Get Thorp and Covich's Freshwater Invertebrates: Keys To Nearctic Fauna 4th Edition James H. Thorp Free All Chapters

Full download test bank at ebook ebookmass.
com
Thorp and Covich's Freshwater

Invertebrates: Keys to Nearctic
Fauna 4th Edition James H. Thorp
CLICK LINK TO DOWLOAD
https://ebookmass.com/product/thorp-and-
covichs-freshwater-invertebrates-keys-to-
nearctic-fauna-4th-edition-james-h-thorp/
ebookmass.com
More products digital (pdf, epub, mobi) instant
download maybe you interests ...
Thorp and Covich's Freshwater Invertebrates: Keys to

Palaearctic Fauna 4th Edition James H. Thorp
https://ebookmass.com/product/thorp-and-covichs-freshwater-
invertebrates-keys-to-palaearctic-fauna-4th-edition-james-h-
thorp/

Neotropical and Antarctic Fauna 4th Edition Cristina
Damborenea
invertebrates-keys-to-neotropical-and-antarctic-fauna-4th-
edition-cristina-damborenea/

Neotropical Hexapoda 4th Edition Neusa Hamada
invertebrates-keys-to-neotropical-hexapoda-4th-edition-neusa-
hamada/
Introduction to Econometrics 3rd Edition James H. Stock
https://ebookmass.com/product/introduction-to-econometrics-3rd-
edition-james-h-stock/
Introduction to Econometrics, Global Edition James H.
Stock
https://ebookmass.com/product/introduction-to-econometrics-
global-edition-james-h-stock/
Business and Professional Communication: KEYS for

Workplace Excellence 4th Edition, (Ebook PDF)
https://ebookmass.com/product/business-and-professional-
communication-keys-for-workplace-excellence-4th-edition-ebook-
pdf/
Probability and Statistics (4th Edition) Morris H.

Degroot
https://ebookmass.com/product/probability-and-statistics-4th-
edition-morris-h-degroot/
Invertebrates: Fourth Edition Richard C. Brusca
https://ebookmass.com/product/invertebrates-fourth-edition-
richard-c-brusca/
Chaos and Nonlinear Psychology : Keys to Creativity in

Mind and Life Shan Guisinger
https://ebookmass.com/product/chaos-and-nonlinear-psychology-
keys-to-creativity-in-mind-and-life-shan-guisinger/
Thorp and Covich’s Freshwater Invertebrates
A Global Series of Books on the Identification,

Ecology, and General Biology of Inland Water Invertebrates
by Experts from Around the World
Fourth Edition
Series Editor: James H. Thorp
Volume I: Ecology and General Biology

Edited by James H. Thorp and D. Christopher Rogers
Published 2015
Volume II: Keys to Nearctic Fauna

Edited by James H. Thorp and D. Christopher Rogers
Published 2016
Volume III: Keys to Palaearctic Fauna

Edited by D. Christopher Rogers and James H. Thorp
Expected Publication Date: 2017
Volumes in Preparation and Under Contract

Keys to Neotropical and Antarctic Fauna
Keys to Neotropical Hexapoda
Keys to Fauna of the Australian Bioregion
Possible Future Volumes of the Fourth Edition

Keys to Oriental and Oceana Fauna
Keys to Oriental and Oceana Hexapoda
Keys to Palaearctic Hexapoda
Keys to Afrotropical Fauna
Keys to Afrotropical Hexapoda
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
AMSTERDAM • BOSTON • HEIDELBERG • LONDON • NEW YORK • OXFORD • PARIS

SAN DIEGO • SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO
Academic Press is an imprint of Elsevier
Academic Press is an imprint of Elsevier
125 London Wall, London EC2Y 5AS, UK
525 B Street, Suite 1800, San Diego, CA 92101-4495, USA
225 Wyman Street, Waltham, MA 02451, USA
The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK
Copyright © 2016 Elsevier Inc. All rights reserved.
No part of this publication may be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopying, recording, or any information storage and retrieval system, without
permission in writing from the publisher. Details on how to seek permission, further information about the
Publisher’s permissions policies and our arrangements with organizations such as the Copyright Clearance
Center and the Copyright Licensing Agency, can be found at our website: www.elsevier.com/permissions.
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
information, methods, compounds, or experiments described herein. In using such information or methods they
should be mindful of their own safety and the safety of others, including parties for whom they have a professional
responsibility.
To the fullest extent of the law, neither the Publisher nor the authors, contributors, or editors, assume any
liability for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise,
or from any use or operation of any methods, products, instructions, or ideas contained in the material herein.
ISBN: 978-0-12-385028-7
British Library Cataloguing-in-Publication Data

A catalogue record for this book is available from the British Library
Library of Congress Cataloging-in-Publication Data

A catalog record for this book is available from the Library of Congress
For information on all Academic Press publications

visit our website at http://store.elsevier.com/
Publisher: Cathleen Sether

Acquisition Editor: Laura Kelleher
Editorial Project Manager: Rowena Prasad
Production Project Manager: Julia Haynes
Designer: Greg Harris
Typeset by TNQ Books and Journals

www.tnq.co.in
Dedications from the Editors
To Henry B. Ward, George C. Whipple, W. Thomas Edmondson, and

