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Thorp and Covich’s Freshwater Invertebrates
Fourth Edition
Series Editor: James H. Thorp
Related Publications
Ecology and Classification of North American Freshwater Invertebrates
Edited by J.H. Thorp and A.P. Covich
First (1991), Second (2001), and Third (2010) Editions
Field Guide to Freshwater Invertebrates of North America
by J.H. Thorp and D.C. Rogers
Keys to Nearctic Fauna
Thorp and Covich’s Freshwater
Invertebrates - Volume II
Fourth Edition
Edited by
James H. Thorp
D. Christopher Rogers
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This book and the individual contributions contained in it are protected under copyright by the Publisher
(other than as may be noted herein).
Notices
Knowledge and best practice in this field are constantly changing. As new research and experience broaden our
understanding, changes in research methods, professional practices, or medical treatment may become necessary.
Practitioners and researchers must always rely on their own experience and knowledge in evaluating and using any
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ISBN: 978-0-12-385028-7
Fernando Álvarez [Chapter 16] Departamento de Cristina Damborenea [Chapter 5] División Zoología
Zoología, Instituto de Biología, U.N.A.M., Circuito Invertebrados, Museo de La Plata, FCNyM-UNLP,
exterior s/n, Ciudad Universitaria, Copilco, Coyoacán, Paseo del Bosque, 1900 La Plata, Argentina; email:
A.P. 70-153, México, Distrito Federal. C.P. 04510, cdambor@fcnym.unlp.edu.ar
México; email: falvarez@servidor.unam.mx R. Edward DeWalt [Chapter 16] Illinois Natural History
Bonnie A. Bain [Chapter 12] Department of Biological Survey, Center for Biodiversity, 607 East Peabody Drive,
Sciences, Southern Utah University, Cedar City, Utah Champaign, Illinois 61820, USA; email: edewalt@inhs.
84720, USA; email: bain@uss.edu illinois.edu
llse Bartsch [Chapter 16] Forschungsinstitut Senckenberg, Genoveva F. Esteban [Chapter 2] Conservation Ecology
c/o DESY, Gebaeude 3, Raum 316, Notkestr. 85, 22607, and Environmental Sciences Group, Faculty of Science
Hamburg, Germany; email: bartsch@meeresforschung.de and Technology, Bournemouth University, Dorset,
Valerie Behan-Pelletier [Chapter 16] Agriculture and United Kingdom; email: gesteban@bournemouth.
Agri-Food Canada, K.W. Neatby Building, 960 Carling ac.uk
Avenue, Ottawa, Ontario K1A 0C6, Canada; email: James W. Fetzner Jr. [Chapter 16] Biodiversity Services
valerie.behan-pelletier@agr.gc.ca Facility, Section of Invertebrate Zoology, Carnegie
Matthew G. Bolek [Chapter 10] Department of Zoology, Museum of Natural History, 4400 Forbes Avenue,
Oklahoma State University, 501 Life Sciences West, Pittsburgh, Pennsylvania 15213-4080, USA; email:
Stillwater, Oklahoma 74078, USA; email: bolek@ FetznerJ@CarnegieMNH.org
okstate.edu Bland J. Finlay [Chapter 2] School of Biological and
Ralph O. Brinkhurst [Chapter 12] 205 Cameron Court, Chemical Sciences, Queen Mary University of London,
Hermitage, Tennessee 37076, USA The River Laboratory, Wareham, Dorset, BH20 6BB,
United Kingdom; email: b.j.finlay@qmul.ac.uk
Francisco Brusa [Chapter 5] División Zoologia Inverte
brados, Museo de La Plata, FCNyM-UNLP, 1900 La Stuart R. Gelder [Chapter 12] Department of Science and
Plata, Argentina; email: fbrusa@fcnym.unlp.edu.ar Math, University of Maine at Presque Isle, Presque Isle,
Maine 04769, USA; email: stuart.gelder@umpi.edu
Richard D. Campbell [Chapter 4] Department of Develop
mental and Cell Biology, University of California, Irvine, Fredric R. Govedich [Chapter 12] Department of Biological
CA, USA; post mail: 2561 Irvine Ave., Costa Mesa, Sciences, Southern Utah University, 351 West University
California, 92627 USA; email: rcampbel@uci.edu Blvd, Cedar City, Utah 84720, USA; email: govedich@
suu.edu
Joo-lae Cho [Chapter 16] Invertebrate Research Division,
National Institute of Biological Resources, Environmental Daniel L. Graf [Chapter 11] The Academy of Natural
Research Complex, Gyoungseo-dong, Incheon, 404-170, Sciences, 1900 Benjamin Franklin Parkway, Philadelphia,
South Korea; email: Joolae@Korea.kr Pennsylvania 19103, USA; email: grad@acnatsci.org
David R. Cook [Chapter 16] 7725 North Foothill Drive Roberto Guidetti [Chapter 15] Department of Biology,
South, Paradise Valley, Arizona 85253, USA; email: University of Modena and Reggio Emilia, via Campi
watermites@msn.com 213/D, 41125, Modena, Italy; email: roberto.guidetti@
Kevin S. Cummings [Chapter 11] Illinois Natural History unimore.it
Survey, Center for Biodiversity, 607 East Peabody Ben Hanelt [Chapter 10] Department of Biology, Univer
Drive, Champaign, Illinois 61820, USA; email: ksc@ sity of New Mexico, 163 Castetter Hall, Albuquerque,
inhs.uiuc.edu New Mexico 87131, USA; email: bhanelt@unm.edu
xi
xii Contributors to Volume II
Brenda J. Hann [Chapter 16] Department of Biological Anna J. Phillips [Chapter 12] Smithsonian Institution,
Sciences, W463 Duff Roblin, University of Manitoba, National Museum of Natural History, Department of
Winnipeg, Manitoba R3T 2N2, Canada; email: hann@ Invertebrate Zoology, 10th and Constitution Ave, NW,
cc.umanitoba.ca Washington, DC 20560-0163, USA; email: phillipsaj@
Tom Hansknecht [Chapter 16] Barry A. Vittor and si.edu
Associates, Inc., 8060 Cottage Hill Rd., Mobile, George O. Poinar Jr. [Chapter 9] Department of Zoology,
Alabama 36695, USA; email: bvataxa@bvaenviro.com Oregon State University, Corvallis, Oregon 97331,
David J. Horne [Chapter 16] School of Geography, USA; email: poinarg@science.oregonstate.edu
Queen Mary University of London, Mile End Road, Wayne Price [Chapter 16] Department of Biology,
London E1 4NS, United Kingdom; email: d.j.horne@ University of Tampa, 401 W. Kennedy Blvd., Tampa,
qmul.ac.uk Florida 33606, USA; email: wprice@ut.edu
Julian J. Lewis [Chapter 16] Lewis & Associates LLC, Roberto Pronzato [Chapter 3] Dipartimento di Scienze
17903 State Road 60, Borden, Indiana 47106-8608, della Terra, dell’Ambiente e della Vita (DISTAV),
USA; email: lewisbioconsult@aol.com Università di Genova, Area Scientifico-Disciplinare 05
Lawrence L. Lovell [Chapter 12] Research Associate, (Scienze biologiche), Settore BIO/05, Genova, Italy;
Polychaetous Annelids, Research & Collections, Natural email: pronzato@dipteris.unige.it
History Museum of Los Angeles County, 900 Exposition Lorena Rebecchi [Chapter 15] Department of Biology,
Blvd., Los Angeles, California 90007, USA; email: University of Modena and Reggio Emilia, via Campi
lllpolytax@gmail.com 213/D, 41125, Modena, Italy; email: lorena.rebecchi@
Tobias Kånneby [Chapter 7] Department of Zoology, unimore.it
Swedish Museum of Natural History, 10405, Stockholm, Janet W. Reid [Chapter 16] Virginia Museum of Natural
Sweden; email: tobias.kanneby@nrm.se History, 1001 Douglas Avenue, Martinsville, Virginia
Renata Manconi [Chapter 3] Dipartimento di Scienze 24112, USA; email: jwrassociates@sitestar.net
della Natura e del Territorio (DIPNET), Università Vincent H. Resh [Chapter 16] Department of Environmental
di Sassari, Muroni 25, I-07100, Sassari, Italy; email: Science, Policy, and Management, University of California,
r.manconi@uniss.it 305 Wellman Hall, Berkeley, California 94720, USA;
William E. Moser [Chapter 12] Smithsonian Institution, email: resh@berkeley.edu
National Museum of Natural History, Department of Dennis J. Richardson [Chapter 12] School of Biological
Invertebrate Zoology, Museum Support Center, 4210 Sciences, Quinnipiac University, 275 Mt. Carmel
Silver Hill Road, Suitland, Maryland 20746, USA; Avenue, Hamden, CT 06518, USA; email: Dennis.
