life, occur in isolated, wet, montane forests,
usually above 1000 m elevation, but two
species (B. bicolor and B. thalassinus) des-
cend to about the 500 m contour.

During the day, these pitvipers rest, usual-

ly well-camouflaged, in the branches or on
tree trunks, making it prudent to check
carefully before leaning on a tree or reach-
ing for a branch when in Bothriechis habi-
tat. Although relatively small, B. schlegelii
possesses highly toxic venom, and bites by
Fig. 852. Bothriechis marchi (Cusuco, Cortes, this species can be life-threatening to
Honduras). Photo: G. Kohler humans (SEIFERT 1983). The Bothriechis
species feed on frogs, lizards, small mam-
mals, and birds (DUELLMAN 1963, ALVAREZ
DEL TORO 1983, CAMPBELL 1998a).
Bothriechis marchi specializes in a diet of
frogs of the genus Eleutherodactylus, and
in Cusuco National Park (Honduras) has
nearly vanished since the Eleuthero-
dactylus populations along the streams
there have plummeted in numbers (J.R.
MCCRANIE pers. comm. 1999). All Bothri-
echis species bear live young. For B. schle-
gelii, the litter size varies from 12 to 20
young (ANTONIO 1980, CAMPBELL 1998a).
Fig. 853. Bothriechis schlegelii (near Bothriechis nigroviridis has a distinctly
Tegucigalpita, Cortes, Honduras). seasonal reproductive cycle, producing 6-8
Photo: G. Kohler
321
 Viperidae
young (165-177 mm total length) during
the rainy season, with coloration identical
to that of the adults (SOLORZANO 1997).
The young of B. lateralis, on the other
hand, are brown in color (HOHMEISTER
1994)
Bothriechis aurifer (SALVIN 1861a), Proc. Zoo}.
Soc. London 1860: 459; type locality: Cohan,
Alta Verapaz, Guatemala. TL to 1000 mm.
Disjunct in west-central Chiapas, Mexico,
and central Guatemala, 1200-2300 m eleva-
tion in cloud forest and pine-oak forest.
Bothriechis bicolor (BocOURT 1868), Ann. Sci.
Nat. Paris (5) 10: 202; type locality: Forests of
St. Augustin, 610 m elevation, on western
slope of Cordillera, Solola, Guatemala. TL to
1000 mm. Disjunct from southeastern
Chiapas, Mexico, to Guatemala, 500-2000 m
elevation in wet forest and cloud forest.
Bothrops ornatus JULIA-ZERTUCHE & VARELA
JULIA 1978 is regarded to be a synonym of
Bothriechis bicolor according to CAMPBELL &
LAMAR (1989).
Bothriechis lateralis PETERS 1863d, Monats.
Akad. Wiss. Berlin 1862: 674; type locality:
Veragua and Volcan Barba, Costa Rica. TL to
1000 mm. Costa Rica and western Panama,
1200-1800 m elevation in rain forest and
cloud forest.
Bothriechis marchi (BARBOUR & LOVERIDGE
1929b), Bull. Antivenin Inst. America 3: 2;
type locality: Quimistan, Departamento de
Santa Barbara, Honduras. TL to 968 mm.
Northwestern and northern, central
Honduras, 500-1550 m elevation in rain
forest and cloud forest.
Bothriechis
.a. B. bicolor
"'B. aurifer
.a. B. thalassinus
• B. lateralis
• B. marchi
• B. rowleyi
322
Bothriechis nigrouiridis PETERS 1859,
Monats. Akad. Wiss. Berlin 1859: 278; type
locality: Volcan Barba, Costa Rica. TL to 800
mm. Costa Rica and western Panama, 1350-
3000 m elevation in rain forest and cloud
forest.
Bothriechis rowleyi (BOGERT 1968b), Amer.
Mus. Novit. 2341: 3; type locality: on a ridge
that extends northward from Rancho Vicente,
1520 m elevation, Colonia Rodolfo Figueroa,
approx. 5 miles W Cerro Baul, Distrito de
Juchitan, Oaxaca, Mexico. TL to 970 mm.
Disjunct in eastern Oaxaca and western
Chiapas, Mexico, 1500-1830 m elevation in
cloud forest and pine-oak forest. Bothrops
nigrouiridis macdougalli SMITH & MOLL 1969
is regarded to be a synonym of Bothriechis
rowleyi according to CAMPBELL & LAMAR
(1989).
Bothriechis schlegelii (BERTHOLD 1845),
Nachr. Georg-Augusts Univ. und K. Ges.
Wiss. Gottingen 3: 42; type locality: western
Colombia (see MYERS & BOHME 1996). TL to
800 mm. Northern Chiapas, Mexico, to north-
ern South America, sea level to 1400 m eleva-
tion (even higher in South America) in wet
forest, rain forest, and cloud forest.
Bothriechis supraciliaris (TAYLOR 1954), Univ.
Kansas Sci. Bull. 36: 791; type locality:
Mountains near San Isidro del General, San
Jose, Costa Rica. TL to 600 mm. Southeastern
Costa Rica and western Panama, 800-1700 m
elevation in rain forest and cloud forest.
Bothriechis thalassinus CAMPBELL & E. SMITH
2000, Rev. Biol. Trop. 48: 1002; type locality:
Finca La Firmeza, 15°22'29"N, 88°41'44"W,
900 m, Sierra de Cara}, Izabal, Guatemala.
TL to almost 1000 mm. Disjunct from eastern
Guatemala to western Honduras, 885-1730 m
elevation in wet forest and cloud forest.
Bothriechis
B. schlegelii
.a. Bothriechis supraciliaris
"' Bothriechis nigroviridis
 Viperidae

Fig. 855. Bothriechis aurifer (Union Barrios, Fig. 858. Bothriechis aurifer.
Cerro Verde, Baja Verapaz, Guatamala). Photo: R. D. Bartlett
Photo: G. Kohler

Fig. 856. Bothriechis bicolor (La Concordia, Fig. 859. Bothriechis bicolor (Chiapas, Mexico).
Chiapas, Mexico). Photo: A. Ramirez V. Photo: A. Ramirez V.

Fig. 857. Bothriechis lateralis (Rio Chevo, Fig. 860. Bothriechis lateralis (La Nevera,
Chiriqui, Panama). Photo: J. Sunyer Panama). Photo: G. Kohler
323
 Fig. 861. Bothriechis nigroviridis (La Nevera, Fig. 864. Bothriechis nigroviridis (Costa Rica).
Panama). Photo: G. Kohler Photo: R. W. Van Devender

Fig. 862. Bothriechis nigroviridis (Jurutungo, Fig. 865. Bothriechis nigroviridis (Jurutungo,
Chiriqui, Panama). Photo: G. Kohler Chiriqui, Panama). Photo: G. Kohler

Fig. 863. Bothriechis nigroviridis (La Nevera, Fig. 866. Bothriechis nigroviridis (San Gerardo,
Panama). Photo: G. Kohler Costa Rica). Photo: R. W. Van Devender

324
Fig. 867. Bothriechis rowleyi (Chiapas, Fig. 870. Bothriechis rowleyi (Chiapas,
Mexico). Photo: A. Ramirez V. Mexico). Photo: A. Ramirez V.

Fig. 868. Bothriechis schlegelii (Bartola, Rio Fig. 871. Bothriechis schlegelii (Rio San Juan,
San Juan, Nicaragua). Photo: G. Kohler Nicaragua). Photo: J. Sunyer

Fig. 869. Bothriechis supraciliaris (Cerro Fig. 872. Bothriechis supraciliaris (Chiriqui,
Paraguas, Costa Rica). Photo: G. Kohler Panama). Photo: G. Kohler

325
 Viperidae
Fig. 873. Bothriechis lateralis (Monteverde,
Costa Rica). Photo: R. W. Van Devender
Key to Bothriechis
1 a Eyelid (superciliaries) scales spinelike,
enlarged, and protruding ............... ......... 2
b Eyelid scales (superciliaries) not spinelike,
enlarged, and protruding ........ ................. 3
2 a Venter with dark spotting; dorsal mark-
ings diffuse with irregular spots, skin be-
tween these markings with numerous
small dots or flecks of secondary pigmenta-
tion (except for the unicolor yellow
"Oropel" color variations); ventrals 138-166
(146-167 in specimens from Costa Rica) .....
............................ Bothriechis schlegelii
b At least the anterior two-thirds of venter
without dark spotting; dorsum with clearly
bordered spots or crossbands, skin between
these markings unicolor and without
secondary pigment (ground color uniform);
ventrals 141-150 ......................................... .
....................... Bothriechis supraciliaris
3 a Dorsal surface of head with dark spots or
black speckles; a black postocular stripe
generally present ...... .... ..... ...... ........ .... 4
b Dorsal surface of head unicolor without
black spots or speckles; black postocular
stripe generally not present, if a postocular
stripe present, it is bluish ... ................. 5
4 a Dorsum without heavy black mottling;
venter unicolor yellow-green; tail tip yel-
326
low; snout covered with relatively large
scales; 1-5 scales between the supraocu-
lars; iris yellowish green or bronze
............. ..... ....... .. ... ........... Bothriechis
aurifer
b Dorsum with heavy black mottling; venter
with black speckling; tail tip black; snout
covered with small scales; 4-8 scales be-
tween the supraocular~; iri~ bl~ckish. ··;··:··
........................ Bothnechis nigrovindis
5 a Venter yellow-green with yellow or white
paraventral stripes; tail tip yellow; gener-
ally 23 dorsal rows at mid body ............. .
... . . . . . . . . . ..... .......... ... Bothriechis lateralis
b Venter yellow-green without pale
paraventral stripes; tail tip black; fewer
than 23 dorsal rows at midbody
......... ... .... .......... 6
6 a Scales on top of head small, each scale with
a single keel, adjacent keels parallel; 7-11,
rarely 5, scales between the supraoculars;
generally 21 dorsal rows at midbody ........ 7
b Scales on head (anterior to level of suprao-
culars) irregular in shape, many large and
flat or with irregular and multiple keels at
different angles; 3-7 scales between the
supraoculars; generally fewer than 21 dor-
sal rows at midbody ................................ 8
7 a 27-33 interrictals ..... Bothriechis bicolor
b 23-25 interrictals .................................... .
........................ Bothriechis thalassinus
8 a Snout scales large and smooth .......... .
........ .. ........ ... ......... Bothriechis rowleyi
b Snout scales smaller and more numerous,
some keeled ... .. ...... Bothriechis marchi
Further Reading
H. SMITH & MOLL 1969, CAMPBELL & LAMAR
1989, CROTHER et al. 1992, WILSON & MCCRANIE
1992, SoL6RZANO 1997, SoL6RZANO et al. 1998,
DEJESUS 2007
"' Bothrops punctatus
Bothrops asper