Robert W. Pennak—pioneers who blazed a publishing trail with books
on the ecology and identification of North American freshwater invertebrates.
To Alan P. Covich, a longtime friend and valued
colleague, who not only helped develop the first three editions
but also made possible the fourth edition’s improved taxonomy and
worldwide coverage by introducing the current editors to each other.
James H. Thorp and D. Christopher Rogers
Contributors to Volume II
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, [email protected]
México; email: [email protected] 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: [email protected] 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: [email protected] 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
[email protected] 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@ [email protected]
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: [email protected]
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: [email protected] Math, University of Maine at Presque Isle, Presque Isle,
Maine 04769, USA; email: [email protected]
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: [email protected] 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: [email protected] Pennsylvania 19103, USA; email: [email protected]
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
[email protected] 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: [email protected]
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: [email protected] Oregon State University, Corvallis, Oregon 97331,
David J. Horne [Chapter 16] School of Geography, USA; email: [email protected]
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: [email protected]
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: [email protected] 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: [email protected]
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
[email protected] 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: [email protected] History, 1001 Douglas Avenue, Martinsville, Virginia
Renata Manconi [Chapter 3] Dipartimento di Scienze 24112, USA; email: [email protected]
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,
[email protected] 305 Wellman Hall, Berkeley, California 94720, USA;
William E. Moser [Chapter 12] Smithsonian Institution, email: [email protected]
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: [email protected] [email protected]
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,
[email protected] Kansas 66047, USA; email: [email protected]
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 [email protected]
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: [email protected]
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
[email protected] 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: [email protected] 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, [email protected]
USA; email: [email protected] 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: [email protected]
Uppsala, Sweden; email: [email protected] 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: [email protected] [email protected]
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: [email protected]
Champaign, 1816 S. Oak, Champaign, Illinois 61820, Timothy S. Wood [Chapters 13, 14] Department of
USA; email: [email protected] 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: [email protected]
USA; email: [email protected] 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, [email protected]
Lawrence, Kansas 66047, USA; email: [email protected]
About the Editors
field and lab. While his research emphasizes aquatic inver-

tebrates, he also studies fish ecology, especially as related
to food webs. He has published more than hundred refereed
journal articles, books, and chapters, including three single-
volume editions of Ecology and Classification of North
American Freshwater Invertebrates (edited by J.H. Thorp
and A.P. Covich) and the first volume (Ecology and General
Biology) in the current fourth edition of Thorp and Covich’s
Freshwater Invertebrates.
Dr. James H. Thorp has been a Professor in the Depart-

ment of Ecology and Evolutionary Biology at the University
of Kansas (Lawrence, KS, USA) and a Senior Scientist in
the Kansas Biological Survey since 2001. Prior to returning
to his alma mater, Prof. Thorp was a Distinguished Profes-
sor and Dean at Clarkson University, Department Chair and
Professor at the University of Louisville, Associate Professor
and Director of the Calder Ecology Center of Fordham Uni-
versity, Visiting Associate Professor at Cornell, and Research
Ecologist at the University of Georgia’s Savannah River Ecol-
ogy Laboratory. He received his Baccalaureate from the Uni- Dr. D. Christopher Rogers is a research zoologist at the
versity of Kansas (KU) and both Masters and PhD degrees University of Kansas with the Kansas Biological Survey and
from North Carolina State. Those degrees focused on zool- is affiliated with the Biodiversity Institute. He received his
ogy, ecology, and marine biology with an emphasis on the PhD degree from the University of New England in Armi-
ecology of freshwater and marine invertebrates. Dr. Thorp has dale, NSW, Australia. Christopher specializes in freshwa-
been on the editorial board of three freshwater journals and ter crustaceans (particularly Branchiopoda and Decapoda)
is a former President of the International Society for River and the invertebrate fauna of seasonally astatic wetlands
Science. He teaches freshwater, marine, and general ecologi- on a global scale. He has numerous peer reviewed publi-
cal courses at KU, and his master’s and doctoral graduate cations in crustacean taxonomy and invertebrate ecology,
students work on various aspects of the ecology of organ- as well as published popular and scientific field guides and
isms, communities, and ecosystems in rivers, reservoirs, and identification manuals to freshwater invertebrates. Christo-
wetlands. Prof. Thorp’s research interests and background pher is an Associate Editor for the Journal of Crustacean
are highly diverse and span the gamut from organismal biol- Biology and a founding member of the Southwest Associa-
ogy to community, ecosystem, and macrosystem ecology. He tion of Freshwater Invertebrate Taxonomists. He has been
works on both fundamental and applied research topics using involved in aquatic invertebrate conservation efforts all over
descriptive, experimental, and modeling approaches in the the world.
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.
Thorp and Covich’s Freshwater Invertebrates. http://dx.doi.org/10.1016/B978-0-12-385028-7.00001-9

Copyright © 2016 Elsevier Inc. All rights reserved. 1
2 Thorp and Covich’s Freshwater Invertebrates
INTRODUCTION
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.
Freshwater Invertebrate Kingdoms and Phyla