email: moserw@si.edu Richardson@quinnipiac.edu
Diane R. Nelson [Chapter 15] Department of Biological D. Christopher Rogers [Chapters 1, 11, 16] Kansas Bio
Sciences, East Tennessee State University, Johnson logical Survey and Biodiversity Institute, Higuchi Hall,
City, Tennessee 37614-1710, USA; email: janddnel- University of Kansas, 2101 Constant Avenue, Lawrence,
son@yahoo.com Kansas 66047, USA; email: branchiopod@gmail.com
Carolina Noreña [Chapter 5] Departamento Biodiversidad S.S.S. Sarma [Chapter 8] Laboratorio de Zoología
y Biología Evolutiva, Museo Nacional de Ciencias Acuática, Unidad de Morfología y Función, Facultad de
Naturales (CSIC), Madrid, España; email: norena@mncn. Estudios Superiores, Universidad Nacional Autónoma
csic.es de México, Av. de lo Barrios, no. 1, Los Reyes,
Roy A. Norton [Chapter 16] SUNY College of Environ Tlalnepantla, Edo. de Méx. C.P. 54090, México; email:
mental Science and Forestry, 134 Illick Hall, 1 Forestry sssarma@gmail.com
Drive, Syracuse, New York 13210, USA; email: ranorton@ Andreas Schmidt-Rhaesa [Chapter 10] Zoological Museum,
esf.edu University Hamburg, Martin Luther-King. Platz 3, 20146
Alejandro Oceguera-Figueroa [Chapter 12] Laboratorio Hamburg, Germany; email: andreas.schmidt-rhaesa@uni-
de Helmintologiá, Instituto de Biologiá, Universidad hamburg.de
Nacional Autoñoma de México, Tercer circuito s/n, Hendrik Segers [Chapter 8] School of Freshwater Biology,
Ciudad Universitaria, Copilco, Coyoacán. A.P. 70-153, Belgian Biodiversity Platform, Royal Belgian Institute
Distrito Federal, C. P. 04510, México; email: aoceguera@ of Natural Sciences, Vautierstraat 29, B-1000, Brussels,
ib.unam.mx Belgium; email: Hendrik.Segers@naturalsciences.be
Contributors to Volume II xiii
Alison J. Smith [Chapter 16] Department of Geology, Robert J. Van Syoc [Chapter 16] California Academy
Kent State University, Kent, Ohio 44242, USA; email: of Sciences, Department of Invertebrate Zoology
alisonjs@kent.edu and Geology, 55 Music Concourse Drive, San
Ian M. Smith [Chapter 16] Systematic Acarology, Environ Francisco, California 94118, USA; email: Bvansyoc@
mental Health Program, Agriculture and Agri-Food calacademy.org
Canada, K.W. Neatby Building, 960 Carling Ave., Ottawa, L. Cristina de Villalobos [Chapter 10] Facultad de Ciencias
Ontario K1A 0C6, Canada; email: smithi@agr.gc.ca Naturales y Museo, Departamento de Invertebrados,
T.W. Snell [Chapter 8] School of Biology, Georgia Institute Paseo del Bosque S/N 1900 La Plata, Argentina; email:
of Technology, 310 Ferst Drive, Atlanta, Georgia 30332, villalo@fcnym.unlp.edu.ar
USA; email: terry.snell@biology.gatech.edu Robert L. Wallace [Chapter 8] Department of Biology,
Malin Strand [Chapter 6] The Swedish Species Information Ripon College, 300 Seward Street, Ripon, Wisconsin
Centre, Swedish University of Agricultural Sciences, 54791, USA; email: wallacer@ripon.edu
Uppsala, Sweden; email: malin.strand@slu.edu Elizabeth J. Walsh [Chapter 8] Department of Biological
Per Sundberg [Chapter 6] Department of Zoology, University Science, University of Texas at El Paso, 500 W.
of Gothenburg, P.O. Box 463, SE-405 30 Gothenburg, University Avenue, El Paso, Texas 79968, USA; email:
Sweden; email: P.Sundberg@zool.gu.se ewalsh@utep.edu
Christopher A. Taylor [Chapter 16] Curator of Fishes Alan Warren [Chapter 2] Department of Life Sciences,
and Crustaceans, Prairie Research Institute, Illinois Natural History Museum, Cromwell Road, London SW7
Natural History Survey, University of Illinois at Urbana- 5BD, United Kingdom; email: a.warren@nhm.ac.uk
Champaign, 1816 S. Oak, Champaign, Illinois 61820, Timothy S. Wood [Chapters 13, 14] Department of
USA; email: ctaylor@inhs.illinois.edu Biological Sciences, Wright State University, 3640
Roger F. Thoma [Chapter 16] Midwest Biodiversity Insti Colonel Glen Highway, Dayton, Ohio 45435, USA;
tute, 4673 Northwest Parkway, Hilliard, Ohio 43026, email: tim.wood@wright.edu
USA; email: cambarus1@mac.com Fernanda Zanca [Chapter 10] Facultad de Ciencias
James H. Thorp [Chapters 1, 11, 12] Kansas Biological Naturales y Museo, Departamento de Invertebrados,
Survey and Department of Ecology and Evolutionary Paseo del Bosque S/N 1900 La Plata, Argentina; email:
Biology, University of Kansas, 2101 Constant Avenue, fmzanca@fcnym.unlp.edu.ar
Lawrence, Kansas 66047, USA; email: thorp@ku.edu
About the Editors
xv
Preface to the Fourth Edition
Those readers familiar with the first three editions of our Our concept for T&C IV included producing one book
invertebrate book (Ecology and Classification of North (Volume I, published in late 2014 with a 2015 copyright
American Freshwater Invertebrates, edited by J.H. Thorp date) with 6 chapters on general environmental issues
and A.P. Covich) will note that the fourth edition has applicable to many invertebrates, followed by 35 chapters
expanded from a North American focus to worldwide cov- devoted to individual taxa at various levels (order to phy-
erage of inland water invertebrates. We gave our book series lum, or even multiple phyla in the case of the protozoa).
on inland water invertebrates the name Thorp and Covich’s Volume I was designed both as an independent book on
Freshwater Invertebrates to: (1) associate present with past ecology and general biology of various invertebrate taxa
editions, unite current volumes, and link to future editions; and as a companion volume for users of the keys in the
(2) establish a connection between the ecological and gen- regional taxonomic volumes, thereby reducing the amount
eral biology coverage in Volume I with the taxonomic keys of information duplicated in the taxonomic volumes. The
in the remaining volumes; and (3) give credit to Professor perhaps 10 taxonomic volumes to be published in the next
Alan Covich for his work on the first three editions. For the decade or so will contain both keys for identifying inverte-
sake of brevity, we refer to the current edition as T&C IV. brates in specific zoogeographic regions and descriptions of
Whether the fifth edition of T&C will ever appear is cer- detailed anatomical features needed to employ those keys.