Bothrops
The "terciopelo" or "barba amarilla", as
Bothrops asper is known in Central
America, is the most common and most
dangerous venomous snake in the region.
It lives not only in forested areas, but also
in secondary undergrowth near human
habitation, preferably close to water.
There, this nocturnal pitviper will hunt
frogs, lizards, and small mammals. This
species is highly prolific and can produce
litters of up to 86 young (usually 20-50),
Fig. 874. Bothrops asper (Parque Nacional
Saslaya, Nicaragua). Photo: G. Kohler
Fig. 875. Bothrops asper (Meseta de Chorcha,
Chiriqui, Panama). Photo: G. Kohler
Viperidae
with newborn young found primarily in
the rainy season (SoL6RZANO & CERDAS
1989). Bothrops punctatus is a large, semi-
arboreal pitviper that reaches only the
extreme eastern portion of Panama.
Bothrops asper (GARMAN 1883), Mem. Mus.
Comp. Zool. 8: 124; type locality: Obispo,
Darien, Panama. TL to 2400 mm. Southern
Tamaulipas and Chiapas, Mexico, to north-
western South America, sea level to 1000 m
elevation in rain forest.
Bothrops punctatus (GARC{A 1896), Los
Ofidios Venenosas de Cauca, Cali, Colombia:
31; type locality: Las Montafias de Dagua,
Colombia. TL to 1500 mm. Eastern Panama
along the Pacific side, to northwestern
Ecuador, sea level to 2300 m elevation in wet
forest, rain forest, and cloud forest.
Key to Bothrops
1 a Dorsal pattern with dark triangular spots;
fewer than 70 subcaudals; 23-25 dorsal
rows at midbody ............ . Bothrops asper
b Dorsal pattern with dark bands arranged
in pairs; more than 70 subcaudals; 25-29
dorsal rows at midbody .... ........ ...... .... .
.. . .. . .. .. .. .. .. .. ..... ...... Bothrops punctatus
Further Reading
SOLORZANO & CERDAS 1989, CAMPBELL & LAMAR
1989, 1992
Fig. 876. Bothrops punctatus (Choco, Colombia).
Photo: M. Lundberg
327
 Viperidae

Cerrophidion

The genus Cerrophidion includes four spe-

cies, two of which are distributed in high-
land Central America (CAMPBELL & LAMAR
1992, LOPEZ LUNA et al. 1999). They are
small, ground-dwelling pitvipers that
usually have a dorsal pattern of dark spots
that are often fused into a zigzag stripe.
Cerrophidion godmani feeds primarily on
insects, salamanders, frogs, lizards, and
small mammals (CAMPBELL & SoL6RZANO
1992). The species of the genus
Cerrophidion are livebearers; C. godmani
produces litters of2-12 young (CAMPBELL &
SoL6RZANo 1992). Fig. 878. Cerrophidion godmani (south-
western Honduras). Photo: G. Kohler

Cerrophidion godmani (GUNTHER 1863), Ann.

Mag. Nat. Hist. (3) 12: 364; type locality:
Duenas and other parts of tableland of
Guatemala. TL to 750 mm. Disjunct from
southeastern Oaxaca, Mexico, to western
Panama, 1600-3200 m elevation in rain forest
and cloud forest, as well as in pine-oak forest.
Cerrophidion tzotzilorum (CAMPBELL 1985), J.
Herpetol. 19: 48; type locality: 10.9 km ESE
San Cristobal de las Casas, 2320 m elevation,
Chiapas, Mexico. TL to 500 mm. Meseta
Central de Chiapas, Mexico, 2000-2500 m ele-
vation in pine-oak forest.
Key to Cerrophidion
1 a Most of the lateral dark body spots are 5-7
scales high; dark postorbital band not pale-
edged above and below ................... ... .... .
. . . . . . . . . . . . ............. Cerrophidion godmani
b Most of the lateral dark body spots are 2-3
scales high; dark postorbital band distinct-
ly pale-edged above and below ........... .. . .
... ...... ..... ...... Cerrophidion tzotzilorum
Further Reading
CAMPBELL & LAMAR 1992, CAMPBELL &
SoL6RZANO 1992, LOPEZ LUNA et al. 1999

Cerrophidion godmani
Fig. 877. Cerrophidion tzotzilorum (Chiapas, "" Cerrophidion tzotziforum
Mexico). Photo: G. Kohler
328
 Viperidae

Crotalus

Only one rattlesnake species, Crotalus

simus, occurs in Central America. It lives
as a ground-dweller in dry areas and,
depending on the season, is diurnal, cre-
puscular, or nocturnal. Usually during the
cooler season, they can be seen during the
day outside of their hiding places (LEE
1996). Because farmers will usually kill
nearly every rattlesnake that they come
across, this species has become extremely
rare in many areas. Crotalus simus feeds
primarily on rodents, but will also consume Fig. 879. Crotalus simus tzabcan (Sian Ka'an
Biosphere Reserve, Quintana Roo, Mexico).
lizards and ground nesting birds CK.LAUBER Photo: H. Bahena B.
1956, CAMPBELL 1998a). The species is a
live bearer, producing 15-47 young per
litter (ARMSTRONG & MURPHY 1979,
CAMPBELL 1998a).

Crotalus simus LATREILLE 1801, in SONNINI DE

MANONCOURT & LATREILLE, Hist. Nat. Rept. 3:
335; type locality: "Ceylan". TL to at least
1800 mm. Mexico to Costa Rica, sea level to
1600 m elevation in dry forest and savanna.
In Central America, the subspecies Crotalus
s. simus (Mexico to Costa Rica) and Crotalus
s. tzabcan Kl.AUBER 1952 (Yucatan Peninsula)
occur.

Further Reading
KLAUBER 1956, CAMPBELL & LAMAR 1989, Fig. 880. Crotalus s. simus (Depto. Jutiapa,
MCCRANIE 1993a, CAMPBELL & LAMAR 2004 Guatemala). Photo: G. Kohler

Crotalus simus Fig. 881. Crotalus s. simus (Morgans Rock,

Nicaragua). Photo: J. Sunyer
329
 Viperidae
Lachesis
The genus Lachesis was earlier seen as
monotypic, until RIPA (1994) and ZAMUDIO
& GREENE (1997) established the morpho-
logical and molecular genetic differences
and were able to show that Lachesis was
made up of at least three species. The cur-
rent view is that Lachesis comprises four
species, three of which occur in Central
America (CAMPBELL & LAMAR 2004).
Bushmasters are the largest of the New
World venomous snakes and the only pitvi-
pers there that lay eggs and actually will
brood them. They live as crepuscular and
nocturnal sit-and-wait predators in pri-
mary rain forest and feed predominantly
on mammals. While L. stenophrys can be
considered quite calm, L. melanocephala is
very nervous and aggressive, particularly
the males during mating season (RIPA
1994). Irritated specimens of L . melano-
cephala will sometimes even follow a per-
son for several meters, lifting their heads
cobra-like up to 50 cm above the ground. In
the case of a venomous snake over two
meters in length, this behavior is certain to
inspire a substantial degree of fear (RIPA
1994). The females will retire to a dark and
secure hiding place in order to lay their
eggs, and appear to like abandoned agoutis
burrows for egg laying (RIPA 1994). The
bushmaster will lay 9-19 eggs per clutch,
which are guarded by the female during
the 60-77 days of the incubation period. The
animal surrounds the eggs with the coils of
her body, not eating during the entire
period, and only leaving the clutch for a
short time in order to drink (RIPA 1994).
Freshly hatched bushmaster babies have a
total length of about half a meter (RIPA
1994).
330
Lachesis acrochorda (GARCiA 1896), Ofidios
Venen. Cauca: 102; type locality: "las selvas
del Choc6, a orillas de los rios Atrato, San
Juan, Dagua y Telembi, y ... el camino de
Buenaventura", Colombia. TL to 3000 mm.
Eastern Panama to northwestern South
America (Pacific side of Colombia and nor-
thwestern Ecuador), sea level to 1600 m ele-
vation in rain forest.
Lachesis melanocephala SoL6RZANO &
CERDAS 1986, J. Herpetol. 20: 463; type local-
ity: 9 km N Ciudad Neily, Puntarenas, Costa
Rica. TL to 3600 mm. Southeastern Costa
Rica (Osa Peninsula and adjacent rain forest
areas), sea level to 500 m elevation in rain
forest.
Lachesis stenophrys COPE 1876, J . Acad. Nat.
Sci. Philadelphia (2) 8: 152; type locality:
Sipurio, Costa Rica. TL to 3600 mm.
Southeastern Nicaragua to central Panama,
sea level to 1000 m elevation in rain forest.
Key to Lachesis
1 a Dorsal surface of head black ...................... .
...... .................. Lachesis melanocephala
b Dorsal surface of head pale brown .. .. ...... 2
2 a 211-228 ventrals ... Lachesis acrochorda
b 191-209 ventrals .... Lachesis stenophrys
Further Reading
VIAL & JIMENEZ-PORRAS 1967, MARTiNEZ &
BoLAN"os 1982, SoL6RZANO & CERDAS 1986, RIPA
1994, ZAMUDIO & GREENE 1997, CAMPBELL &
LAMAR 2004
• Lachesis acrochorda
Lachesis stenophrys
.a. Lachesis melanocephala
 Viperidae

Fig. 882. Lachesis acrochorda (Darien region, Fig. 885. Lachesis acrochorda (Ecuador).
Panama). Photo: W. E. Duellman Photo: E. L. Neuschulz

Fig. 883. Lachesis melanocephala (San Vito, Fig. 886. Lachesis melanocephala (Tinamastes,
Costa Rica). Photo: R. W. Van Devender San Jose, Costa Rica). Photo: G. Kohler