1 Multicellular, heterotrophic organisms as individuals or colonies (sometimes with symbiotic autotrophs)........................... kingdom Animalia
.................................................................................................................................................................................................................................. 2
1’ Unicellular (or acellular) organisms present as individuals or colonies with nuclei irregularly arranged; heterotrophic and/or autotrophic;
multiple phyla within the autotrophic protozoa phyla ....................................................................................... kingdom Protista [Chapter 2]
2(1) Radially symmetric or radially asymmetric organisms living individually or in colonies ............................................................................. 3
2’ Individuals bilaterally symmetric ................................................................................................................................................................... 4
3(2) Surface not porus; oral tentacles always present around a closeable mouth; colonial or single, mostly single polyp forms (primarily hydra)
or rarely medusoid form (freshwater jellyfish); adults with a single central body cavity opening to the exterior and surrounded by cellular
endoderm, acellular mesoglea, and cellular ectoderm ....................................................................................... phylum Cnidaria [Chapter 4]
3’ Surface porus; colonial; tentacles absent; no closable orifices; without discrete organs; cellular-level (or incipient tissue-level) construction;
variable, non-distinct colony shapes, including encrusting, rounded, or digitiform growth forms; skeleton of individual siliceous spicules
and a collagen matrix; internal water canal system; may contain symbiotic algae; the sponges ....................... phylum Porifera [Chapter 3]
4(2) Oral region with numerous tentacles or cilia distributed around the mouth; organism never with eversible jaws and never vermiform as
adult ................................................................................................................................................................................................................ 5
4 Oral region with two or no tentacles, or tentacles behind the mouth ............................................................................................................. 7
5(4) Oral region with tentacles, organisms in gelatinoids or branching colonies .................................................................................................. 6
5’ Oral region ringed with cilia, muscular pharynx (mastax) with complex set of jaws; single free swimming, or semi-sessile living singly or
in small colonies; wheel animals, or rotifers ...................................................................................................... phylum Rotifera [Chapter 8]
6(5) Oral tentacles (the lophophore) in a “U” or “horseshoe” shape around mouth; anus opens outside of lophophore; colonial animals, often in
massive colonies attached to hard surfaces; true bryozoans ....................................................... phylum Ectoprocta (Bryozoa) [Chapter 13]
6’ Both mouth and anus open within lophophore; individual (non-colonial) animals with a calyx containing a single whorl of 8–16 ciliated
tentacles ........................................................................................................................................................ phylum Entoprocta [Chapter 14]
7(4) Not with the combination of characteristics described below ........................................................................................................................ 8
7’ Small (50–800 μm), spindle- or tenpin-shaped, ventrally flattened with a more or less distinct head bearing sensory cilia; cuticle usually
ornamented with spines or scales of various shapes; posterior of body often formed into a furca with distal adhesive tubes; gastrotrichs
(pseudocoelomates) ...................................................................................................................................... phylum Gastrotricha [Chapter 7]
8(7) Anterior mouth and posterior anus present ..................................................................................................................................................... 9
8’ Flattened or cylindrical, acoelomate worms with only one, ventral digestive tract opening; sometimes with evident head; turbellarian flat-
worms (commonly called planaria, a non-specific, and usually incorrect name) .................................. phylum Platyhelminthes [Chapter 5]
9(8) Vermiform or not, eversible oral proboscis not present, although eversible jaws or other mouthparts may occur ...................................... 10
9’ Long, flattened, unsegmented worms with an eversible proboscis; ribbon worms ......................................... phylum Nemertea [Chapter 6]
10(9) Body not enclosed in a single, spiraled shell or in a hinged, bivalved shell; or if a bivalved shell is present, then animal has jointed legs .......... 11
10’ Soft-bodied coelomates whose viscera is covered (in freshwater species) by a single or dual (hinged), hard calcareous shell; with a ventral
muscular foot; fleshy mantle covers internal organs; snails, clams, and mussels ........................................... phylum Mollusca [Chapter 11]
11(10) Segmented legs absent in all life stages; if jaws are present, then body with at least 20 segments .............................................................. 12
11’ Adults and most larval stages with legs; if larvae without legs or prolegs (some insects), then cephalic region with paired mandibles, or
eversible head, always with less than 15 body segments .............................................................................................................................. 14
12(11) Organism vermiform, not segmented ............................................................................................................................................................ 13
12’ Organism vermiform or not, body segmented ................................................................................................. phylum Annelida [Chapter 12]
13(12) Body cylindrical, usually tapering at both ends; cuticle without cilia, often with striations, punctuations, minute bristles, etc.; 1 cm long
(except family Mermithidae, <6 cm); nematodes, roundworms .......................................................................... phylum Nemata [Chapter 9]
INTRODUCTION
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
INTRODUCTION amoebozoans, opisthokonts, rhizarians, and excavates

(Cavalier-Smith, 2010; Adl et al., 2012). Protozoa typi-
During the last 20 years, studies on the systematics and evo- cally measure 5 to 1000 μm in size, and most are visible
lution of unicellular eukaryotes (algae, protozoa, and lower only with the aid of a microscope. There is considerable
fungi) have been in a state of great activity. Over this period, morphological and physiological diversity within the
many taxonomic boundaries, including those between the group. Because actively feeding protozoa need water,
algae and protozoa, have been broken down and new rela- all free-living (non-parasitic) protozoa are essentially
tionships established (Cavalier-Smith, 2010; Adl et al., aquatic, living in freshwater (including soil), brackish,
2012). As a result, the constituent organisms are grouped and marine environments.
together by some workers as protists, reviving the term orig-
inally coined by Haekel (1866), or as protoctists (Margulis
et al., 1989), although many systematists believe that such
LIMITATIONS
groups have no evolutionary or systematic validity. By con- There are a number of factors that pose significant limita-
trast, other workers have proposed systems that retain the tions to the taxonomy of protozoa. These include: (1) the
Kingdom Protozoa, albeit with much modified definitions lack of adequate methods for the fixation and long-term
and boundaries (Cavalier-Smith, 2010). Nevertheless, the preservation of specimens for much of the ca. 350-year his-
terms algae and protozoa are still useful in a functional or tory of the discipline of protozoology; (2) an absence of type
ecological sense, defining (primarily) photoautotrophic and specimens for most species; (3) a lack of sufficient morpho-
heterotrophic protists, respectively. logical features for species circumscription; (4) inadequate
Protozoa sensu lato, which means first animals, species descriptions for reliable identification; (5) high
are a diverse assemblage that comprises a number of rates of synonymy; (6) insufficient numbers of trained tax-
separate lineages representing almost all the major onomists; (7) undersampling and a large unknown species
eukaryote clades, including alveolates, stramenopiles, diversity; and (8) technical difficulties in culturing many
Thorp and Covich’s Freshwater Invertebrates. http://dx.doi.org/10.1016/B978-0-12-385028-7.00002-0

Copyright © 2016 Elsevier Inc. All rights reserved. 5
species, which is sometimes a prerequisite for adequate

characterization.
It is often difficult and time-consuming to identify pro-
tozoa to the level of species. In many cases, unambiguous
identification requires specialized staining techniques or the
use of electron microscopy. The taxonomic grouping used
in our key is an amalgamation of publications by special-
ists on the different groups (e.g., Lee et al., 2000; Lynn,
2008; Bass et al., 2009; Cavalier-Smith, 2010; Smirnov
et al., 2011; Adl et al., 2012). While the taxonomy of many
groups is based on a combination of cell morphology, ultra-
structural features, and molecular data, this key is designed
to make possible the identification of many protozoa to the
family level using light microscopy alone. Although obser-
vation of living organisms is important for identification,
the key should still be useful for many fixed samples. The
Protozoa
illustrations used as examples here are of one or more spe-

cies considered typical of a genus.
Although this key primarily deals with free-living pro-
tozoa, some ciliates that are commensal or parasitic, e.g.,
certain groups of suctorians and oligohymenophoreans, are
also included.
FIGURE 2.1 Examples of the main functional groups of protozoa.