tainly problematic, but who knows! At present we are con- While the vast majority of authors in T&C editions
sidering producing up to 11 volumes in the fourth edition. I–III were from the United States or Canada, we attempted
While I am the sole editor of the book series at this in T&C IV to attract authors from many additional coun-
point, Christopher has been a major and highly valued part- tries in six continents. Although we largely succeeded in
ner in developing ideas for the fourth edition and is thus this goal, we expect the fifth edition of T&C—if it is ever
far an editor on the first three volumes (senior editor on the published—to continue increasing the proportion of authors
third). He will also play a major role in many of the remain- from outside North America as our books become better
ing volumes because of his diverse and global knowledge of known internationally.
freshwater invertebrates, especially in the area of taxonomy. Our goals for T&C IV are to improve the state of taxo-
As we made significant progress on the first three volumes, nomic and ecological knowledge of inland water inverte-
we began contacting some potential coeditors and authors brates, help protect our aquatic biodiversity, and encourage
to develop volumes for other zoogeographic regions and more students to devote their careers to working with these
negotiations with a few of those volumes are now under- fascinating organisms. These goals are especially impor-
way. However, we are still seeking experts in fields of tant because the verified and probable losses of species in
invertebrate taxonomy for various zoogeographic regions to wetlands, ponds, lakes, creeks, and rivers around the globe
serve as highly dependable coeditors, especially those who exceed those in most terrestrial habitats.
both work and live in the zoogeographic regions covered by
the various future volumes. James H. Thorp
xvii
Preface to Volume II
This is the second volume of the fourth edition of Thorp and We have asked authors to include only taxa that are
Covich’s Freshwater Invertebrates (T&C IV) and the first to recognized internationally by publication in reputable sci-
focus almost exclusively on taxonomy. Information on the entific journals that follow the International Code of Zoo-
ecology and general biology of the groups can be found in logical Nomenclature. Thus, no taxa that have merely been
Volume I (Ecology and General Biology, edited by Thorp proposed should be included even if they have been identi-
& Rogers, 2015), the companion text for the current and all fied by the world’s expert on that group. “Common” species
remaining books in this series. All taxonomic volumes (other are not designated because a common species in one area
than those focused exclusively on Hexapoda) are expected may not be common in another, and this designation can
to consist of an introductory chapter, a chapter on protozoa lead to overly frequent and false identifications. Authors
(multiple kingdoms), and 14 chapters on individual phyla have been encouraged to end the keys at the point where
from Cnidaria to Arthropoda. Some of the chapters are very further identification without genetic analysis is not practi-
small (e.g., Chapter 14 on Entoprocta), whereas others are cal or when it is clear that too many of the extant fauna have
huge, especially Chapter 16 on Arthropoda. yet to be described in scientific publications.
A typical chapter includes a short introduction, a brief Users of these keys need to realize that taxonomy is
discussion of limits to identification of taxa in that chapter, a growing and vibrant field in which new taxa are being
important information on terminology and morphology that described and previously accepted relationships reevalu-
is needed to use the keys, techniques for preparing and pre- ated. For some users, this volume may be sufficient for their
serving material for identification (also covered in Volume I), needs, but for others, a companion text listing known species
the taxonomic keys, and a few references. In the large chap- in a smaller geographic region may also be helpful.
ters on Mollusca (11), Annelida (12), and Arthropoda (16), This edition is strongly focused on species found in
different individuals have contributed separate sections, and fresh through saline inland waters, with a nonexclusive
thus there are multiple sections on introduction through emphasis on surface waters, thereby reflecting the bias
keys and references. While this may confuse some readers, of existing scientific literature. Again, most estuarine and
it has allowed us to gain contributions from an increased parasitic species are not covered in this book, but we do
number of experts around the world. discuss species whose life cycle includes a free-living
The multilevel keys are formatted to enable users to work stage (e.g., Nematomorpha) and species that live in hard
easily at the level of their taxonomic expertise and the needs freshwaters through to brackish waters even though they
of their project. For that reason, we separated keys by major may be normally associated with estuarine or marine habi-
taxonomic divisions. For example, a student in a college tats in some parts of their life cycles (e.g., some shrimp
course might work through one or more of the initial crus- and crabs).
tacean keys to determine the family in which a freshwater It is our hope that scientists and students from around
shrimp belongs. In contrast, someone working on an environ- the world will benefit from this volume. Suggestions for
mental monitoring project might need to identify a crayfish improving future volumes are welcome.
or crab to genus or even species, and thus would use the rele-
vant, detailed keys that require more background experience. Editors
We also designed the keys, where possible, to proceed from a James H. Thorp
general to a specific character within a couplet. D. Christopher Rogers
xix
Acknowledgments for Volume II
Many people contributed to this volume in addition to the production from the original concept to the final market-
chapter authors and those acknowledged in individual chap- ing. In particular, we appreciate our association with Else-
ters. We greatly appreciate all our colleagues who have con- vier editors and production team including Candace Janco,
tributed information, figures, or reviews to Volume II, and Rowena Prasad, Laura Kelleher, and the entire United States
also thank those who provided similar services for the earlier and overseas production teams, especially Julia Haynes.
editions, upon which the present book partially relies. We are
again grateful to the highly competent people at Academic James H. Thorp
Press/Elsevier who helped in many aspects of the book’s D. Christopher Rogers
xxi
INTRODUCTION
Chapter 1
Introduction1
James H. Thorp
Kansas Biological Survey and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS, USA
D. Christopher Rogers
Kansas Biological Survey and Biodiversity Institute, University of Kansas, Lawrence, KS, USA
Chapter Outline
Introduction to This Volume and Chapter 1 1 Key to Kingdoms and Phyla in This Volume 3
Components of Taxonomic Chapters 1 References4
How to Use This Volume 2
INTRODUCTION TO THIS VOLUME among volumes vary in specificity of their taxonomic keys.
AND CHAPTER 1 This reflects both the likely percent of the fauna that has
been named and how easily taxa can be separated by alpha
This is the second volume in the fourth edition of Thorp and taxonomic methods and associated keys.
Covich’s Freshwater Invertebrates. Unlike the first three
editions of Ecology and Classification of North American
Freshwater Invertebrates (edited by Thorp and Covich in
COMPONENTS OF TAXONOMIC CHAPTERS
1991, 2001, and 2010), the fourth edition has been split This volume is an identification manual to the inland water
into multiple texts, with Volume I (Thorp & Rogers, 2015) invertebrates of the Nearctic Region where we present infor-
providing global coverage of the ecology, general biol- mation needed to diagnose and determine these organisms
ogy, phylogeny, and collection techniques for inland water to various taxonomic levels. Other information concerning
invertebrates. Subsequent volumes provide keys to identify ecology, morphology, physiology, phylogeny, and both col-
fauna in specific zoogeographic regions. This division of lecting and culturing techniques can be found in Volume I of
volumes enabled us to produce reasonable sized volumes this series. Each of the remaining 15 chapters in the current
at relatively moderate prices instead of publishing one mas- volume is limited to a single phylum, except Chapter 2’s
sive, high priced tome. While some labs may have multi- coverage of multiple phyla of unicellular protists. Chapter 2
ple copies of the “Keys to Fauna” in their region, we also is designed for readers who only need general information
recommend that they have at least one copy of Volume I, about protists. We have attempted to include the following
in order to obtain useful background information on each five sections in those chapters: (1) a brief introduction to
invertebrate group. the broader taxon; (2) a description of identification limi-
The current chapter is organized into an introduction, tations for each taxon; (3) details of pertinent terminology
a section explaining the organization of most taxonomic and morphology; (4) information on preparing and preserv-
chapters, and a key to larger taxonomic groups. This chap- ing specimens for identification; and (5) taxonomic keys
ter’s key is designed to help the reader locate the most per- (separated by level of identification). A restricted number
tinent chapter (important probably only for students and of especially pertinent references are given in each chapter
beginning taxonomists) and begin identifying organisms in following appropriate taxonomic sections. Readers can find
their samples. Readers will note that chapters within and a much more extensive list of references to their group in
1. This chapter was written to be a useful starting point for taxonomic volumes (II, III, etc.) in all zoogeographic regions. Consequently, there will be only
minor differences among volumes.