Fig. 884. Lachesis stenophrys (Siquirres, Costa Fig. 887. Lachesis stenophrys (Guayacan de
Rica). Photo: R. D. Bartlett Siquirres, Costa Rica). Photo: G. Kohler
331
 Porthidium
The genus Porthidium consists of a group
of small (to 900 mm total length), slender,
terrestrial pitvipers, some of which have a
snout tip that points conspicuously upward
(these are commonly referred to as the
hognosed pitvipers). Recent revisions of
the genus Porthidium (CAMPBELL & LAMAR
1992, WERMAN 1992, MCDIARMID et al.
1999) have restricted the generic name to
the seven species (those listed below, plus
the poorly known P. hespere CAMPBELL
from the Mexican state of Colima). Despite
their modest body size, bites from the spe-
cies of this genus can be life-threatening to
humans.
The Porthidium species are generally cre-
puscular and nocturnal, but can be found
outside of their hiding places in shady
spots during the day. Their diet is made up
of frogs, lizards, blind snakes, and small
mammals, with the young including inver-
tebrates, such as earthworms (PORRAS et
al. 1981, LEE 1996). All Porthidium species
bear live young; litter size: P. nasutum 8-18
young (CAMPBELL 1998a), P. ophryomegas
19 young (LAWSEN 1997), P. yucatanicum 4-
10 young (McCOY & CENSKY 1992). Sexual
dichromatism has been reported in P.
ophryomegas, with the ground color being
gray in males and tan to grayish brown in
females (aside from differences in color
pattern; see SOLORZANO et al. 1988, LAWSEN
1997).
P. ophryomegas
•P. nasutum
• P. yucatanicum
• P. dunni
T P. lansbergii
• P. volcanicum
-Y P. porrasi
Porthidium
332
Porthidium dunni (HARTWEG & OLIVER 1938),
Occ. Pap. Mus. Zool. Univ. Michigan 390: 6;
type locality: Tehuantepec, Oaxaca, Mexico.
TL to 540 mm. Southwestern Oaxaca to
western Chiapas, Mexico, sea level to 500 m
elevation in dry forest.
Porthidium lansbergii (SCHLEGEL 1841), Mag.
Zool. Rept. (1-3), pl. 1; type locality: Turbaco,
Colombia. TL to 900 mm. Pacific side of
eastern Panama, as well as northern South
America, sea level to 1270 m elevation in dry
forest and wet forest.
Porthidium nasutum (BOCOURT 1868), Ann.
Sci. Nat. Paris (5) 10: 202; type locality:
Panz6s, on banks of Rio Polochic, Guatemala.
TL to 600 mm. Northwestern Chiapas,
Mexico, to northwestern Ecuador, sea level to
900 m elevation in rain forest
Porthidium ophryomegas (BOCOURT 1868),
Ann. Sci. Nat. Paris (5) 10: 201; type locality:
Warm regions on western [actually southern]
slope of mountains at Escuintla, Guatemala.
TL to 770 mm. South central Guatemala to
Costa Rica (occurrence in Panama question-
able), sea level to 1000 m elevation in dry
forest and savanna.
Porthidium porrasi LAMAR & SASA 2003, Rev.
Biol. Trop. 51: 799; type locality: "Drake Bay,
Peninsula de Osa, Puntarenas Province,
Costa Rica". TL to 385 mm. Golfo Dulce regi-
on inlcuding the Peninsula de Osa,
Puntarenas Province, Costa Rica, sea level to
about 500 m elevation in wet forest.
Porthidium volcanicum SoL6RZANO 1994, Rev.
Biol. Trop. 42: 696; type locality: Ujarras de
Buenos Aires (Valle de General) in the south-
west of Puntarenas Province, Costa Rica. TL
to 259 mm. Valle de General in the southwest
of Puntarenas Province, Costa Rica, 400-450
m elevation in dry forest and wet forest.
Porthidium yucatanicum (SMITH 19410,
Zoologica 26: 62; type locality: Chichen ltza,
Yucatan, Mexico. TL to 550 mm. The northern
portion of the Yucatan Peninsula, sea level to
250 m elevation in dry forest.
 Fig. 890. Porthidium nasutum (Cerro Saslaya,
Atlantico Norte, Nicaragua). Photo: G. Kohler

Fig. 888. Porthidium dunni (Arriaga, Chiapas,

Mexico). Photo: A. Ramirez V.

Fig. 891. Porthidium nasutum (Cerro Saslaya,

Nicaragua). Photo: A. Quintana

Fig. 889. Porthidium lansbergii. Fig. 892. Porthidium ophryomegas (El Rosario,
Photo: T. Leenders Zacapa, Guatemala). Photo: G. Kohler
333
 Scalation Characters

Key to Porthidium
1 a Snout tip points conspicuously upward
(Fig. 895) ........ ......................................... 2
b Snout tip does not point conspicuously
upward (Fig. 896) ....................... ........... 3
2 a Usually 23 dorsal scale rows at midbody;
underside of tail tip dark in adults; dorsal
blotches usually juxtaposed on either side
of the vertebral line ................... ............... ..
............................... Porthidium nasutum Fig. 895. Porthidium nasutum .
Drawing: M. Vesely
b Usually 25-27 dorsal scale rows at midbo-
dy; underside of tail tip pale in adults; dor-
sal blotches usually arranged opposite
each other on either side of the vertebral
line ........ ..... .. ......... Porthidium porrasi
3 a Snout tip moderately upturned ........ ....... 4
b Snout not or only slightly upturned ..... .... 5
4 a 21-23 dorsal rows at midbody, fewer than
21 dorsal rows one headlength anterior of
cloaca .................. ........ Porthidium dunni
b 23-27 dorsal rows at midbody; 21 dorsal
rows one headlength anterior to cloaca ...... Fig. 896. Porthidium ophryomegas .
.. . . . .... .. ........ ... Porthidium yucatanicum Drawing: M. Vesely
5 a Two canthals per side ................................ .
........................ Porthidium ophryomegas
b One canthal per side .......... ...... .... ......... 5
6 a Ornate white head pattern; venter mostly
dark brown with small, irregular, pale late- Further Literature
ral spots .......... Porthidium volcanicum AMARAL 1929b, PORRAS et al. 1981, WILSON &
b No ornate white head pattern; venter tan MCCRANIE 1984, CAMPBELL & LAMAR 1989,
or cream with darker mottling .. .......... .. McCOY & CENSKY 1992, SOLORZANO 1994,
............................. Porthidium lansbergii CAMPBELL & LAMAR 2004, BRYSON et al. 2008

Fig. 893. Porthidium yucatanicum (Quintana Fig. 894. Porthidium yucatanicum (Dzibalchen,
Roo, Mexico). Photo: H. Bahena B. Campeche, Mexico). Photo: R. Cedeno V.
334
 Scalation Characters

Scalation Characters in Snakes

Data from WILSON & MEYER 1985, PEREZ-SANTOS & MORENO 1988, LEE 1996, ROZE 1996, CAMPBELL
1998, original descriptions (in poorly known species), JAY SAVAGE (pers. comm. 2000) and Kohler
unpubl. data. Sea snakes and blind snakes are not inlcuded. Abbreviations: Dorsals-MB (number
of dorsal rows at mid-body); Dorsals-RC (number of dorsal rows one head length anterior to cloa-
ca); k-post (dorsal scales keeled on posterior body); m (mostly).

~~
~~
?>~~
~
~ ~
..;# ..;~c ~
~ ~

~e~"""'""'~
c 4~ 4~ 4~ ~c """'e
~~ ""'(: ""'(: ""'(: c"" ~c~
Boidae
Boa constrictor 225-228 45-65 smooth 60-71 35-40 undivided
Corallus annulatus 251-269 76-88 smooth 50-57 25-30 undivided
Corallus ruschenbergerii 250-272 94-115 smooth 38-48 25-30 undivided
Epicrates cenchria 233-238 55-58 smooth 46-53 22-25 undivided
Loxocemus bicolor 234-270 39-52 smooth 31-35 23-26 divided
Trachyboa boulengeri 132-152 21-33 keeled 31-33 23-25 undivided
Ungaliophis continentalis 204-258 39-47 smooth 25 15 undivided
Ungaliophis panamensis 226-254 41-48 smooth 19-23 17 undivided

Colubridae
Adelphicos daryi 120-132 19-28 smooth 15 15 divided
Adelphicos ibarrorum 135-138 21-26 smooth 15 15 divided
Adelphicos latifasciatus 125-138 37-51 smooth 15 15 divided
Adelphicos nigrilatus 113-135 21-36 smooth 15 15 divided
Adelphicos quadrivirgatus 117-155 29-50 smooth 15 15 divided
Adelphicos veraepacis 120-142 24-41 smooth 15 15 divided
Amastridium sapperi 144-170 79-85 smooth 17 17 divided
Amastridium veliferum 111-134 69-86 smooth 17 17 divided
Atractus clarki 181 33 smooth 17 17 undivided
fl1
Atractus darienensis 159 25 smooth 17 17 undivided ~
~
Atractus depressiocellus 167 30 smooth 17 17 undivided tU
Atractus hostilitractus
Atractus imperfectus
160 27 smooth
smooth
17
17
17 undivided =
00.
Chapinophis xanthocheilus 178-196 29-40 smooth 17 17 divided
Chironius carinatus 143-165 109-135 keeled 12 8 divided
Chironius exoletus 123-162 111-160 keeled 12 8 variable
Chironius grandisquamis 151-167 129-155 keeled 10 8 variable
Clelia clelia 198-247 57-93 smooth 19 17 undivided
Clelia equatoriana 202-217 56-80 smooth 17 17 undivided
Clelia scytalina 203-221 75-90 smooth 17 17 undivided
Coluber constrictor 151-172 78-108 smooth 17 15 divided
Coniophanes alvarezi 134-143 54-64 smooth 19 17 divided
Coniophanes bipunctatus 124-145 72-100 smooth 21 17-19 divided
Coniophanes fissidens 109-146 57-103 smooth 19-21 17 divided
Coniophanes imperialis 114-141 62-94 smooth 19 15-17 divided
Coniophanes joanae 131-132 49-53 smooth 17 15 divided
Coniophanes meridanus 120-135 78-90 smooth 17 15 divided
335
 Scalation Characters