TERMINOLOGY AND MORPHOLOGY (A) Peranema trichophorum—a flagellate; (B) Amoebae proteus—an
amoeba; (C) Actinophrys sol—a heliozoan; (D) Tetrahymena sp.—a ciliate.
Traditionally, free-living protozoa have been divided into
After Vickerman & Cox (1967) A, B; Siemensa (1991) C; Curds (1982) D.
three main groups according to their morphology and means
of locomotion: flagellates, amoebae (including heliozoans),
and ciliates (Fig. 2.1). Of these, only the ciliates are a truly Other groups of flagellates contain mostly or entirely
natural, monophyletic group, the flagellates and amoebae autotrophic forms with chloroplasts. However, many of the
being polyphyletic and include groups that may be only dis- pigmented, autotrophic taxa are also capable of phagot-
tantly related. Nevertheless, from a practical viewpoint, it rophy, producing an overall condition called mixotrophy
is still sometimes useful to refer to these groupings because (Sanders, 1991; Esteban et al., 2010), and also among these
isolation, cultivation, and identification methods used are groups are some wholly heterotrophic species. The groups
often the same within each group. with many mixotrophic or heterotrophic taxa include cryp-
tophytes, chrysophytes, dinoflagellates, and euglenoids,
and are usually considered phyla. Pigmentation and chlo-
Flagellates roplast morphology are important taxonomic characters for
Flagellates are characterized by the possession of one or some of these groups.
more flagella, which are long, tapering, hair-like append- Choanoflagellates, or collared flagellates, are dis-
ages that act as organelles of locomotion and feeding tinctive for the collar that surrounds the single flagellum
(Fig. 2.1 A). In free-living taxa, as opposed to parasitic spe- (Fig. 2.2 B–H). They bear a strong resemblance to sponge
cies, the number of flagella is limited; Paramastix has two choanocytes. Most choanoflagellates attach to the substrate
rows of 8–12 flagella, but most others have 1–4 (usually 2). or are colonial, and many have an external, loose-fitting
Typically, where two flagella are present, one may project covering or lorica, although this may be difficult to see with
forward, and the other trails behind. Often, the organism’s the light microscope.
flagella are longer than its body. There are several groups Bicoecids (Fig. 2.2 I) resemble choanoflagellates,
of heterotrophic flagellates in freshwater: choanoflagel- although they lack a collar. Like choanoflagellates, they are
lates, kinetoplastids, diplomonads, and bicoecids. These enclosed in a lorica and have a flagellum that is used to cre-
are raised to phyla by some authors, while bicoecids are ate a feeding current. A second flagellum lies along the cell
occasionally put with chrysophytes. Some amoeboid forms, and continues posteriorly to become an attachment to the
such as cercomonads and the Schizopyrenida, or amoebo- base of the lorica.
flagellates, also have flagella but are treated here with the Kinetoplastids (Fig. 2.2 J, N–P) are known mostly as
amoebae. parasites, especially Trypanosoma and its relatives,
Chapter | 2 Protozoa 7
(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
(A) (B) (C)
(E) (F) (G)

(H)
Protozoa
(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).
(A) (B)
(C) (D) (E)
(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.
Amoebae simultaneously, as in Amoeba (Figs. 2.1 B and 2.4 F), or

by moving as a single mass on a broad front (2.4 E, P),
The primary characteristic of amoebae is their possession
or as a cylinder (limax amoebae, Fig. 2.4 I, K, L). Not only
of pseudopodia, retractile processes that serve as organelles
do pseudopodia have characteristic shapes, but the tail end
of locomotion and feeding (Fig. 2.1 B). There is consider-
or uroid may be distinctive (Fig. 2.4 J, L), and the cell sur-
able diversity of structure in the amoebae, particularly in
face may be distinctly sculptured, as in Thecamoeba (Fig.
the character of any shell or skeletal material that may be
2.4 D). The classification of the naked, lobose amoebae was
present, and in the type of pseudopodium, for example,
recently revised by Smirnov et al. (2011).
broadly lobed, needle-like, or reticulate. Amoebae range
Other groups of amoebae, notably the testate amoebae,
in size from only a few micrometers to 2 mm in diameter.
possess shells (or tests) that may be proteinaceous, agglu-
Although many lack a fixed external morphology, the char-
tinate, siliceous or calcareous in composition (Figs. 2.5
acteristic morphologies shown by the various taxa are sur-
A–Q and 2.6 B–K). These are generally vase-shaped, with
prisingly distinctive, even if difficult to quantify (Fig. 2.4).
a single opening through which pseudopodia emerge. Many
By using also the number, size, and structure of organelles
are terrestrial, but benthic forms are common, and a few
and characteristics of tests (where present), identification is
are planktonic. Identification of testate amoebae is mainly
not as difficult for living specimens as might be imagined.
based on shell characters, i.e., size, shape, and composition.
The morphology of amoebae is plastic. Many adopt a stel-
late morphology if suspended in water, but few are truly
planktonic; rather, they live on surfaces or in sediments.
Heliozoans
In most, for example, Amoeba (Figs. 2.1 B and 2.4 F), the Heliozoans and pseudoheliozoans are roughly spheri-
cytoplasm is divided into an inner granular endoplasm and cal amoebae with many stiff projections called axopo-
an outer hyaline ectoplasm, or hyaloplasm, with a charac- dia radiating outward from the cell surface (Figs. 2.1 C,
teristic thickness and distribution around the cell. Loco- 2.7, and 2.8 D, E, I, J, L). The axopodia give heliozoans
motion may be achieved by extending many pseudopodia their characteristic sun-like appearance for which they
(D)
(C)
(B)
(A) (E)
(G) (H) (I)

(F)
(J) (K) (L) (M) (N)

Protozoa
(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,
FIGURE 2.6 (A) Vampyrella lateritia;

(B) Paraeuglypha reticulata; (C) Euglypha
tuberculata; (D, E) Trinema enchelys, oral
and side views; (F) Sphenoderia lenta;
(G, H) Cyphoderia ampulla; (I, J) Campascus
(A) (B) (C) (D) (E) (F) triqueter; (K) Paulinella chromatophora;
(L) Diplophyrys archeri; (M) Liekerkuehnia
wagnerella; (N) Microcometes paludosa;
(O) Microgromia haeckeliana; (P) Biomyxa
vegans; (Q) Chlamydomyxa montana;
(R) Reticulomyxa filosa. Scale 10 μm for L, N,
O; 15 μm for K; 25 μm for A, B, C; 40 μm for
(K) D, E, F, G, H; 50 μm for I, J, M, Q; 80 μm for
P; and 10,000 μm for R. After: Bovee (1985)
(G) (H) (I) (J) B, D, E, F, G, H, I, J, K, L, M, N, O, P, Q, R;
Deflandre (1959) A, C.
Protozoa
(L) (M)
(O)
(N)
(R)
(P) (Q)
FIGURE 2.7 (A) Clathrulina elegans;