Volume I (Chapters 3 and 7–41) along with more details on correct identification answer is always present in the key. This
collecting, preparation, and preserving major taxa. Figures assumption generally takes one of the following three forms:
in each chapter are limited to those needed for effective use
1. A ll species are identifiable using a given key. Many new
of the keys. For additional anatomical information, including
species have yet to be described, let alone discovered.
figures, see the relevant chapter in Volume I.
Generalized geographic ranges are provided for most
taxa presented herein, yet species ranges shrink, swell,
HOW TO USE THIS VOLUME and change elevation constantly, particularly as weather
and climate patterns shift. Species disperse, colonize, and
There is an old maxim that says “keys are written by people
suffer stochastic local extinctions. In addition to these
who do not need them for people who cannot use them.” We
natural processes, some species are introduced inten-
have made every effort to make these keys as user friendly
tionally or accidentally by humans, and sometimes their
as publication limitations would permit.
establishment allows other species to invade as well.
Each section begins with a basic introduction to the
2. All variation is accounted for in the key. As stated above,
morphology and terminology used in diagnosing the taxa of
identification keys use specific, primary, diagnostic
that section. Limitations to the current state of taxonomic
characters. Problems in identification are compounded
knowledge are also presented so that the reader may gauge
by taxa that: (a) have different character states at differ-
the reliability of the information presented. Only the estab-
ent times; (b) only have diagnostic characters at certain
lished, peer reviewed scientific literature was used to define
life stages or in certain genders; and/or (c) have severely
the taxonomic categories and epithets included. All names,
truncated morphology (often due to lack of sexual selec-
as far as we are aware, conform to the International Code
tion) and lack morphological characters to separate the
of Zoological Nomenclature (ICZN). All nomina and tax-
species. Furthermore, new variation within taxa is con-
onomic arrangements used, as well as the rejection of old
tinually developing, and thus, one cannot assume that
names was based on peer reviewed scientific literature.
species are immutable or develop tools predicting those
Names from unpublished manuscripts, dissertations, “in
changes.
house” designations, or records that have not been validated
3. The key is a sufficient identification tool in and of itself.
are not acceptable. Provisional names and species designated
A key is just a tool. The fact that one has a bolt that needs
“taxon 1” or “species 1” were not used unless they were pre-
removing and a wrench of the correct size does not mean
viously recognized and accepted in the peer reviewed scien-
that the bolt can be loosened. Similarly, identification
tific literature (Richards & Rogers, 2011). No new species
keys are tools to aid in taxon identification. They are pri-
descriptions or previously unpublished taxonomic arrange-
marily tools to eliminate incorrect taxa from the range
ments are presented.
of possible choices, narrowing the field to the names
The keys are dichotomus (no triplets or quadruplets are
that may be applicable. Keys are the process of elimina-
used) and are hierarchical. Thus, for a given group, the first
tion. The possibility that the specimen to be identified is
keys are to the highest taxonomic category. The second set
new, a hybrid, anomalous, or a recent invasive colonist is
of keys is to the next level, the third set to the level below
always a possible answer. This is fundamental to using
that one, and so on, down to the lowest justifiable taxonomic
any identification key.
level based on current knowledge of that group. This level
is different for different groups depending upon the state of Once one arrives at a name or group of possible names
resolution in the scientific literature. Organisms not identifi- for a specimen in hand, the specimen should then be com-
able beyond a particular taxonomic level are left at that level. pared against descriptions, distribution maps, and figures
Properly prepared keys typically employ specific, pri- of that and other taxa in that group. The descriptions, fig-
mary, diagnostic characters. Older keys often use different ures, and maps are other tools to be used in identification.
characters than the more recent keys. This shift in primary Direct comparison of the specimen at hand with identified
characters results from systematists and taxonomists testing museum material or using molecular comparisons is also
the importance of characters. The ultimate goal of the sys- sometimes necessary for a correct identification.
tematist is to ensure that the interpretation of which char- Species are not immutable, fixed in location and form.
acters are important will converge with biological reality. They change constantly and will continue to do so, con-
To a non-taxonomist, this process may seem merely to be founding keys and any other identification method, such
“lumping and splitting,” rather than the result of employing as trait tables, character matrices, or even genetic analyses.
the scientific method to reveal natural relationships. This is why biology is far behind physics in the develop-
Surprisingly, many users do not know how to interpret a ment of unified theories: biology is far more complex than
dichotomus key, making the fundamental assumption that a physics, as it involves more interacting parts and processes.
Chapter | 1 Introduction 3
INTRODUCTION
KEY TO KINGDOMS AND PHYLA levels of interest, need, and skill without having to wade
through extraneous taxa not in the direct line to the taxon
IN THIS VOLUME
of interest.
A major change in the identification keys for our fourth The following key was derived in part from Chapter 1 in
edition has been to include multiple keys per chapter that Volume I of the fourth edition. It is meant to allow you to
generally start with a class level key and proceed to finer move to the next level of keys, which will be in individual
and finer divisions. These allow users to work at their chapters.
13’ Body with anterior tip normally obtusely rounded or blunt, posterior tip may be bi- or trilobed; cuticle opaque to dark brown or black, and
epicuticle usually crisscrossed by minute grooves; length several cm to 1 m, width 0.25–3 mm; only adults with free-living stage; hair-
worms or horsehair worms .................................................................................................................... phylum Nematomorpha [Chapter 10]
14(11) Four pairs of clawed, non-jointed legs; water bears ..................................................................................... phylum Tardigrada [Chapter 15]
14’ Adults and most larvae with jointed legs, or legs lacking, or more or less than four pairs.......................... phylum Arthropoda [Chapter 16]
REFERENCES Thorp, J.H. & D.C. Rogers (eds.). 2015. Ecology and General Biology.
Volume I of Thorp and Covich’s Freshwater Invertebrates, Fourth
Richards, A.B. & D.C. Rogers. 2011. Southwest Association of Freshwa- Edition. Academic Press, Elsevier, Boston, MA.
ter Invertebrate Taxonomists (SAFIT) list of freshwater macroinver-
tebrate taxa from California and adjacent states including standard
taxonomic effort levels. 266 pp.