~'(,~ti)
?>~ti)
~~ ~
~ ~
~~ ~~~c~
~
~~ ~~ ~~ ~" ~e
4e~ ~~" ~o'(. ~o'(. ~o'(. c"-" ~"~
Coniophanes piceivittis 153-174 78-115 smooth 23-25 19-21 divided
Coniophanes quinquevittatus 157-164 63-70 smooth 21-23 17-21 divided
Coniophanes schmidti 153-175 83-115 smooth 23-25 19 divided
Conophis lineatus 155-178 56-80 smooth 19 17 divided
Conophis vittatus 149-181 55-76 smooth 19 17 divided
Crisantophis nevermanni 173-183 71-89 smooth 19 17 divided
Dendrophidion nuchale 153-175 132-163 keeled 17 15 variable
Dendrophidion paucicarinatum 179-195 119-139 keeled 17 15 variable
Dendrophidion percarinatum 153-169 133-164 keeled 17 15 divided
Dendrophidion vinitor 148-165 111-128 keeled 17 15 variable
Diaphorolepis wagneri 193-197 133-138 keeled 19 17 undivided
Dipsas articulata 196-214 108-135 smooth 15 15 undivided
Dipsas bicolor 186-199 111-129 smooth 15 15 undivided
Dipsas brevifacies 162-180 70-101 smooth 15 15 undivided
Dipsas maxillaris 180 84 smooth 15 15 undivided
Dipsas temporalis 206-208 125-132 smooth 15 15 undivided
Dipsas tenuissima 225 128 smooth 15 15 undivided
Dipsas nicholsi 198-208 92-98 smooth 15 15 undivided
Dipsas viguieri 190-203 102-127 smooth 15 15 undivided
Drymarchon corais 188-215 56-88 smooth 17 13-15 undivided
Drymobius chloroticus 151-171 107-125 keeled 17 15 divided
Drymobius margaritiferus 137-158 103-138 keeled 17 15 divided
Drymobius melanotropis 150-161 91-108 keeled 17 15 divided
Drymobius rhombifer 148-163 84-102 keeled 17 15 divided
Elaphe fl,avirufa 245-269 96-122 k-post 25-31 19-25 divided
Enuliophis sclateri 130-155 88-109 smooth 15 15 divided
Enulius bifoveatus 168-181 100-120 smooth 17 15 divided
Enulius fl,avitorques 165-216 85-121 smooth 17 (15-)17 divided
00
Enulius roatanensis 165-176 103-121 smooth 17 17 divided
Erythrolamprus bizona 181-201 47-60 smooth 15 15 divided
=
~
Erythrolamprus mimus 171-199 42-51 smooth 15 15 divided
rl
I'll Ficimia publia 127-157 26-44 smooth 17 17 divided
Ficimia ramirezi 136 38 smooth 17 17 divided
Geophis bellus 131 33 k-post 15 15 undivided
Geophis brachycephalus 119-153 30-51 keeled 15 15 undivided
Geophis cancellatus 171 21-23 smooth 15 15 undivided
Geophis carinosus 120-134 41-48 k-post 17 17 undivided
Geophis championi 125-130 23-33 smooth 15 15 undivided
Geophis damiani 136-143 34-41 smooth 15 15 undivided
Geophis downsi 122-133 41-46 k-post 15 15 undivided
Geophis dunni 140 36 keeled 17 17 undivided
Geophis fulvoguttatus 135-157 34-36 smooth 17 17 undivided
Geophis godmani 132-145 26-36 smooth 15 15 undivided
Geophis hoffmanni 117-135 24-37 smooth 15 15 undivided
Geophis immaculatus 130 29 smooth 17 17 undivided
Geophis laticinctus 175 33 smooth 15 15 undivided
Geophis nasalis 115-142 23-37 keeled 17 17 undivided
336
 Scalation Characters

~.;,
~'(,'b' CJ'b'
~.;,
~'bl ~.;,
i;,'b'
~<e-# i;,'b'~<e~cfb-CJ'b'~ ~e
i;,'b'
40~ ~~ ~o~ ~o~ ~o~ c'" ~CJ~
Geophis nephodrymus 120-138 22-32 smooth 17 17 undivided
Geophis rhodogaster 129-147 29-48 smooth 17 17 undivided
Geophis ruthveni 123-135 32-41 k-post 15 15 undivided
Geophis talamancae 138 33 k-post 15 15 undivided
Geophis zeledoni 141-150 37-46 smooth 15 15 undivided
Hydromorphus concolor 171-180 31-52 smooth 17 15 divided
Hydromorphus dunni 164 52 smooth 15 13 undivided
Imantodes cenchoa 228-261 134-178 smooth 17 17 divided
Imantodes gemmistratus 195-262 106-155 smooth 17 17 divided
Imantodes inornatus 196-218 110-132 smooth 17 17 variable
Imantodes phantasma 235-236 156-161 smooth 17 (15-)17 divided
Imantodes tenuissimus 242-246 142-146 smooth 17 17 divided
Lampropeltis triangulum 205-244 39-63 smooth 19-23 17-19 undivided
Leptodeira annulata 156-184 54-89 smooth 21-25 15-19 divided
Leptodeira frenata 170-192 64-86 smooth 21-23 13-17 divided
Leptodeira nigrofasciata 161-196 54-76 smooth 19 15-17 divided
Leptodeira rubricata 177-182 82-97 smooth 21-23 17-19 divided
Leptodeira septentrionalis 174-211 72-107 smooth 19-25 11-17 divided
Leptodrymus pulcherrimus 195-210 145-152 smooth 17 15 divided
Leptophis ahaetulla 150-183 137-185 keeled 15 11 divided
Leptophis depressirostris 144-158 158-170 keeled 15 11 divided
Leptophis diplotropis 167-184 134-161 keeled 15 11-13 divided
Leptophis mexicanus 145-174 140-181 keeled 15 11 divided
Leptophis modestus 168-183 166-186 keeled 13 11 divided
Leptophis nebulosus 150-160 146-151 keeled 15 11 divided
Leptophis riveti 133-149 135-145 keeled 15 11 divided
Liophis epinephelus 128-152 49-78 smooth 17 15 divided
Liophis lineatus 159-179 70-97 smooth 19 15-17 divided
Manolepis putnami 167-181 57-82 smooth (17-)19 15 divided rn
Masticophis mentovarius 166-205 95-126 smooth 17 13 divided QI
~
Mastigodryas dorsalis 177-196 109-137 smooth 17 15 divided cu
Mastigodryas melanolomus 163-195 85-136 smooth 17 15 divided r::
00
Mastigodryas pleei 171-191 83-105 smooth 17 15 divided
Nerodia rhombifera 135-142 64-86 keeled 25-31 21-25 divided
Ninia atrata 133-156 39-70 keeled 19 19 undivided
Ninia diademata 123-159 73-106 keeled 19 19 undivided
Ninia celata 123-127 33-45 keeled 19 19 undivided
Ninia espinali 139-157 49-63 keeled 19 19 undivided
Ninia maculata 125-155 44-60 keeled 19 19 undivided
Ninia pavimentata 130-148 63-76 keeled 19 19 undivided
Ninia psephota 139-163 51-77 keeled 17 17 undivided
Ninia sebae 130-156 40-74 keeled 19 19 undivided
Nothopsis rugosus 146-162 81-105 keeled 26-28 22-26 undivided
Omoadiphas aurula 162-170 24-39 smooth 17 17 divided
Omoadiphas texiguatensis 172 47 smooth 17 17 divided
Oxybelis aeneus 173-205 137-203 keeled 17 13 divided

337
 Scalation Characters

Oxybelis brevirostris 166-186 159-179 keeled 15 13 undivided

Oxybelis fulgidus 198-217 139-186 keeled 17 13 divided
Oxybelis wilsoni 201-217 164-189 keeled 17 13 divided
Oxyrhopus petola 193-244 79-117 smooth 19 17 undivided
Phimophis guianensis 168-204 53-72 smooth 19 17 undivided
Pituophis lineaticollis 234-246 57-65 keeled 27-37 19-27 undivided
Pliocercus elapoides 122-144 85-115 smooth 17 17 divided
Pliocercus euryzonus 120-137 97-121 smooth 17 17 divided
Pseudoboa neuwiedii 177-198 64-96 smooth 19 17 undivided
Pseustes poecilonotus 181-220 95-145 keeled 19-25 13-15 undivided
Rhadinaea anachoreta 139-147 74-80 smooth 17 17 divided
Rhadinaea calligaster 141-156 46-68 smooth 17 17 divided
Rhadinaea decorata 110-134 67-122 smooth 17 17 divided
Rhadinaea godmani 156-186 71-95 smooth 21 21 divided
Rhadinaea hannsteini 140-152 62-75 smooth 17 17 divided
Rhadinaea hempstedae 156-175 79-101 smooth 19 19 divided
Rhadinaea kanalchutchan 159-169 71-90 smooth 19 19 divided
Rhadinaea kinkelini 136-163 70-83 smooth 17 17 divided
Rhadinaea lachrymans 154-183 61-89 smooth 17 17 divided
Rhadinaea macdougalli 119-131 60-74 smooth 17 17 divided
Rhadinaea montecristi 157-176 71-87 smooth 19 19 divided
Rhadinaea pilonaorum 151-164 98 smooth 17 17 divided
Rhadinaea pegosalyta 163 86 smooth 21 21 divided
Rhadinaea posadasi 136-146 86-95 smooth 17 17 divided
Rhadinaea pulveriventris 119-134 63-80 smooth 17 17 divided
Rhadinaea rogerromani 161 96 smooth 21 21 divided
Rhadinaea sargenti 117-122 67-74 smooth 17 17 divided
Rhadinaea serperaster 156-172 66-79 smooth 19 19 divided
Rhadinaea stadelmani 169-191 74-94 smooth 19-21 17-19 divided
Rhadinaea tolpanorum 157-164 68-73 smooth 17 17 divided
Rhadinaea vermiculaticeps 117-121 63-82 smooth 17 17 divided
Rhinobothryum bovallii 239-246 115-125 k-post 19 17 divided
Salvadora lemniscata 196-208 118-146 smooth 17 13 divided
Scaphiodontophis annulatus 132-166 110-145 smooth 17 17 divided
Scaphiodontophis venustissimus 92-156 107-121 smooth 17 17 divided
Scolecophis atrocinctus 181-198 45-54 smooth 15 15 divided
Senticolis triaspis 241-282 87-126 k-post 27-39 21-25 divided
Sibon annulatus 161-192 108-133 smooth 15 15 undivided
Sibon anthracops 166-188 73-91 smooth 13 13 undivided
Sibon argus 181-201 112-121 smooth 15 15 undivided
Sibon carri 157-173 41-51 smooth 13 13 undivided
Sibon dimidiatus 175-200 106-126 smooth 15 15 undivided
Sibon lamari 162-171 77-119 smooth 15 15 undivided
Sibon longifrenis 151-173 82-106 smooth 15 15 undivided
Sibon manzanaresi 171-176 106-112 smooth 15 15 undivided
Sibon miskitus 168-172 95-112 smooth 15 15 undivided