(B) (C) (B) Actinophrys sol; (C) Actinosphaerium
eichhorni; (D) Heterophrys myriopoda;
(E) Ciliophrys infusorium; (F) Acanthocystis turfa-
cea; (G) Lithocolla globosa; (H) Raphidiophrys
elegans. Scale 15 μm for E; 30 μm for B, D, G;
50 μm for A; 75 μm for F, H; and 160 μm for C.
After: Deflandre (1959) H; Kudo (1966) B, C,
D, E; Rainer (1968) A, F, G.
(A)
(D)
(E)
(F)
(H)
(G)
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).
(D) (E)
(C)
(A) (B)
(F) (G) (H) (I) (J)
(K)
Protozoa
(M) (O)
(N)
(P)
(Q)
(L)
(R) (S) (T)
(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).
(B) (C) (D) (E)
(A) (G) (H) (I) (J)

(F)
Protozoa
(O)
(L) (N)
(K)
(M)
(T) (U)
(S)
(P)
(Q) (R) (V) (W)
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
(D) (E) (F)
(B) (C)
(A) (G) (H)
(K)
(J)
(Q)
(P)
(N)
(M) (O)
Protozoa
(I)
(L)
(W)
(U)
(R) (S) (T) (V)

(X)
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),
(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.
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.
wheat or rice, which will promote the growth of bacteria and