Chapter 2
Protozoa
Alan Warren
Department of Life Sciences, Natural History Museum, London, UK
Genoveva F. Esteban
Bournemouth University, Faculty of Science and Technology, Dorset, UK
Bland J. Finlay
Protozoa
School of Biological and Chemical Sciences, Queen Mary University of London, The River Laboratory, Wareham, Dorset, UK
Chapter Outline
Introduction5 Material Preparation and Preservation 16
Limitations5 Isolation17
Terminology and Morphology 6 Cultivation18
Flagellates6 Preservation22
Amoebae9 Acknowledgments23
Heliozoans9 Keys to Protozoa 23
Ciliates10 References37
(D)
(C)
(A)
(B)
(F)
(E)
(G)
Protozoa
(H)
(I)
(K)
(J)
(L)
(M)
(O)
(N)
(P)
FIGURE 2.2 (A) Uroglena americana (mixotrophic); (B) Desmarella moniliformis; (C, D) Sphaeroeca volvox, individual and colony; (E) Codosiga
botrys; (F) Diploeca plactita; (G) Salpingoeca fusiformis; (H) Monosiga ovata; (I) Bioeca lacustris; (J) Bodo caudatus; (K) Cercomonas sp.;
(L) Cephalomonas cyclopum; (M) Hexamita inflata; (N, O) Pleuromonas jaculans, attached and amoeboflagellate forms; (P) Rhynchomonas nasuta.
Scale 2.5 μm for P; 5 μm for F, G, H, I, K, L; 10 μm for A, B, C, J, M, N, O; 20 μm for E; and 30 μm for D. After: Bourelly (1968) L; Calaway & Lackey (1962)
N, O, P; Lackey (1959) B, F; Lee et al. (1985) K; Pascher (1913) C, D, E, G, H, I, J, M.
but many members of the suborder Bodina live in fresh- The pellicle is covered with plates, although these also are
water (Vickerman, 1976). The best-known genus is Bodo, not generally visible.
which, like other bodonids, has two flagella (Fig. 2.2 J) one The dinoflagellates (Fig. 2.3 A–C) form a very large
of which trails, while the other extends ahead. and unique group, which is probably more important
The cryptomonads include many common heterotrophs in marine than freshwater environments. Their unique
and autotrophs and a few mixotrophs. The two flagella are arrangement of flagella, one spiraling around the cell in
unequal in length and arise from a subapical invagination a groove (girdle) and a second distally directed in another
commonly referred to as a “gullet,” although it does not groove (sulcus), makes them distinctive. Again, heterot-
appear to be the site of ingestion in heterotrophic forms. rophy and mixotrophy are common. A covering of plates
8 Thorp and Covich’s Freshwater Invertebrates
(D)
(I)
(J)
(K)
(N)
(L) (M)
FIGURE 2.3 (A) Peridinium; (B) Gymnodinium; (C) Gyrodinium; (D) Khawkinea halli; (E) Polytomella citri; (F) Entosiphon sulca-
tum; (G) Petalomonas abcissa; (H) Peranema trichophorum; (I) Urceolus; (J) Chilomonas paramecium; (K) Paraphysomonas vestita;
(L) Spumella (Monas) vivipara, two cell shapes; (M) Ochromonas variabilisa; (N) Dinobryon sertularia (mixotrophic). Scale 5 μm for E; 10 μm for A, B,
C, G, I, J, K, L, M; and 20 μm for D, F, H, N. After: Bourelly (1968) L; Calaway & Lackey (1962) E, F, J, N; Eddy (1930) A; Jahn & McKibben (1937)
D; Leedale (1985) H; Pascher (1913) M; Lemmerman (1914) K; Shawhan & Jahn (1947) G; Smith (1950) I.
may or may not be present (hence the terms armored and in recognizing the members of this group. Chrysophytes con-
naked dinoflagellates). tain both colorless heterotrophs and pigmented mixotrophs.
Chrysophytes are generally small, and they prey on bac- Euglenids are generally large flagellates with two fla-
teria. They have two unequal flagella, one long and directed gella, although in many taxa, only one flagellum emerges
anteriorly, the other short and directed laterally (Fig. 2.3 K–M). from the gullet (Fig. 2.3 D). Several heterotrophic species
They are naked or covered in fine siliceous scales (Esteban creep over the substrate with the second flagellum trail-
et al., 2012), which are not always visible with light micros- ing and hidden beneath the cell (Fig. 2.3 F–H), as in some
copy; many are amoeboid. Their carbohydrate storage product, bodonids. The euglenids are currently assigned to the super-
chrysolaminarin, occurs in liquid globules and may be useful group Excavata (Adl et al., 2012).
Chapter | 2 Protozoa 9
(A) (B)
(I)
(H)
(G)
(F)
Protozoa
(J)
(K) (L)
(P)
(O)
(M) (N)
FIGURE 2.4 (A) Vahlkampfia avaria; (B) Naegleria; (C) Stachyamoeba lipophora; (D) Thecamoeba sphaeronucleolus; (E) Vanella miroides; (F) Amoeba pro-
teus; (G) Mayorella bigomma; (H) Vexillifera telemathalassa; (I) Hartmannella vermiformis; (J) Chaos illinoisense; (K) Saccamoeba lucens; (L) Trichamoeba
cloaca; (M) Echinamoeba exudans; (N) Acanthamoeba; (O) Filamoeba nolandi; (P) Hylodiscus rubicundus. Scale 10 μm for A, B, C, E, I, M; 15 μm for H,
N, O, P; 30 μm for D, G, K, L; 50 μm for F; and 100 μm for J. After: Bovee (1985) A, B, C, D, H, I, J, M, N, O, P; Kudo (1966) F; Page (1988) E, G, K, L, P.
(D)
(C)
(B)
(A) (E)
(O) (Q)
(P)
(R)
(T)
(S) (W)
(U) (V)
FIGURE 2.5 (A) Cochliopodium bilimbosum; (B, C) Phryganella nidulus, side and oral views; (D) Pyxidicula operculata; (E) Plagiopyxis callida;
(F, G) Arcella vulgaris, side and dorsal views; (H, I) Penardochlamys arcelloides, side and oral views; (J, K) Difflugia corona, side and oral views;
(L, M) Hyalosphenia cuneata; (N) Lesquereusia spiralis; (O, P) Quadrulella symmetrica; (Q) Nebela collaris; (R) Penardia granulose; (S) Chlamydophrys
minor; (T, U) Lecythium hyalinum, dorsal and side views; (V) Pelomyxa palustris; (W) Pseudo difflugia gracilis. Scale 10 μm for C, D, R, S; 30 μm for
H, I, L, M, T, U, W; 45 μm for G, N, O, P; 60 μm for Q; 90 μm for B, E, F, J, K; and 500 μm for V. After: Bovee (1985) A, B, C, H, I, J, K, N, O, P, R, T, U;
Deflandre (1959) D, E, F, G, L, M, Q, S, W; Kudo (1966) V.
are named, and are variously used for capturing food, near the benthos. Some heliozoans traverse the bottom
sensation, movement, and attachment. Axopodia are with a unique tumbling motion, resulting from controlled
strengthened by a microtubular array called an axoneme changes in the length of the axopodia. Many sessile forms
or stereoplasm. The term axoneme is also used to describe with stalks are known. In sessile forms, cell division is
the microtubular core of cilia and flagella, but this does likely to be unequal, producing a dispersal stage that may
not imply homology, and the origin and ultrastructure of be flagellated or amoeboid.