338
 Scalation Characters

~,,,/~~ ~~
~~~
~~ ~~
# #
~...~ ~
Ci ~~ ~~ ~ ~Ci ~e
4e~ 't:>~ ~()~ ~o~ ~o~ c'o <t:>ei~
Sibon nebulatus 159-200 64-114 smooth 15 15 undivided
Sibon sanniolus 143-162 57-88 smooth 15 15 undivided
Siphlophis ceruinus 223-264 105-128 smooth 19 15-17 undivided
Spilotes pullatus 198-241 100-142 keeled 14-18 10-14 undivided
Stenorrhina degenhardtii 136-159 31-48 smooth 17 17 divided
Stenorrhina freminuillei 160-182 25-42 smooth 17 17 divided
Storeria dekayi 112-148 36-69 keeled 15-17 15-17 divided
Symphimus leucostomus 162-181 106-122 smooth 15 15 divided
Symphimus mayae 150-165 115-146 smooth 15 15 divided
Tantilla albiceps 183 65 smooth 15 15 divided
Tantilla alticola 128-145 32-60 smooth 15 15 divided
Tantilla armillata 155-177 46-60 smooth 15 15 divided
Tantilla bairdi 163-164 34-36 smooth 15 15 divided
Tantilla breuicauda 139-155 21-26 smooth 15 15 divided
Tantilla cuniculator 139-154 48-58 smooth 15 15 divided
Tantilla hendersoni 153 64 smooth 15 15 divided
Tantilla impensa 162-171 65-72 smooth 15 15 divided
Tantilla jani 144-147 44-47 smooth 15 15 divided
Tantilla lempira 138-154 36-52 smooth 15 15 divided
Tantilla moesta 138-152 52-62 smooth 15 15 divided
Tantilla reticulata 158-173 58-70 smooth 15 15 divided
Tantilla rubra 148-162 59-68 smooth 15 15 divided
Tantilla ruficeps 139-155 59-83 smooth 15 15 divided
Tantilla schistosa 117-147 24-42 smooth 15 15 divided
Tantilla supracincta 138-151 52-65 smooth 15 15 divided
Tantilla taeniata 143-161 62-70 smooth 15 15 divided
Tantilla tayrae 140-154 44-51 smooth 15 15 divided
Tantilla tecta 148 54 smooth 15 15 divided
Tantilla triseriata 163-165 63 smooth 15 15 divided fll
Tantilla tritaeniata 155-161 59-65 smooth 15 15 divided Q)
~
Tantilla uermiformis
Tantilla uulcani
115-129
139-154
19-27
37-51
smooth
smooth
15
15
15
15
divided
divided
=
=
00
Tantillita breuissima 116-126 28-34 smooth 15 15 divided
Tantillita canula 103-114 32-44 smooth 15 15 divided
Tantillita lintoni 103-115 43-56 smooth 15 15 divided
Thamnophis cyrtopsis 144-166 75-101 keeled 19 17 undivided
Thamnophis fuluus 132-154 56-80 keeled 19 15-17 undivided
Thamnophis marcianus 134-173 56-82 keeled 19 17 undivided
Thamnophis proximus 141-181 82-131 keeled 19 17 undivided
Tretanorhinus nigroluteus 127-151 56-82 keeled 21 17-19 divided
Tretanorhinus mocquardi 166-177 69-85 keeled 19 17 divided
Trimetopon barbouri 138-153 58 smooth 15 15 divided
Trimetopon gracile 141-153 60-69 smooth 15 15 divided
Trimetopon pliolepis 141-154 59-73 smooth 17 17 divided
Trimetopon simile 122-131 69 smooth 15 15 divided
Trimetopon sleuini 155-161 45-53 smooth 17 17 divided

339
 Scalation Characters

~'<-~~ ~~
~~~
~~
#'
~~ ~~~
#
~ 4~ ~ ~c ~~
~~~ "
~~ -Q"'(. -Q"'(. -Q"'(. c'" ~c~
Trimetopon viquezi 161 <40- smooth 17 17 divided
Trimorphodon biscutatus 232-286 77-105 k-post 23-27 15-19 divided (m)
Trimorphodon quadruplex 249-266 73-99 k-post 23-27 15-19 divided
Tripanurgos compressus 228-258 110-125 smooth 19 15 undivided
Tropidodipsas fasciatus 172-200 58-87 smooth 17 17 undivided
Tropidodipsas fischeri 167-194 43-82 keeled 17 17 undivided
Tropidodipsas sartorii 165-197 52-77 keeled 17 17 undivided
Urotheca decipiens 122-143 90-121 smooth 17 17 divided
Urotheca fulviceps 136-143 98-122 smooth 17 17 divided
Urotheca guentheri 135-176 82-110 smooth 17 17 divided
Urotheca myersi 132-138 120 smooth 17 17 divided
Urotheca pachyura 130-138 104-124 smooth 17 17 divided
Xenodon rabdocephalus 124-153 35-52 smooth 19 17 undivided

Elapidae
Micrurus alleni 211-240 32-60 smooth 15 15 divided
Micrurus ancoralis 244-286 30-37 smooth 15 15 divided
Micrurus bogerti 214-230 38-56 smooth 15 15 divided
Micrurus browni 204-230 36-58 smooth 15 15 divided
Micrurus clarki 190-221 34-58 smooth 15 15 divided
Micrurus diastema 188-227 32-57 smooth 15 15 divided
Micrurus dissoleucus 182-211 19-27 smooth 15 15 divided
Micrurus elegans 202-226 32-49 smooth 15 15 divided
Micrurus hippocrepis 196-226 37-56 smooth 15 15 divided
Micrurus latifasciatus 186-212 37-56 smooth 15 15 divided
Micrurus mipartitus 242-281 23-33 smooth 15 15 divided
Micrurus multifasciatus 235-311 24-38 smooth 15 15 divided
Micrurus nigrocinctus 182-230 31-53 smooth 15 15 divided
Micrurus ruatanus 178-203 34-48 smooth 15 15 divided
00.
=
~
ts;"
Micrurus stewarti
Micrurus stuarti
200-228
210-231
36-55
37-49
smooth
smooth
15
15
15
15
divided
divided
ID
f/J
Viperidae
Agkistrodon bilineatus 129-144 50-68 keeled 23 19 undivided
Atropoides nummifer 114-135 22-39 keeled 23-31 17-23 undivided
Atropoides picadoi 138-155 30-40 keeled 23-29 19-21 undivided
Bothriechis aurifer 148-167 48-65 keeled 18-21 undivided
Bothriechis bicolor 156-175 57-75 keeled 21 15-17 undivided
Bothriechis lateralis 155-171 54-70 keeled 21-23 undivided
Bothriechis marchi 156-174 55-69 keeled 19-21 15-17 undivided
Bothriechis nigroviridis 134-158 44-58 keeled 17-21 undivided
Bothriechis rowleyi 154-166 53-66 keeled 19-21 undivided
Bothriechis schlegelii 137-169 42-64 keeled 21-25 17-21 undivided
Bothriechis supraciliaris 141-150 45-54 keeled 21-23 17-21 undivided
Bothriechis thalassinus 161-168 60-67 keeled 21-23 15-17 undivided
Bothrops asper 185-220 56-70 keeled 23-25 19-21 undivided

340
 Snakebite

Snakebite - Prevention and First Aid

Snakes in the tropics pose a health risk

that should not be overlooked. Life-threat-
ening snakebites in Central America occur
most commonly among natives working in
the field, with the highest mortality report-
ed in incidents involving pitvipers (partic-
ularly Bothrops asper) (D. HARDY 1994).

Taking appropriate precautions can dimin-

ish the risk of snakebite. A study of the
incidence of snakebite amongst field biolo-
gists provides evidence that the risk of
being bitten by a pitviper, even in areas
with a high population of these snakes, is
slight. The use of proper equipment and
procedures within snake habitat will en-
sure safety. Fig. 898. Bothrops asper (Nicaragua).
Photo: G. Kohler

Safety Precautions

• Suitable clothing: long, loose-fitting
pants and sturdy footwear, preferably
rubber boots. A large pitviper, however,
can bite through rubber boots.
• One should only proceed through areas
where it is clear that there are no
snakes; do not walk through dense bush
or high grass. Likewise, be certain there
are no snakes under or behind fallen
logs before stepping over them.
• Only hold onto tree trunks or branches
once one has determined that there are
no snakes present. Unless it moves, a
pitviper can be virtually undetectable in
the twilight of the rain forest, due to its
cryptic coloration. Snakes reflect light
quite easily, and at night can be spotted
with a flashlight from a distance of
several meters.

• When suddenly faced with a snake, it is

important to remain as still as possible,
as snakes have relatively poor eyesight
and will mostly strike in the direction of
a moving object; sudden movement must
absolutely be avoided. One should then
calmly withdraw.

Fig. 897. Difficult to spot: a Bothrops asper

(Saslaya, Nicaragua). Photo: G. Kohler
342
 Possible Resting Places of Pitvipers

Fig. 899. Possible resting places of pitvipers in

the rain forest:
Bothriechis schlegelii and Bothriechis bicolor in
the vegetation, Bothrops asper on the floor.
Photos: G. Kohler
 Snakebite
Recommendations for First Aid
The elaboration of a snakebite protocol can
improve the outcome of a venomous snake-
bite in the field. This protocol should at
least include (liARDY 1994):
• First aid measures to be used and those
which will not be used.
• Route and means of transportation for
evacuation; name and location of nearest
medical facility.
• List of medical consultation resources
with expertise in snakebite treatment
including locations and telephone num-
bers both in country and overseas.
The psychological reaction of the snakebite
victim will greatly affect the out-come of
envenomation. The victim should be encour-
aged to remain calm and avoid panic.
Under no circumstances should a person
who has been bitten be left alone. The
affected extremity should be immobilized
(arm in a sling, leg splinted, as appropria-
te), in order to slow the spread of the
venom. Rings and bracelets should be
immediately removed to avoid cutting off
the circulation as swelling occurs. When-
ever possible, the snake should be identi-
fied in order to determine specific therapy.
Capture and killing of the animal should
only be undertaken if no threat is posed to
the victim or helper. Do not handle a see-
mingly dead snake. Snakes are experts at
feigning death. They may even twist them-
selves upside down and lie with their
mouth open and the tongue lolling out.
Also, a freshly killed snake often still has
some kind of biting reflexes and might be
able to deliver bites. The snakebite victim
needs to be brought to the nearest point of
medical treatment as quick as possible. At
any time, respiration, blood pressure, and
consciousness of the victim needs to be
monitored. A brief outline of the circum-
stances of the accident, the progress of the
envenomation, information about the vic-
tim, any evidence, personal observations,
and measures taken can be helpful for
later treatment. Due to possible complicati-
ons, therapeutic measures should only be
undertaken by a qualified physician.
344
Antivenin is the only specific therapy for
venomous snakebite. Laymen often have
the impression that once the antivenin has
been administered, the patient is out of
danger. However, the use of antivenin car-
ries its own risk. Occasionally, a patient
may develop symptoms of life-threatening
anaphylactic shock: facial swelling, respi-