Isolation thereby produce a food source for the protozoa. Some com-
There is a wide variety of methods of isolation, and these can monly used enrichment methods are described in Finlay et al.
broadly be classified into three categories: enrichment meth- (1988) and Lee & Soldo (1992). The development of bacteri-
ods, dilution methods, and physical methods. Enrichment is the vores may also encourage the growth of carnivorous protozoa
inoculation of a field sample into an equal or greater volume that will feed upon them. Dilution methods are most effective
of suitable medium and incubation under favorable conditions. for use on preponderantly uniprotozoan samples. Material is
By inoculation of parallel cultures in a range of media, differ- sequentially diluted in an appropriate medium and incubated
ent organisms will be selected. For bacterivorous protozoa, the under favorable conditions. The greatest dilution in which
simplest way to enrich a sample is to add boiled grains of barley, growth occurs is likely to be uniprotozoan and usually isolates
(A)
(B)
(D)
(E) (F)
(C)
(H) (I) (J)
(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
Protozoa
FIGURE 2.15 (A) Dendrosoma radians; (B) Rhynchophrya palpans; (C) Periacineta linguifera; (D) Rheichenowella nigricans; (E) Pseudourostyla
levis; (F) Stichotricha sp.; (G) Kerona pediculus; (H) Ophryoglena rhabdocaryon; (I) Ichthiophthirius multifiliis; (J) Pseudokeronopsis similis;
(K) Parabryophrya penardi; (L) Lagenophrys nassa. Scale = 200 μm B, C, I; 100 μm E, K; 50 μm A, G, H, J, L; 25 μm D, F. After Corliss (1979) I; Curds
(1982) A, B, C, D, E, F; Curds et al. (1983) G, H, L; Shi et al. (2007) J; Foissner (1985) K.
appropriate food. Various publications provide comprehen- feeds. Many flagellates and ciliates eat bacteria, and in these
sive information or refer to media preparations for protozoa cases, non-selective media, designed for the growth of bacte-
(Kirsop & Doyle, 1991; Lee & Soldo, 1992; Nerad, 1993; rial populations, may be used. For many species, isolates may
Tompkins et al., 1995; Finlay et al., 2000; Day et al., 2007). be cultured in the presence of mixed bacterial flora that coex-
However, certain isolation techniques, growth media, and isted with the target organism in its original habitat. Alterna-
culture conditions suit a wide range of organisms. Some of tively, selective cultures may be obtained by incubating the
these are discussed below. protozoa in an inorganic salt solution along with an appropri-
The methods for the cultivation of flagellates and ciliates ate food organism. In some cases, non-pathogenic laboratory
are often identical or similar, so these two groups will be dealt cultures of bacteria may be used, whereas in others it may be
with together and the amoebae separately. The choice of cul- necessary to isolate bacteria from the original sample and use
ture medium will depend largely upon what the protozoan one or more of these strains as the selected food organisms.
Protozoa
FIGURE 2.16 (A) Lophophorina capronata; (B) Acinetopsis elegans; (C) Encelyomorpha vermicularis; (D) Psilotricha viridis; (E) Plagiocampa
ovata; (F) Cyrtolophosos mucicola; (G) Kreyella minuta; (H) Trimyema compressum; (I) Pseudochlamydonella rheophyla; (J) Pseudogemma pachystyla
(trophont on Acineta tuberosa); (K) Trypanococcus rotiferorum; (L) Manuelophrya parasitica; (M) Woodfruffia spumacola. Scale = 40 μm B, D, H, J, L,
M; 30 μm A, E; 20 μm C, K; 15 μm G, F, I. After Batisse (1968) J; Batisse (1994) K; Curds (1982) A, B, C, F, G, H, M; Curds et al. (1983) D, E; Foissner
(1993) I; Matthes (1988) L.
For omnivores and carnivores, an examination of the contents commonly used are lettuce, hay, powdered cereal leaf, and
of the food vacuoles may give an indication of the preferred grains of rice, wheat, and barley. Soil extract media are simi-
food. The organism can then be incubated in the presence of lar to plant infusions in the sense that organic compounds
its natural prey. In some cases, it may be necessary to carry out that will support the growth of bacteria are extracted from the
replicated feeding experiments using a range of food organ- soil. Thus, these media may also be used for the cultivation of
isms to determine which will support the growth of the isolate. bacterivorous species. Inorganic salt solutions provide a bal-
Culture media may be categorized into four main types: anced medium for the growth of many protozoa. However,
plant infusions, soil extract-based media, inorganic salt solu- they contain negligible quantities of organic matter, so the
tions, and specific (organically rich) media. Plant infusions addition of food organisms, or a carbon source, is essential.
are commonly used for bacterivorous flagellates and cili- Such media are commonly used for the cultivation of car-
ates. The principle is that organic compounds leach out of nivorous species. Specific, defined, media may be used for
plant material, and these support bacterial growth. The most producing axenic cultures. Such media invariably contain
(D)
(B) (E)
(A)
(C)
Protozoa
(H) (I) (J)
(F) (G)
(M) (N) (O)
(K) (L)
(T)
(R) (S)
(P) (Q)
FIGURE 2.17 (A) Frontonia leucas; (B) Stokesia vernalis; (C) Glaucoma scintillans; (D) Urocentrum turbo; (E) Parastokesia bütschlii (called
Disematostoma bütschlii); (F) Turaniella vitrea; (G) Lembadion magnum; (H) Colpidium colpoda; (I) Paramecium caudatum; (J) Clathrostoma viminale;
(K, L) Maryna socialis, individual and colony; (M) Plagiopyla nasuta; (N) Bresslaua vorax; (O) Tetrahymena pyriformis; (P, Q) Tillina magna, live and
line drawing of silver-stained specimen; (R, S) Colpoda steini, live and silver-stained; (T) Chilodonella uncinata. Scale 15 μm for G, O, R; 25 μm for C,
H, S, T; 30 μm for D, F; 40 μm for B, E, J, M; 60 μm for I, N; 75 μm for A, K, Q; 100 μm for P; and 300 μm for P. After: Corliss (1979) O, R; Dragesco
(1966b) B; Kahl (1930–1935) A, C, D, E, F, G, H, I, J, K, L, M, P; Kudo (1966) N; Lynn (1976) S; Lynn (1977) Q; Noland (1959) T.
high concentrations of dissolved organic compounds, usually whereas bacterivorous and carnivorous species may best be
derived from animal sources. Optimal maintenance conditions cultivated in the dark in order to control algal contaminants.
(e.g., temperature, pH, oxygen tension) and the frequency of Amoebae may be cultured in liquid media, on agar, or in
subculturing may be highly variable among different species biphasic media. Generally, the larger forms are grown in liquid
and should usually take account of the conditions in the natural culture, and the smaller forms are grown on agar. In most cases,
habitat and the feeding strategy of the isolate. For example, amoebae can be cultivated on a n on-nutrient medium (liquid
algivorous species should be kept in conditions of illumina- or agar) with a suitable bacterial food organism, although the
tion that allow an adequate algal food supply to be maintained, medium can be enriched to stimulate growth of the bacterial
Protozoa
FIGURE 2.18 (A) Furgasonia blochmanni; (B) Spelaeophrya troglocardis; (C) Erastophrya chattoni (growing on the peritrich Apiosoma);
(D) Cometodendron eretum; (E) Phacodinium metchnicoffi; (F) Uroleptoides kihni; (G) Cirrophrya haptica; (H) Lagynus elegans; (I) Rovinjella sphaero-
mae; (J) Rhyncheta cyclopum; (K) Mucophrya pelagica; (L) Echinophrya horrida; (M) Pseudoholophrya terricola. Scale = 200 μm C, I; 100 μm B, E;
50 μm A, G, H, J, K, L; 25 μm D, F, M. After Berger et al. (1984) M; Curds (1982) A, B, C, D, E, F, J, K, L; Curds et al. (1983) G; Dovgal (1985) L;
Matthes (1972) I; Sola et al. (1990) H.
food. Cultures of bacterivores should normally be cultivated only be visible in live cells. Nevertheless, it may also be
in the dark, whereas algivorous amoebae, or those with algal necessary to employ preservation methods, for example:
symbionts, will require a light-dark cycle. Testate amoebae are (1) if it is not possible to observe the sample for a long
often maintained on cereal leaf agar overlaid with a cereal leaf period after collection; (2) to observe certain features
infusion. Details of media formulations and culture methods not visible in live specimens; or (3) to maintain a refer-
may be found in (Lee & Soldo, 1992; Nerad, 1993; Tompkins ence collection of the organisms or permanent record of
et al., 1995; Finlay et al., 2000; Day et al., 2007). the sample. A useful general fixative is Lugol’s Iodine,
1% volume/volume, or higher in saline or hard water
(Taylor & Heynen, 1987). Mercuric chloride has been
Preservation used extensively but should probably be discontinued
Wherever possible, protozoa should be observed in vivo for safety and environmental reasons. These fixatives do
to determine their behavior and certain features that may not lend themselves to the identification of ciliates, nor
to the detection of chromatophores in small flagellates. cultivation, and preservation of protozoa is given in
Filtration methods may aid with both of these problems. Lee & Soldo (1992).
The quantitative protargol method or QPS (Montagnes
& Lynn, 1987a,b; Skibbe, 1994) produces permanent,
ACKNOWLEDGMENTS
quantitative, stained preparations for identification
of ciliates and flagellates, although it requires that The contents of this chapter are based heavily on the
samples be fixed in a concentrated Bouin’s fixative. Vari- chapter on Protozoa by Bill Taylor and Bob Sanders in
ous types of silver-staining techniques, which highlight the third edition of Ecology and Classification of North
ciliary patterns, have been used in the identification of American Freshwater Invertebrates (Thorp & Covich,
ciliates (Lee et al., 1985; Foissner, 1991). Other fixa- 2010), which we have expanded and updated where nec-
tives, such as glutaraldehyde and/or osmium tetroxide essary. We would therefore like to thank Bill and Bob
(OsO4), are used if cells are to be examined by electron for allowing us to use so much of their material, and in
microscopy. A comprehensive account many of the main doing so, acknowledge their important contribution to
commonly used methods for the collection, isolation, this chapter.
Protozoa
KEYS TO PROTOZOA
Key to Major Functional Groups of Protozoa
The phylum Cercozoa includes organisms that are amoeboid, flagellate, or both. This group is therefore included in the keys
to both these major functional groups.
1 Flagella never or rarely present; primary organelles of locomotion and feeding are either pseudopodia or cilia .......................................... 2
1’ Commonly 1–4 flagella (Fig. 2.1 A), 16–24 in one free-living genus (Paramastix).......................................... Flagellated protozoa [p. 23]
2(1) Main organelles of locomotion and feeding are pseudopodia as in Fig. 2.1 B, C. Rarely, flagella may be present as well (e.g., Figs. 2.5 S–W
and 2.6 A–I, K, P)................................................................................................................................................ Amoeboid protozoa [p. 24]
2’ Simple cilia or compound ciliary organelles characteristic (but see Fig. 2.14 A–P) and present in at least one part of life cycle; subpellicular
infraciliature present even when cilia are not; two types of nuclei (macronucleus and micronucleus) with rare exceptions (Fig. 2.1 D);
ciliated protozoa ..................................................................................................................................................... phylum Ciliophora [p. 27]
Key to Major Groups of Flagellated Protozoa