axonemes is diverse (Yabuki et al., 2012). Most helio-
zoans lack the skeleton that is so characteristic of their
Ciliates
marine counterparts such as Radiolaria and Acantharia,
although some are covered in siliceous or organic scales The ciliates (phylum Ciliophora) form a natural group dis-
(Fig. 2.7 F, H), and some have a perforated shell or capsule tinguishable from other protozoa by a number of special-
(order Desmothoracida, Fig. 2.7 A). Although heliozoans ized features, including the possession of cilia, which are
are frequently planktonic, they are found primarily on or short hair-like processes, at some stage in their life cycle,
Chapter | 2 Protozoa 11
Protozoa
(L) (M)
(O)
(N)
(R)
(P) (Q)
(F)
(H)
(G)
12 Thorp and Covich’s Freshwater Invertebrates
Protozoa
FIGURE 2.8 (A) Amphitrema stenostoma; (B) Microchlamys patella; (C, I) Pinaciophora fluviatilis; (D) Rabdiophrys anulifera; (E) Rabdiaster
pertzovi; (F) Heliomorpha depressa; (G) Limnofila mynlikovi; (H) Clathrella foreli; (I) Pinaciophora fluviatilis; (J) Acanthoperla ludibunda; (K)
Acinetactis mirabilis; (L) Pompholyxophrys punicea. Scale = 200 μm C, I; 100 μm D; 50 μm A, B, G, H, J, K, L; 25 μm E, F. After Greef (1869) I;
Lemmermann (1914) K; Mikrjukov (1999) J; Mikrjukov (2001) E; Mikrjukov & Mylnikov (1995) (called Penardia cometa) G; Penard (1902) A, B;
Penard (1905) H; Rainer (1968) C, D; Schoutenden (1907) F; Siemensma (1991) L.
the presence of two types of nuclei, and a unique form of mixotrophic due to the presence of endosymbiotic algae,
sexual reproduction called conjugation. A representative or by sequestering chloroplasts from ingested algae that
ciliate is shown in Fig. 2.1 D. The body surface is covered are kept functional in the ciliate cytoplasm (Esteban et al.,
with cilia, which are mostly aligned in rows called kine- 2010).
ties. The pattern of kineties is interrupted in the region of The ciliates are divisible into 12 classes (Adl et al.,
the mouth where there may be specialized oral cilia used 2012). Members of the class Karyorelictea are thought
for feeding. The cilia may be reduced in number, espe- primitive for the group, with numerous non-dividing mac-
cially in sessile forms, or organized into larger compound ronuclei that are not highly polyploid. They are largely
ciliary organelles, such as cirri. The only large group that benthic, the best-known freshwater example being Loxodes
does not always possess cilia is the Suctoria; these are (Fig. 2.9 J). Compound ciliary organelles associated with
sessile predators whose dispersal stages are, however, the cytostome are prominent in the classes Heterotrichea
ciliated. This distinctive group is easily recognized by its and Spirotrichea. Large heterotrichs, such as Stentor and
feeding tentacles. The novice should take care not to con- Spirostomum (Fig. 2.10 A–F), are familiar as teaching mate-
fuse small, ciliated animals with ciliates; the size range of rial. Spirotrichs are abundant in many freshwater habitats,
ciliates overlaps that of several metazoan groups, such as from plankton (choreotrichs and oligotrichs, Fig. 2.11 S–W)
turbellarians, rotifers, and gastrotrichs. Some ciliates are to the benthos (e.g., many stichotrichs and hypotrichs).
Chapter | 2 Protozoa 13
(D) (E)
(C)
(A) (B)
(K)
Protozoa
(M) (O)
(N)
(P)
(Q)
(L)
(U)
(V)
(W) (X)
FIGURE 2.9 (A) Prorodon teres; (B) Pseudoprorodon ellipticus; (C) Holophyra simplex; (D) Trachelius ovum; (E) Paradileptus robustus;
(F) Amphileptus claparedi; (G) Litonotus fascicola; (H) Dileptus anser; (I) Loxophyllum helus; (J) Loxodes magnus; (K) Cyrtolophosis mucicola; (L, M,
N) Philasterides armata, live, silver-stained, and oral detail of silver-stained specimen; (O) Loxocephalus plagius; (P) Urozona bütschlii; (Q) Balanonema
biceps; (R) Pleuronema coronatum; (S) Histiobalantium natans; (T) Cohnilembus pusillus; (U) Uronema griseolum; (V) Cinetochilum margaritaceum;
(W) Cyclidum glaucoma; (X) Calyptotricha pleuronemodies. Scale 10 μm for K, Q; 15 μm for P, V; 20 μm for T, U, W, X; 25 μm for G, H, L, M; 30 μm
for C, I, S; 40 μm for B, R; 50 μm for F; 60 μm for A, O; and 75 μm for D, E, J. After: Corliss (1979) R; Dragesco (1966a) I; Grolière (1980) M, N; Kahl
(1930–1935) A, B, C, F, G, J, K, O, P, Q, S, V, W, X; Kudo (1966) I; Noland (1959) L, T, U.
Stichotrichs and hypotrichs (Figs. 2.11 A–H, N–Q; and established on the basis of small subunit (SSU) rRNA gene
2.12 X, Y) are mostly dorsoventrally flattened crawlers sequence data. Armophoreans are found only in anoxic hab-
with compound ciliary structures called cirri. itats, benthic, pelagic, or as endosymbionts in the digestive
The Nassophorea are named for their basket-like nasse or systems, mainly of invertebrates. Armophoreans are free-
cyrtos supporting the cytopharynx (Fig. 2.12 V, W, Z). The swimming, typically small to medium-size, with multiple
armophoreans were formerly placed in the Heterotrichea adoral polykinetids and a somatic ciliature that is typically
but are now recognized as a separate class, Armophorea, holotrichous but sometimes reduced (Fig. 2.11 K, R).
14 Thorp and Covich’s Freshwater Invertebrates
(O)
(L) (N)
(K)
(M)
(T) (U)
(S)
(P)
FIGURE 2.10 (A) Spirostomum minus; (B) Blepharisma lateritium; (C) Bursaria truncatella; (D) Climacostomum virens; (E) Condylostoma tardum;
(F) Stentor polymorphus, half extended; (G) Actinobolina radians; (H) Coleps hirtus; (I) Bryophyllum lieberkühni; (J) Metacystis recurva; (K) Lacrymaria
olor; (L) Askenasia volvox; (M) Urotricha farcta; (N) Mesodinium pulex; (O) Vasicola ciliata; (P) Trachelophyllum apiculatum; (Q) Enchelyodon ele-
gans; (R) Homalozoon vermiculare; (S) Enchelys simplex; (T) Chaenea teres; (U) Spathidium spathula; (V, W) Didinium nasutum, live and silver-stained.
Scale 10 μm for M, N; 20 μm for H, J, L, P, S; 30 μm for G, O, U; 40 μm for B, K, T; 60 μm for E, Q, R; 80 μm for D, V, W; 100 μm for A, F, I; and 200 μm
for C. After: Dragesco (1966a) K, S, V, W; Dragesco (1966b) P, R; Kahl (1930–1935) A, B, D, E, F, G, H, I, J, L, M, N, O, Q, T, U; Kent (1882) C.
Classes Prostomatea (Fig. 2.10 J, O) and Litostomatea the cytostome, on a proboscis, on tentacles, or elsewhere on
(Figs. 2.9 D, E, H; and 2.13 J, M) are largely predators, the body. A number of short, specialized kineties (rows of
often of other ciliates. Prostomes generally have apical cyto- kinetosomes) are often found near the anterior. This brosse
stomes, while many litostomes have subapical, sometimes (brush) probably assists in prey recognition.