ratory distress, rapid drop in blood pressu-
re, and pallor.
Various snakebite kits (e.g., "Sawyer
Extractor", "Aspivenin") are available in
the trade. They have been proven useless
under field conditions (D. MEBS pers.
comm. 2000). Some extractors can lead to
injuries and actually be dangerous: in
situations where the venom produces an
anticoagulant effect on the blood, hemor-
rhage can occur. There is a popular mis-
conception that the venom can be somehow
removed by removing the surrounding tis-
sue (e.g., by amputating a finger or making
an incision in the skin). Most surgical
interventions for snakebite actually cause
further injury and either delay or entirely
prevent recovery (MEBS 2000). Under no
circumstances should procedures, such as
incision, excision, squeezing, or applying suc-
tion, be performed on the wound site (see
also snakebite kits). Such measures often
cause damage to the larger vessels and
cause the venom to spread more rapidly
through the circulatory system. There is
also the danger of secondary infection, and
tissue damage can lead to hemorrhage if
the venom has an anticoagulant effect. The
venom of most Central American pitvipers
has both pro-coagulant and anti-coagulant
properties, which lead to the consumption
of the blood's clotting factors (in a process
known as consumption coagulopathy or dis-
seminated intravascular clotting). In
humans, the fibrinolytic system is activa-
ted to prevent clotting, but once the blood
clotting factors are used up, the blood beco-
mes incoagulable. If hemorrhagic factors
(e.g., enzymes that perforate the basal
membranes of blood vessels) are present in
the venom, this can lead to spontaneous
major internal and external bleeding.
 Snakebite

Tying off the affected extremity is not

recommended, as this will disrupt the
blood flow and lead to tissue damage.
Extreme heat (cauterization) or cooling of
the wound site are equally detrimental.
Application or injection of home remedies,
administration of alcohol or coffee, or the
ingestion of food (which may cause vomit-
ing) are also contraindicated. The applica-
tion of a pressure bandage is not advisable,
as a great deal of swelling will result from
a viper bite and the circulation can be cut
off (D. MEBS pers. comm. 2000). Fig. 900. Conophis lineatus ; note the enlarged
tooth of this rear-fanged snake.
Photo: G. Kohler

Rear-fanged Snakes

Many colubrids produce a secretion in the

Duvernoy gland, located in the upper jaw,
that in some species is highly toxic. The
rear-fanged snakes have enlarged, grooved
teeth in the rear of the upper jaw (compare
Figs. 900 and 901), that enable them to
inject this secretion into their prey or into
potential predators. While some Old World
rear-fanged snakes are classified as dead-
ly (e.g., the boomslang, Dispholidus typus),
there have been no known fatalities from
envenomation by such snakes in Central
America (MINTON 1990). Fig. 901. Skull of Xenodon rabdocephalus;
the enlarged tooth in the upper jaw is clearly
evident. Photo: R. W. Van Devender
Yet, the rear-fanged snakes of Central
America cannot be considered entirely
harmless as certain species (e.g., Conio-
phanes spp., Conophis spp. , Pliocercus
spp., Xenodon spp.) produce a highly
potent venom, which causes not only local-
ized symptoms (pain and swelling) but
toxic systemic effects (headache and
persistent bleeding) (MINTON & MEBS
1978, GREENE 1997). In order to avoid bites
from rear-fanged snakes, these animals
should only be handled, if at all, with
leather gloves and protective, long-sleeved
clothing.

Further Reading
CAMPBELL & LAMAR 1989, MINTON 1990, D.
HARDY 1992, 1994, D. HARDY & SILVA HAAD Fig. 902. Xenodon rabdocephalus (Costa Rica).
1998, MEBS 1992, 2000 Photo: R. W. Van Devender
345
 Mimicry

Mimicry in Snakes
Mimicry is defined as deceptive imitation
by an organism with regard to form, color-
ation, or behavior, usually to frighten off
potential predators, but, in some cases, to
attract prey. Batesian mimicry (two similar
species, of which one, the model, is
venomous or inedible and the mimic is
non-venomous and edible) is differentiated
from Mullerian mimicry (both species are
venomous and are model and mimic of
each other). The concept of Mertensian
mimicry (a mildly venomous model, a high-
ly venomous and a non-venomous mimic)
has been dismissed as superfluous by some Fig. 903. A true coral snake (Micrurus hippo-
authors (GREENE & MCDIARMID 1981). crepis) from Belize. Photo: P. Stafford

In the latter half of the 19th century, re-

searchers had already noted the remark-
able similarity between venomous and
non-venomous coral snakes and certain
colubrids (COPE 1860d, WALLACE 1867,
1870). The possibility that the brightly
colored true coral snakes (approximately
50 species of the genera Micrurus and
Micruroides) and the many similarly col-
ored non-venomous or mildly venomous
colubrids of several genera (among others,
Erythrolamprus, Lampropeltis, Oxyrhopus,
and Pliocercus) are involved in a complex
mimicry system has created much contro-
versy (DUNN 1954, BRATTSTROM 1955, Fig. 904. Pliocercus elapoides (a specimen from
HECHT & MARIEN 1956, MERTENS 1956). No Belize) exhibits a very similar color pattern as
fewer than 115 species of colubrid (18% of several species of venomous coral snakes.
all American snakes) display elements of Photo: P. Stafford
coral snake coloration and might be con-
sidered as mimics of Micrurus (SAVAGE &
SLOWINSKI 1992). The coral snake mimicry
hypothesis states that the conspicuous
colors of the venomous species act as a
warning to potential predators and these
predators will also avoid similarly colored
non-venomous and mildly venomous snakes.
One of the primary arguments against this

Fig. 905. Erythrolamprus bizona has pairs of

black rings, whereas in the Central American
Micrurus species, the black rings are either
single or in triads. Photo: G. Kohler
346

hypothesis, that the true coral snakes
serve as a model for a sizable number of
colubrids, is that a potential predator
would not survive the bite of a Micrurus
and, thus, benefit from the acquired experi-
ence. GREENE & MCDIARMID (1981) sugges-
ted several possibilities for circumventing
this "deadly model", and report that the
bite of a Micrurus need not be deadly, even
for small vertebrates. In predators that
hunt in groups, an individual can learn to
avoid the coloration of coral snakes by
observing the fate of a conspecific that has
been bitten by a Micrurus. The possibility
of the innate avoidance of snakes with
bright red and black bands, which are the
warning colors of coral snakes, should not
be overlooked. Those individuals with a les-
ser innate inclination to attack coral snakes
have a higher probability of surviving, and,
therefore, a selective advantage over con-
specifics that are not deterred by coral
snake coloration. Contrary to previous
assertions that they are strictly nocturnal,
coral snakes (Micrurus and the bulk of
their supposed mimics) are also active
during the day, so that predators can easi-
ly spot their bright colors.
In recent literature, several important
arguments for mimicry among New World
snakes with black-red or black-yellow-red
markings have been published. Experi-
mental work relating to this phenomenon
has also been performed (GREENE &
MCDIARMID 1981, SAVAGE & SLOWINSKI
1992, BRODIE & JANZEN 1995, BRODIE &
MOORE 1995, WILSON et al. 1996). An exact
geographic correspondence exists among
several supposed model-mimic pairs
(GREENE & MCDIARMID 1981, WILSON et al.
1996). Purported mimics among colubrid
species with a broad range display conspic-
uous color variations that correspond to
their supposed models. This is particularly
notable in the various Pliocercus and
Erythrolamprus populations, whose colors
correspond remarkably to those of sympat-
ric Micrurus species.
Mimicry
The idea that Micrurus species are not just
mimicked, but themselves mimic certain
less venomous rear-fanged snakes
(Erythrolamprus, Pliocercus) needs to be
examined (MERTENS 1956). Despite plenti-
ful evidence that mimicry is the cause of
the above-named similarities between
many unrelated colubrid and Micrurus
species, less obvious explanations for this
type of color similarity should not be disre-
garded. The correspondence may be the
result of other environmental factors that
have nothing to do with mimicry, or it may,
in fact, be coincidence. Evidence contrary
to these alternative explanations is provid-
ed by the fact that the supposed models
and mimics inhabit different microhabi-
tats subject to different environmental fac-
tors yet they correspond exactly to the geo-
graphic variability of models and mimics
with a broad range. There are also several
documented cases of supposed model and
mimic pairs in which unusual color varia-
tions occur which are otherwise unknown
in their respective genera, as for example
spotted Micrurus and Pliocercus as well as
black Micrurus and Erythrolamprus.
Findings to date suggest that certain spe-
cies of Atractus, Erythrolamprus,
Lampropeltis, Micruroides, Micrurus and
Pliocercus and other genera are integrated
into a complex system of mimicry whose
dynamics and relationships have yet to be
investigated (GREENE & MCDIARMID 1981).
The phenomenon of mimicry is not limited
among snakes in Central America to the
above mentioned venomous and non-veno-
mous coral snakes. Even the highly
dangerous pitvipers (Bothrops and related
species) also have imitators, such as cer-
tain Sibon species as well as Leptodeira
frenata, and especially the "Falso
Terciopelo" (Xenodon rabdocephalus),