1 Without a median groove................................................................................................................................................................................. 2
1’ With median groove (annulus and sulcus); two flagella, one extending transversely around the cell (Fig. 2.3 A–C).......................................
....................................................................................................................................................................................... phylum Dinoflagellata
[Note: This group contains many or mostly photosynthetic members; several genera, however, have one or more species that lack chloro-
plasts and are thus heterotrophic, or contain chloroplasts but also exhibit heterotrophic nutrition.]
2(1) Without pharyngeal rods or paramylon as a reserve material ......................................................................................................................... 3
2’ Often large flagellates with pharyngeal rods (sometimes difficult to see) or containing the reserve material paramylon; one or usually two
flagella arise from within an anterior invagination (reservoir); contractile vacuole associated with reservoir; elongate; body shape often
plastic when living (Fig. 2.3 D, F–I) .................................................................................................................................... phylum Euglenida
3(2) Cell with 1–4 flagella (commonly two), usually of unequal length................................................................................................................ 4
3’ Cell usually with two or four equal flagella; plastids two-membraned and containing chlorophyll a and b and starch as a carbohydrate stor-
age product (Fig. 2.3 E) ................................................................................................................................................... phylum Chlorophyta
4(3) Cells without deep gullet................................................................................................................................................................................. 5
4’ Small cells with a deep, subapical gullet; two nearly equal length flagella arise from gullet (Fig. 2.3 J) ...................... phylum Cryptophyta
5(4) Cells lacking the characteristics described below ........................................................................................................................................... 6

5’ Typically with two unequal flagella; long flagellum usually directed forward during swimming, with short flagellum directed backward if
emergent; compact Golgi body frequently visible anterior to nucleus of larger cells; chrysolaminarin vesicle(s) often fill posterior of cell;
contractile vacuole usually present in extreme anterior end of cell; many species capable of simultaneous photosynthesis and phagocytosis
(Figs. 2.3 K–N and 2.2 A)................................................................................................................................................ phylum Chrysophyta
6(5) Without collar, or with indistinct collar........................................................................................................................................................... 7
6’ Single anterior flagellum encircled laterally by a tentacular, funnel-shaped collar; solitary or colonial; with or without theca (Fig. 2.2 B–H)
....................................................................................................................................................................................... class Choanoflagellida
7(6) Without protoplasmic collar............................................................................................................................................................................. 8
7’ Like Choanoflagellida above, but with second trailing flagella attached to base of lorica; protoplasmic collar; bicoecid flagellates (affinities
uncertain) (Fig. 2.2 I).................................................................................................................................................................. class Bicoecea
8(7) Ventral cytostome or flagellar pocket, with associated flagella; either kinetoplast or flagellar mastigont system present; ingestion not by
pseudopodia..................................................................................................................................................................................................... 9
Protozoa
8’ Flagellates with two unequal length flagella, one trailing; no kinetoplast or flagellar mastigont system; ingestion by pseudopodia (Fig. 2.2 K)
............................................................................................................................................................................................... phylum Cercozoa
9(8) With one or usually two flagella arising from a flagellar pocket (depression); characteristically elongate or bean-shaped. Unique organelle,
the kinetoplast, usually associated with the flagella (Fig. 2.2 J, L, N–P) ......................................................................... class Kinetoplastida
9’ Cells with one or two nucleus-flagella complexes (karyomastigonts) each with 1–4 flagella; no Golgi apparatus; when two karyomasti-
gonts, mirrored symmetry of nuclei and flagella; diplomonads (Fig. 2.2 M) ................................................................. order Diplomonadida
Key to Major Groups of Amoeboid Protozoa

1 Pseudopodia either blunt and hyaline, or filiform and sometimes branching or anastomosing, or thread-like and often branching and anas-
tomosing, or stiff and radiating ....................................................................................................................................................................... 2
1’ Large, cylindrical or ovoid multinucleate amoeba; bacterial symbionts; usually containing mineral particles; non-motile flagella-like exten-
sions; oxygen-poor habitats (Fig. 2.5 V) ......................................................................... phylum Amoebozoa, class Archamoebea Pelomyxa
2(1) Pseudopodia hyaline, usually blunt, eruptive but filiform process may occur (Figs. 2.4, 2.5 A–Q, and 2.8 B); flagellated stages may be
present.............................................................................................................................................................................................................. 3
2’ Pseudopodia not blunt and eruptive................................................................................................................................................................. 4
3(2) Small, usually <65 μm long, with eruptive pseudopodia, flagellated stage common in one group....................................................................
........................................................................................................................................ class Heterolobosea, order Schizopyrenida [p. 24]
3’ No flagellated stages....................................................................................................................................................... class Lobosea [p. 24]
4(2) Pseudopodia not as filopodia........................................................................................................................................................................... 5
4’ Filiform pseudopodia (filopodia) sometimes branching or anastomosing, granular or hyaline, many with flagella at some stage in their life
cycle; some with test (Figs. 2.5 S–W and 2.6 A–I, K, P)......................................................................................... phylum Cercozoa [p. 25]
5(4) Pseudopodia thread-like and delicate with finely granular appearance, often branching and anastomosing to form complex reticulum
(reticulopodia); test often present (Figs. 2.6 M, O, Q, R and 2.8 A)................ traditionally grouped as phylum Granuloreticulosea [p. 26]
5’ Long slender axopodia or filopodia radiating 3-dimensionally from cell; with or without skeletal elements (Figs. 2.1 C, 2.7, 2.8 D, I–J, L)
...................................................................................................................................................... Heliozoans and Pseudoheliozoans [p. 26]
Protozoa: Amoeboid Protozoa: Heterolobosea: Schizopyrenida: Families