slit-like cytostomes. The mouth is encircled by a crown of Class Phyllopharyngea contains the distinctive Suctoria
cilia from whose bases (kinetosomes) arise the rhabdos, (Figs. 2.13 B, F, I; 2.14; 2.15 A–C; and 2.16 B, C, J, L), ses-
a cylinder of microtubules surrounding and supporting the sile or free-floating predators of other ciliates. Suctoria are
cytopharynx. Toxicysts are found in most species and are unusual in that most have several “sticky” feeding tentacles
used to subdue active prey. Toxicysts may be found around rather than a single mouth. Suctoria reproduce by unequal
Chapter | 2 Protozoa 15
(B) (C)
(A) (G) (H)
(K)
(J)
(Q)
(P)
(N)
(M) (O)
Protozoa
(I)
(L)
(W)
(U)
FIGURE 2.11 (A) Gastrostyla steini; (B) Uroleptus piscis; (C) Oxytricha fallax; (D) Urostyla grandis (dorsal view); (E) Stylonychia mytilus
(dorsal view); (F) Gonostomum affine; (G) Tetrastyla oblonga(called Amphisiella oblonga); (H) Stichotricha aculeata; (I) Hypotrichidium conicum; (J)
Discomorphella pectinata; (K) Metopus es; (L) Myelostoma flagellatum; (M) Saprodinium dentatum; (N,O) Chaetospira mülleri, contracted and extended
forms; (P) Strongylidium crassum; (Q) Psilotricha acuminata; (R) Caenomorpha medusula; (S) Tintinnidium fluviatile; (T) Tintinnopsis cylindricum; (U)
Strombidinopsis setigera; (V) Strombidium viride; (W) Halteria grandinella; (X) Strobilidium gyrans. Scale 15 μm for L; 25 μm for H, W, X; 30 μm for F,
I, J, P, Q, R, T; 40 μm for A, G, K, M, N, O, S, U, V; 60 μm for B; 80 μm for C, E; and 140 μm for D. After: Jankowski (1964a,b) J, M; Kahl (1930–1935)
F, G, H, I, K, L, N, O, P, Q, R, V, W, X; Kent (1882) A, B, C, D, E; Noland (1959) S, T, U.
binary fission (budding), which yields a ciliated dispersal sequence data, has been characterized, based on which the
stage or “swarmer.” Other groups within the Phyllopha- class Cariacotrichea was established (Orsi et al., 2011).
ryngea include the Cyrtophoria, which contains surface- Members of the Oligohymenophorea are mostly
associated algivores such as Chilodonella (Fig. 2.17 T), plus microphagous, and this class is named for the compound
a diverse array of epizooic and free-living forms such as cho- ciliary organelles that are found in a buccal cavity sur-
notrichians and rhynchodians (Gong et al., 2009). rounding the cytostome. The most common pattern (in
Colpodeans (Figs. 2.16 F, G, M; 2.17 K, L, N, P, S; and 2.18 subclasses Hymenostomatia, Scuticociliatia, and Penicu-
G) are not common in freshwater environments, most being lia; Figs. 2.9 L–X; 2.15 H, I; and 2.17 A–J) is three
terrestrial bacterivores. They are more likely to be encountered polykinetids on the left side of the buccal cavity and an
in small, temporary waters. Plagiopylea is a riboclass whose undulating membrane on the right. The net result is three
monophyly, like the class Armophorea, is based only on the brushes, the polykinetids, working against a curved wall,
evidence of sequences of the SSU rRNA gene. Also like the the undulating membrane, to deliver small particles to the
armophoreans, plagiopyleans are considered to be anaerobic cytostome. The large subclass Peritrichia (Figs. 2.12 A–U,
or microaerophilic and include groups not formerly thought to 2.13 H, and 2.18 I) contains sessile bacterivores in which
be phylogenetically related, e.g., the “classic” plagyopyleans the buccal cavity is deepened as an infundibulum, and the
(Fig. 2.17 M), which were formerly placed in the Colpodea polykinetids wind down it to the cytostome after encir-
and resemble colpodids in form, and the odontostomes (Fig. cling a prominent peristome. Somatic ciliature is absent
2.11 J, M). Recently, another anoxic ciliate lineage, which in most species. Many are attached to the substrate
was initially known only from marine environmental rRNA by a stalk, as in the common Vorticella (Fig. 2.12 K),
16 Thorp and Covich’s Freshwater Invertebrates
(C)
(G)
(E)
(A) (B)
(F)
(D) (N)
(L)
(I) (J) (K)
(H)
(M)
Protozoa
(W)
(O)
(R) (V)
(S)
(T) (U)
(P) (Y)
(Q)
(X) (Z)
FIGURE 2.12 (A) Hastatella radians; (B) Astylozoon faurei; (C) Urceolaria mitra; (D) Trichodina pediculis; (E) Scyphidia physarum; (F) Cothurnia
imberbis; (G) Vaginicola ingenita; (H, I) Zoothamnium arbuscula, individual and colony; (J) Ophrydium eichhorni; (K) Vorticella campanula; (L) Pyxicola
affinis; (M) Platycola decumbens (called Platycola longicollis); (N) Thuricola folliculata; (O) Epistylis plicatilis; (P) Rhabdostyla pyriformis; (Q, R)
Carchesium polypinum, individual and colony; (S) Opercularia nutans; (T, U) Campanella umbellaria, individual and colony; (V) Pseudomicrothorax
agilis; (W) Microthorax pusillus; (X) Aspidisca costata; (Y) Euplotes patella; (Z) Nassula ornata. Scale 15 μm for V, W; 20 μm for A, B, G, P; 25 μm for
D, E, H, F, X; 30 μm for C, Z; 40 μm for L, M, S, Y; 50 μm for O; 75 μm for K, N, Q, U; and 200 μm for I, J. After: Corliss (1979) V, Y; Kahl (1930–1935)
A, B, C, D, E, H, L, N, Q, R, T, U, W; Kent (1882) I, J, K, O, S, X; Noland (1959) F, G, M, P.
and a few are secondarily free-swimming. Peritrichs may Likewise, it is optimal that suitable temperature, light, and
be either solitary or colonial. oxygen tension regimes should also be maintained through-
out the isolation and culturing processes. Numerous meth-
ods for the isolation and cultivation of protozoa have been
MATERIAL PREPARATION
reported, and these have been reviewed or summarized on a
AND PRESERVATION number of occasions (Finlay et al., 1988; Kirsop & Doyle,
To identify certain species of protozoa, it may be necessary 1991; Nerad, 1993; Lee & Soldo, 1992; Tompkins et al.,
to cultivate them. This involves isolating them from other 1995; Day et al, 2007).
(contaminant) organisms and then growing them in a cul- Methods to collect protozoa are described in Volume I’s
ture medium. In general, all initial manipulations and trans- chapter on protozoa, but below we describe more detailed
fers should be performed where possible in media with pH methods for isolating, culturing, and preserving selected
and osmotic potential similar to those at the site of isolation. groups.
Chapter | 2 Protozoa 17
Protozoa
FIGURE 2.13 (A) Gastronauta sp; (B) Paracineta patula; (C) Metacineta micraster var. pentagonalis (called M. pentagonalis in Nozawa 1939);
(D) Choanophrya infundibulifera; (E) Solenophrya micraster; (F) Prodiscophrya collini; (G) Bryometopus pseudochilodon; (H) Usconophrys aperta;
(I) Endosphaera engelmanni in cytoplasm of Opisthonecta henneguyi; (J) Apertospathula armata; (K) Apsikrata gracilis; (L) Lecanophryella paraleptas-
taci; (M) Lagynophrya fusidens; (N) Trachelostyla ciliophorum; (O) Wallackia schiffmanni. Scale = 200 μm C, I; 100 μm B, E, O; 50 μm A, G, H, J, K,
L, M, N; 25 μm D, F. After Clamp (1991) H; Curds (1982) A, B, C, D, E, F, M; Curds et al. (1983) G, N, O; Dovgal (1985) L; Foissner & Xu (2006) J;
Foissner (1984) K; Matthes (1971) I.