which at first glance appears quite similar
to the pitvipers.
Further Reading
HECHT & MARIEN 1956, MERTENS 1956, GREENE
& MCDIARMID 1981, WILSON et al. 1996, KUCH
1997
347
 Mimicry

Fig. 743. Scaphiodontophis annulatus (Finca Fig. 909. Rhinobothryum bovallii (Rio
La Giralda, La Libertad, El Salvador). Siquirres, Costa Rica). Photo: A Hohmeister
Photo: E. Greenbaum

Fig. 907. Erythrolamprus bizona (Rio Fig. 910. Erythrolamprus mimus (near Pueblo
Siquirres, Costa Rica). Photo: A Hohmeister Wiso, Jinotega, Nicaragua). Photo: G. Kohler

Fig. 908. Scolecophis atrocinctus (Nicaragua?). Fig. 911. Tantilla supracincta (Osa Peninsula,
Photo: R. W. Van Devender Costa Rica). Photo: R. W. Van Devender
348
 Mimicry

Fig. 912. Pliocercus euryzonus (Selva Negra, Fig. 915. Ninia sebae (albino; Cusuco,
Matagalpa, Nicaragua). Photo: G. Kohler Honduras. Photo: J. Kolby

Fig. 913. Tropidodipsas sartorii (near Fig. 916. Oxyrhopus petola (La Fortuna, Prov.
Escarcega, Campeche, Mexico). Photo: J . C. Lee Alajuela, Costa Rica).
Photo: R. W. Van Devender

Fig. 914. Ficimia publia (La Ceiba, Honduras). Fig. 917. Dipsas breuifacies (Yucatan, Mexico).
Photo: G. Kohler Photo: J. C. Lee

349
 Acknowledgments

Numerous people have assisted me in the to these people for their valuable help.
various phases of my work in Central Many students and associates of the
America over a more than 15-year period department of herpetology at Senckenberg
and contributed to this book project. Museum, Frankfurt, Germany, have tested
the identification keys included in this
I appreciate the cooperation I have re- book. Their help is greatly appreciated.
ceived from forestry and wildlife officials in
Central America. Collecting and exporta- I am grateful to numerous colleagues for
tion permits were provided by R. Rivera, A photos of Central American reptile
Salazar, Y. Hidalgo, and Sra. Cruzcaya, species-these photographers are acknow-
Autoridad N acional de Ambiente (ANAM),
Panama City, Panama; J. Guevara
Sequeira and R. Tenorio Jimenez,
Ministerio de Recursos Naturales, Energia
y Minas, San Jose, Costa Rica; M. Fonseca
Cuevas, S. Tijerino, M. G. Camacho, and C.
Peres-Roman, Ministerio del Ambiente y
los Recursos N aturales (MARENA),
Managua, Nicaragua; A. Barahona, A. P.
Martinez, T. Garcia, and C. Romero,
Corporaci6n Hondureiia de Desarollo
Foresta! (COHDEFOR), Tegucigalpa,
Honduras; L. R. Arevalo and A. Sanchez,
Ministerio de Agricultura y Ganaderia, El
Salvador; J.I. Galvez Duiiionez, Consejo
Nacional de Areas Protegidas (CONAP), Fig. 918. Expedition to Pico Bonito, Honduras
Guatemala City, Guatemala; F. Ramirez (March 1996). Left to right: Webb Munoz,
Ruiz de Velasco and L. Lozano, Secretaria Ernesto Rodriguez; Twan Leenders, Elke
de Medio Ambiente, Recursos Naturales y Kohler, Gunther Kohler. Photo: G. Kohler
Pesca, Mexico D.F., Mexico.

For logistic support for fieldwork I thank

Robert Dilger, Hauke Hoops, Siegfried
Kastl, Mausi and Eddi Kuhl, Juan Carlos
Martinez Sanchez, Ivan Ortega, Roger
Roman, and Francisco Serrano. For field
assistance, I thank Manuel E. Acevedo,
Abel Batista, Martin Dolling, Jorge
Ferrari, Martin Jansen, Elke Kohler, Jorg
Kreutz, James Randy McCranie, Fernando
Mendoza, Kirsten Nicholson, Michael
Obermeier, Marcos Ponce, Ardiel Z.
Quintana, Antonio Ramirez Vasquez,
Fig. 919. Return from Quebrada Grande,
Fabian Schmidt, Robert Seipp, Jorge Soto, Honduras (October 1998). Left to right: James
Pablo Soto, Javier Sunyer, and Jose Randy McCranie, Elke Kohler, Gunther
Manuel Zolotoff. I am especially thankful Kohler. Photo: G. Kohler

350
 Acknowledgments

Fig. 920. Manuel E. Acevedo (left) and Gunther Kohler in the highlands of Guatemala (June
2005). Photo: E. Kohler

Fig. 921. Field work in Panama. Left to right: Abel Batista, Marcos Ponce, Gunther Kohler, Jose
Vega. Photo: E. Kohler
351
 Acknowledgments
ledged in the captions of their respective
photos. Many of the excellent pen and ink
drawings are the painstaking efforts of
Lara Czupalla and Milan Vesely.
For the loan of specimens or access to the
collections under their care I thank Linda
Ford and Darrel R. Frost, American
Museum of Natural History (AMNH), New
York City; Ted Daeschler, Academy of
Natural Sciences of Philadelphia (ANSP),
Philadelphia; Colin J. McCarthy, The
Natural History Museum (BMNH),
London; Jens Vindum, California Academy
of Sciences (CAS), San Francisco; Alan
Resetar and Harold Voris, Field Museum of
Natural History (FMNH), Chicago; John E.
Simmons, Eli Greenbaum, Linda Trueb,
and William E . Duellman, University of
Kansas Natural History Museum (KU),
Lawrence; Jeff A. Seigel, Natural History
Museum of Los Angeles County (LACM),
Los Angeles; Douglas A. Rossman and
Frank T. Burbrink, Museum of Natural
Science, Louisiana State University
(LSUMZ), Baton Rouge; Jose Rosado and
James Hanken, Museum of Comparative
Zoology (MCZ), Harvard; Alain Dubois,
Ivan Ineich and Annemarie Ohler,
Museum National d'Histoire Naturelle
(MNHN), Paris; Eunice E. Echeverria,
Museo de Historia Natural de El Salvador
(MUHNES), San Salvador; David B. Wake
and Carla Cicero, Museum of Vertebrate
Zoology (MVZ), Berkeley; Franz
Tiedemann, Naturhistorisches Museum
Wien (NMW), Wien; Janalee P. Caldwell,
Amy D. Estep, and Laurie J . Vitt, Sam
Noble Oklahoma Museum of Natural
History (OMNH), Norman; James R. Dixon
and Katherine Vaughan, Texas
Cooperative Wildlife Collection CTCWC),
College Station; F. Wayne King and
Kenneth Krysko, Florida Museum of
Natural History (UF), Gainesville;
Christopher A. Phillips, John Petzing, and
Steven D. Sroka, Museum of Natural
History, University of Illinois at Urbana
(UIMNH), Urbana; Ronald Nussbaum,
Arnold Kluge, and Greg Schneider,
University of Michigan Museum of Zoology
352
(UMMZ), Michigan; Ronald Crombie,
Ronald Heyer and Robert Wilson,
Smithsonian Institution, National
Museum of Natural History (USNM),
Washington DC; Jonathan Campbell,
University of Texas at Arlington (UTA),
Arlington; Victor J. Hellebuyck (VH), San
Salvador; Wolfgang Bohme, Zoologisches
Forschungsinstitut und Museum Alexan-
der Koenig (ZFMK), Bonn; and Rainer
Gunther, Museum fur Naturkunde der
Humboldt-Universitat zu Berlin CZMB),
Berlin; Frank Glaw and Dieter Fuchs,
Zoologische Staatssammlung Munchen
CZSM), Munich.
I owe an immense debt of gratitude to AJ
Gutman, James Randy McCranie, and
Larry David Wilson, who patiently sup-
ported this project and read the various
stages of the manuscript; the final manu-
script version was also read by Helmut
Diethert, Peter Stafford, and Harlan
Walley; all individuals and provided valu-
able comments and advice that signifi-
cantly improved the text. Manuel E .
Acevedo, Antonio Ramirez Vasquez, Jay M.
Savage and Hobart M. Smith have
reviewed and improved the distribution
maps for the species included in this book.
I especially wish to thank AJ Gutman who
translated the major part of the text from
German to English. James Randy
McCranie, Charles W. Myers, Jay M.
Savage, and Larry David Wilson generous-
ly shared their extensive knowledge on
Central American herpetology with me
and provided important unpublished data.
Special thanks are due to my friend and
mentor, Alfred A. Schmidt, for his constant
support and encouragement.
Finally, I am especially grateful to my
beloved wife Elke Kohler for her longtime
emotional support and inspiring encour-
agement. Elke often accompanied me in
the field and provided valuable assistance.
She also merits special thanks for her very
careful editing of the manuscript. My in-
debtedness to her cannot be exaggerated.
Glossary
abdominal scute (plate) in turtles, a paired
scale located on the plastron between the
pectoral scale and the femoral scale (Fig.
32c)
anal spur anal claw; in boids two claw-like
structures on either side of the cloaca,
which form the externally visible remnants
of rudimentary hind limbs
anal scale the hindmost pair of scales on the
plastron of turtles (Fig. 32c); in lizards and
snakes, a single or divided scale covering
the cloacal opening (Fig. 532)
apical at the tip (apex)
arboreal living in trees
aridization the process of climatic drying
arthropods animals with jointed appendages
(insects, spiders, crustaceans)
autochthon evolved in a particular location
autotomy ability to drop body parts (e.g. tail)
as a protective measure
axillary scute a small scale on either side of a
turtle shell between the bridge and the pec-
toral scute (Fig. 32c)
axillary pocket tube-shaped pocket of skin in
the shoulder area (Fig. 350)
basal located at the base
basis foundation, starting point
biotope habitat
bridge in turtles, the segment joining the
carapace and the plastron (see Fig. 32a)
canthal scales on the canthus rostralis bet-
ween the eye and the nasal opening (Figs.
82,429,433)
carapace dorsal shell
carnivore meat-eater
caudal on the tail, pertaining to the tail
conspecific of the same species
convex curved toward the outside
cryptic coloration coloration matching the
surrounding environment
distal further from the center of the body or
from the point of reference
dorsal on the back, towards the back
dorsolateral transition from the back to the
side
endemic restricted in distribution to a parti-
cular location, limited distribution
femoral pores specifically arranged glandu-
lar openings on the underside of the upper
thigh
femoral scale thigh scale; a paired scale on
the plastron between the abdominal scale
and the anal scale (Fig. 32c)
frontal (pl. frontals) frontal scale, scale(s) on
the top of the head between the eyes (Figs.
102, 159, 160, 162, 168,488)
frontonasal (pl. frontonasals) scales between