(A)
(B)
(D)
(E) (F)
(C)
(G)
Protozoa
(M) (N)
(K)
(P)
(L)
(O)
FIGURE 2.14 (A) Thecacineta cothurniodes; (B, C) Metacineta mystacina, top and side views; (D) Paracineta crenata; (E) Podophrya fixa, show-
ing trophont, encysted form, and swarmer; (F) Acineta limnetis; (G) Sphaerophyra magna; (H) Trichophyra epsitylidis; (I) Dendrocometes paradoxus;
(J) Heliophrya reideri; (K) Tokophrya quadripartita; (L) Multifasciculatum elegans; (M) Squalorophyra macrostyla; (N) Discophrya elongata; (O) Stylocometes
digitalis; (P) Dendrosoma radians. Scale 15 μm for E, H, J, O; 30 μm for A, D, F, G; 50 μm for I, L, M, N; 75 μm for B, K, 150 μm for C; and 2000 μm for P.
After: Corliss (1979) P; Goodrich & Jahn (1943) F, K, L, M; Kent (1882) G, I; Matthes (1954) J, O; Noland (1959) A, B, C, D, N; Small and Lynn (2000) E, H.
the most abundant species in a sample. Details of various on discerning colony growth of isolated clones on agar sur-
dilution methods are described in Cowling (1991) and Finlay faces or within agar, and are particularly useful for amoebae
et al. (2000). Once isolated, it may be important to reduce the and some flagellates. Usually one or two drops of sample are
volume of liquid in which the cell is contained, thereby initiat- placed onto a non-nutrient agar plate that has been streaked
ing a quorum-sensing mechanism. Physical methods involve with a suitable food organism, and then incubated. Amoebae
the selection of individual protozoan cells and their transfer into then migrate across the agar surface away from site of inocula-
a growth medium. Micropipetting with thin capillary pipettes, tion, thereby isolating themselves from other organisms in the
working under a dissecting microscope, can be used for a wide sample. The amoebae may then be picked off and subcultured
variety of protozoa, particularly those that are relatively large (Lee & Soldo, 1992; Day et al., 2007). Electromigration is a
and/or slow. Other methods of isolation include silicone oil method for obtaining concentrated suspensions of ciliated and
plating, flow cytometry, agar plating, and electromigration. flagellated protozoa relatively free of bacteria and other organ-
Silicone oil plating involves the isolation of clone-founding isms. It works on the principle that many ciliates and flagel-
cells within microdroplets formed from vortex-mixed oil/cul- lates orient themselves in a direct current and migrate toward
ture emulsions (Soldo & Brickson, 1980). Flow cytometry is the cathode (Schmidt, 1982). It is particularly useful for the
an automated means of discriminatory cell sorting and isola- isolation of organisms from mud and sediment samples.
tion on the basis of various cell attributes including size and
density. It is particularly useful for cells that contain pigments
Cultivation
that give a fluorescent signal and has also been applied suc-
cessfully to isolate protozoa using their fluorescent food vacu- To maintain cultures of protozoa long term, it is necessary
ole contents (Keenan et al., 1978). Agar plating methods rely to provide a medium that suits each species and a supply of
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d. Artistic qualities of Hawthorne’s versions; how far can these be shown in
the school?
e. The ethical element: how treated?
f. Typical lessons.
7. Whittier’s “Snow Bound.”
a. Its historical value as a presentation of a type of civilization: occupation,
pleasures, interests, types of character.
b. Its literary value as an ideal treatment of its theme.
c. The point of view that of an old man’s retrospect. How far is this
appreciable by children?
d. The study of this poem involves also attention to structure, diction,
allusions, poetic descriptions, and metrical form.
e. Typical passages treated in lessons.
8. Longfellow’s “Evangeline.”
a. The poet’s departure from historical fact; its justification.
b. The idyllic element; the descriptions.
c. The central theme, and its treatment in the first and second parts of the
poem.
d. The different quality of the two parts; predominance of description and
the meditative element in the second.
e. Metrical structure.
f. What things in the poem can be made especially interesting to young
people?
9. Scott’s “Ivanhoe.”
a. Its free treatment of historical fact. The difference between historic and
poetic truth.
b. The historical novel: its general relations to history; to be regarded
primarily as literature, not as history.
c. The portrayal of ideals and customs of a past age: types of characters;
structure (plot) of the book built in accordance with this purpose.
d. Difficulties of language, allusions, etc.
e. Means of arousing interest in romantic literature. Comparisons with other
books commonly read by children.
10. Shakspere’s “Julius Cæsar.”
a. Historical basis, anachronisms, etc.
b. The nature of its appeal to young readers.
c. Treatment of verbal difficulties and of the dramatic form.
d. The action, the characters, the dramatic motives and situations.
V. Composition
VI. Grammar
1. Historical Review.
1. Historical changes in the idea of English grammar.
2. Attempt, in the Renaissance period, to Latinize the grammar of English.
Persistence of this point of view.
3. Recent changes due to philological study.
4. What now constitutes English grammar?
5. What problems remain unsolved?
2. Objects of Teaching Grammar.
1. Various theories: for correctness of expression and for discipline.
2. Modern notions of authority in usage, and of the province of grammar.
3. Amount of Grammar to be Taught.
1. How much grammar shall be taught in the schools?
2. What things are of most value?
3. Importance of syntax; of study of forms.
4. The Order of Treatment.
1. Syntax or etymology first?
2. A study of the methods of development adopted by some of the
representative textbooks.
References: Laurie, Hinsdale, Carpenter, Baker and Scott, and Chubb. Liddell,
“English Historical Grammar,” Atlantic Monthly, Vol. LXXXII. Sweet, New English
Grammar, Part II. Barbour, The Teaching of English Grammar; Goold Brown,
Grammar of Grammars (Introduction). Krapp, Syllabus of English Language and
Grammar (Columbia University Extension Syllabi, Series A, No. 5).
I I . T H E T E A C H I N G O F M AT H E M AT I C S I N E L E M E N TA RY
SCHOOLS
VI. Method
Professor Suzzallo’s list of details as set forth in The Teachers College Record,
January, 1909, p. 43, and in Smith, The Teaching of Arithmetic, chap. xiv.
I I I . T H E T E A C H I N G O F G E O G R A P H Y.
1. Course as a Whole.
A. Must be capable of being judged as good by geographers.
B. Must lead to knowledge and power.
C. Must be arranged so as to lead from known to unknown along lines of
least resistance but not least effort.
D. Method of approach of mature mind must not always be followed.
2. Some Fundamental Considerations.
A. No one course available for all localities.
B. Course should start with home geography.
C. Should lead next to elementary knowledge of world whole.
D. The emphasis to be given to continental work.
Division of work by grades.
E. Disadvantages of teaching all continents twice.
The following compromise is suggested as in general workable:
a. Twice: North America, United States, Europe, and perhaps portions of
Asia.
b. Once: Asia, Africa, Australia, and South America.
Present importance of South America.
F. Plan of intermediate work should differ from plan of upper grades.
“Concentric Circles.”
G. The place of physical geography in the course of study:
a. Home geography: observational side.
Danger of overemphasis, of giving wrong outlook.
b. Intermediate grades: give setting to life side.
Larger facts only.
c. Upper grade: basis of work.
Not to be taught as a topic by itself but as a means to an end.
H. Emphasis of industries and commercial side.
I. “Following interests of children,”—best meaning of phrase. Value and
dangers.
Recent interest in industrial education places a renewed obligation on
geography to be practical and to be free from fads.
Developing interest. Temporary and permanent interests.
References: Teachers College Record, March, 1901, pp. 9-15. McMurry, C. A.,
Special Method in Geography, chap. ii. Redway, J. W., New Basis of Geography,
chap. x. Bagley, W. C., Function of Geography in Elementary Schools, Journal of
Geography, Vol. III, p. 222. Dodge, Richard E., “Some Suggestions Concerning a
Course of Study in Geography,” Journal of Geography, vii, pp. 7-14.