the rostral and prefrontals (Figs. 102, 111,
159, 430)
granular small, conical, non-overlapping (Fig.
352d)
gular shield the foremost scale on the plastron,
may be paired or unpaired (Fig. 32c)
habitat the physical and biotic characteristics
of an area in which plants or animals live
hemipenis (pl. hemipenes) paired copulatory
organ in lizards and snakes
herbivore plant-eating
heterogeneous non-uniform (heterogeneous
scalation see Fig. 353b)
homogeneous similar, uniform
humeral scale a paired scale on the plastron
between the gular scale and the pectoral
scale (Fig. 32c)
imbricate overlapping (Fig. 353a,b,e)
incubation developmental phase of eggs,
brooding of eggs
infralabial infralabial scale, scales on the
lower lip (Figs. 82a, 102, 488)
inframarginals scales on the bridge below
the marginal scales on a turtle shell
inguinal scale a small scale present on both
sides of a turtle shell between the bridge
and the femoral scale (Fig. 32c)
insectivore insect eater
intercalary scales "pushed in scales", in igu-
anas, the small scales between the whorls
of enlarged spiky tail scales (Fig. 398)
internasal internasal scale, scales between
the nasals (Figs. 102, 488)
interparietal scale on the dorsal surface of
the head between the parietals (Figs. 82,
102, 159,191, 430)
interrictals scales counted from one corner of
the mouth across dorsum of head to the
corner of the mouth on the opposite side;
391
 Glossary
this count includes the ultimate supralabial
on each side
invertebrates animals without a vertebral
column
juvenile young animal
keeled with a raised ridge (Figs. 352a,b,
527c,d)
labial pits in some boids (e.g. Corallus,
Epicrates) characteristic, tubular depressi-
on in the lip scales; infrared sensing organs
lateral at or pertaining to the side of the body
or a structure
loreal scale(s) between the canthals and
supralabials in lizards (Figs. 82a, 102, 429);
scale(s) between the nasal and preocular in
snakes (Figs. 488, 845)
loreal pit for pit vipers (subfamily crotalinae)
characteristic, tubular depressions between
the nasal opening and the eye (Fig. 845);
infrared sensing organ
lorilabial scales between the loreals and the
supralabials (Fig. 429)
marginal scute small scales on the lateral
edges of the carapace (Fig. 32a,b)
median in the middle
mental chin scale, foremost, usually particu-
larly large scale on the lower jaw (Figs.
82a,429,488,845)
mental groove extensible groove between the
pairs of enlarged chin scales (Fig. 488)
monophyletic forming a closed category des-
cended from a common ancestor
monotypic species a species not divided into
subspecies
monotypic genus a genus consisting of only a
single species
morphologic concerning the external form
mucronate bearing a projecting spine on the
rear edge (Fig. 352a)
nasal nose scale, scale encircling the nostril
(Fig. 429, 484)
nuchal scale neck scale, the foremost scale on
the carapace (Fig. 32b)
occipital scales at the back of the head behind
the parietals (Fig. 102)
omnivore an animal that eats both plant and
animal food
osteoderm superficial dermal bones under-
lying the epidermal scales
oviparous egglaying
ovoviviparous live bearing from eggs;
development of fertilized eggs in the womb,
with young hatching at or shortly after egg
laying
392
ocelli eye markings, markings consisting of a
light ring with a dark center or a dark ring
with a light center
paravertebral of or pertaining to the area
immediately lateral to the dorsal midline
parietal eye pineal eye, light sensitive organ
below the interparietal scale (Fig. 4 77)
parietal parietal scale, scale on the top of the
head posterior to level of eyes (next to the
interparietal, if present) (Figs. 159, 160,
161,464,488)
pectoral scale thoracic scale; a paired scale
on the plastron between the humeral scale
and the abdominal scale (Fig. 32c)
phylogenetic concerning evolutionary history
physiography landscape features
plastron ventral shell
pleural scale rib scale, the large lateral scales
of the plastron between the marginal scales
and the vertebral scales (Fig. 32a,b)
population a group of individuals of the same
species living in a particular area
postanal scales posterior to the cloaca} slit; 2-
4 enlarged scales are present in this area
in some lizard species (Fig. 355)
postfemoral pocket slitlike skin pocket at
the posterior insertion of the upper thigh
(Fig. 431)
postmental scales posterior to the mental
(Figs. 362, 397, 484)
postnasal postnasal scale, scale between the
nasal opening and the loreal or, in its
absence, the preocular (Figs. 488, 845)
postocular (pl. postoculars) postocular scale,
scale posterior to the eye (Figs. 102, 488,
845)
postorbital located posterior to the eye
preauricular (pl. preauriculars) scales
anterior to the external ear opening
prefrontal prefrontal scale(s), lying anterior
to the frontals (Figs. 111, 159, 430, 484,
488,489,528,529)
precloacal pores species specific arrange-
ment of glandular openings anterior to the
cloacal slit (Fig. 484)
prenasal prenasal scale, scale between the
rostral and the nasal opening (Figs. 488,
524, 845)
preocular scale(s) anterior to the eye (Figs.
102,429,488,525,845)
proximal situated near or toward the point of
reference
revision literaly a "new perspective" (on a pro-
blem), comprehensive treatment of a taxo-
nomic group

rostral rostral scale, usually particularly
enlarged scale on the tip of the upper jaw
(Figs.82,83,84,102, 159,429,430,484,
488,489,845)
rudimentary diminished, degenerated
scute (pl. scutes) large flat scale
semiaquatic lifestyle partially adapted to an
aquatic environment; most semiaquatic
species live at the edge of a body of water
and spend part of the time in the water
semiarid half dry
sexual dichromatism differential coloration
of males and females in a species
subcaudal subcaudal scale, scales on the ven-
tral surface of the tail; may be paired or
unpaired (Figs. 523, 846)
subdigital lamellae widened scales on the
underside of the toes (Figs. 187d, 358)
subimbricate only very slightly overlapping
(Fig. 352f)
subocular scale below the eye (Figs. 82a,
190b,429)
superciliary scale(s) above the eye at the
edge of the eye opening (Figs. 82a, 159, 359,
429)
supra-anal tubercle tubercular keels on the
dorsal scales above the level of the cloaca
(Fig. 838)
supra-auricular scales scales above the
external ear opening; these may be enlarged
and spiky
supra-auricular spines enlarged spiky scales
above the external ear opening (Fig. 96)
supralabial supralabial scale, upper lip scale
(Figs.82a, 102,429,484,488,845)
supranasal scale(s) above the nasal (Fig. 102)
supraocular supraocular scale, scale on the
dorsal surface of the head above the eyes
(Figs. 159,430,488,847)
supraorbital semicircles usually slightly
enlarged scales on the dorsal surface of the
head arranged in a semicircular pattern
bordering the inside of the supraoculars
(Fig. 82b)
Glossary
SVL snout-vent length, distance from snout tip
to cloaca! opening
sympatric co-occuring in the same geographic
area
synonym two or more names for the same
taxon; in systematics, a senior synonym is
the earliest available name for a taxoil' a
junior synonym is any available name '
other than the senior synonym and is
invalid
syntopic occuring together in the same location
taxonomy study of the systematic classifica-
tion of organisms
temporal temporal scale, lateral head scales
in the temporal region behind the post-
oculars, between the supralabials and the
parietal (Figs. 102, 437, 484, 488)
tubercle a small, rounded scale or bump on
the skin scale
ventral on the belly, towards the belly; of or
pertaining to the lower surface of the body
or other structure
ventral ventral scales, belly scales
ventrolateral in transition from the belly to
the lateral surface of the body or other
structure
vertebral scale central scale, scales running
along the midline of the plastron or body
(Figs. 32a,b)
vertebrates animals with a vertebral colomn
vicariance event process in which a previous-
ly uniform population is divided into two or
more subpopulations; e.g. by the formation
of barriers, flooding or mountain upheaval
viviparous live-bearing
xerophytic suited to dry areas and climate
zoogeography study of the distribution of
animals
 This book is a completely up-dated and fully
'\'UC11tjn colored illustrated guide to the 557 species
Peninsula
of crocodiles, turtles, lizards and snakes that
are known to occur in the region extending
from the Isthmus of Tehuantepec in Mexico
Guatomo~
Hondur•• to the southern extent of the Panamanian
El~r isthmus.

Special features:
• The rich illustrated keys for identification
will prove useful to anyone who has a
doubt about what reptile is in hand .
• The text is accompanied by professional
quality color photos of 466 of the 557
species.
• With expertly done line drawings of
diagnostic features.
• Distribution maps to all species.
• and more ...

About the author:

Dr. Gunther Kohler is the Head of the
Department of Herpetology at the Research
Institute Senckenberg in Frankfurt am Main,
Germany, and Director of the Conservation
Project Utila Iguana, Honduras. To date, he
has produced 15 books, 145 popular articles,
and 110 scientific papers on herpetology,
most of which pertain to Central American
amphibians and reptiles .

2nd revised and updated edition 2008:

• 300 new photographs.
• 33 more pages.
• Identification keys revised and updated.
• This new edition treats 557 species, an
increase of 20 species over the 537
species of crocodilians, turtles, lizards,
amphisbaenians, and snakes covered in
10er ISBN 3-936180-28-8
the 2003 edition.
13er ISBN 9783936180282

9 LJl,L