1 Amoeboid form usually cylindrical, often monopodial; usually uninucleate; nucleolus divides to form polar masses in mitosis; temporary
flagellate stage common (Fig. 2.4 A, B) .................................................................................................................................. Vahlkampfiidae
1’ Amoeba flattened or limax; often multinucleate, nucleolus disintegrates during mitosis, no flagellate stage known (Fig. 2.4 C) ..................
.................................................................................................................................................................................................... Gruberellidae
Protozoa: Amoeboid Protozoa: Lobosea: Families

1 Lacking external test, traditionally grouped as subclass Gymnamoebia......................................................................................................... 2
1’ Incompletely enclosed in a test or other flexible cuticle of microscales traditionally grouped as subclass Testacealobosia ....................... 10
2(1) Cylindrical or flattened; flattened forms with regular outline; no trailing uroidal filaments, with rare exceptions; not strikingly eruptive .....
order Euamoebida............................................................................................................................................................................................ 3
2’ Usually flattened; frequent changes in shape typical; sometimes eruptive; subpseudopodia usually present, often furcate.......................... 8
3(2) Without subpseudopodia ................................................................................................................................................................................. 4
3’ With subpseudopodia ...................................................................................................................................................................................... 6
4(3) Cell body flattened .......................................................................................................................................................................................... 5
4’ Cell body subcylindrical ................................................................................................................................................................................. 7
5(4) Cell usually oblong; crescent-shaped hyaline margin at anterior end; pellicle-like layer, with dorsum often wrinkled and/or ridged; usually
uninucleate; no cytoplasmic crystals (Fig. 2.4 D).................................................................................................................... Thecamoebidae
5’ Body usually fan-shaped, oval, or spoon-shaped, with hyaline margin occupying up to half of length (Fig. 2.4 E)..................... Vannellidae
6(3) Subpseudopodia hyaline, blunt, digitiform, usually from anterior hyaline margin; uninucleate; nucleolar material in central body (Fig. 2.4 G)
.................................................................................................................................................................................................... Paramoebidae
6’ Few slender, conical, or linear subpseudopodia, from anterior hyaline margin or cell surface; uninucleate (Fig. 2.4 H) ................................
..................................................................................................................................................................................................... Vexilliferidae
7(4) Most species polypodial; length usually more than 75 μm; uni- or multinucleate; numerous cytoplasmic crystals (Fig. 2.4 F, J, L) ..............
Protozoa
......................................................................................................................................................................................................... Amoebidae
7’ Cell monopodial, pseudopods rare; uninucleate with central nucleolus; cytoplasmic crystals in some; cysts common (Fig. 2.4 I, K) ...........
................................................................................................................................................................................................ Hartmannellidae
8(2) Cell flattened, triangular, trapezoid or irregular in outline.............................................................................................................................. 9
8’ Cell regularly discoid, flattened ovoid, or fan-shaped; usually broader than wide; postcentral granular mass, usually surrounded, sometimes
completely, by hyaline border with short subpseudopodia (Fig. 2.4 P)...................................................................................... Hyalodiscidae
9(8) Cell flattened, broad and irregular in outline, though sometimes elongate during locomotion; slender tapering subpseudopodia, sometimes
furcate, produced from broad, hyaline lobopodium; often with small lipid globules; uninucleate (Fig. 2.4 N) ...................... Acanthamoebidae
9’ Several to many fine, sometimes furcate subpseudopodia, finer than in Acanthamoebidae (Fig. 2.4 M, O)..................................... Echinamoebidae
10(1) Test more or less rigid with distinct aperture ...................................order Arcellinida.................................................................................. 11
10’ Discoid or sometimes globose amoeba incompletely closed in a flexible tectum, no well-defined aperture (Fig. 2.5 A).................................
................................................................................................................................................................................................ Cochliopodiidae
11(10) Pseudopodia digitate and finely granular....................................................................................................................................................... 12
11’ Pseudopodia conical, clear, sometimes anastomosing; test with siliceous material embedded or attached (Fig. 2.5 B, C)..............................
................................................................................................................................................................................................... Phryganellidae
12(11) Test membranous or chitinoid, pliable or rigid; no plates or scales, but may have attached debris.............................................................. 13
12’ Test chitinoid or not, rigid, with embedded and/or attached plates, scales, siliceous granules.................... suborder Difflugina................ 16
13(12) Test round; aperture ventral .............................................................suborder Arcellina................................................................................ 14
13’ Test oval to flask-shaped; non-areolar, clear; aperture terminal (Fig. 2.5 L, M).................................................................... Hyalospheniidae
14(13) Test flexible to semi-rigid; finely or not areolate............................................................................................................................................15
14’ Test rigid, areolar, smooth; aperture ventral, round (Fig. 2.5 D, F, G) ............................................................................................ Arcellidae
15(14) Test not areolate; cytoplasm not enclosed in a separate membrane sac (Fig. 2.5 H, I) ......................................................... Microcoryciidae
15’ Test finely areolate; cytoplasm enclosed in a separate membrane sac (Fig. 2.8 B)............................................................. Microchlamyiidae
16(12) Aperture round, broadly oval or wavy (Fig. 2.5 J, K, N)............................................................................................................... Difflugiidae
16’ Aperture slit-like or narrow oval ................................................................................................................................................................... 17
17(16) Aperture terminal........................................................................................................................................................................................... 18
17’ Aperture anterioventral, invaginated, slit-like with overhanging lip (Fig. 2.5 E)....................................................................... Plagiopyxidae
18(17) Test particles rectangular (Fig. 2.5 O, P) ................................................................................................................................ Paraquadrulidae
18’ Test particles not rectangular (Fig. 2.5 Q) ....................................................................................................................................... Nebelidae
Protozoa: Amoeboid Protozoa: Cercozoa: Subphyla

1 Filopodia very fine, branching or unbranching, often with obvious granules (extrusomes); many biciliate; with or without a test.................
.................................................................................................................................................................................................... Filosa [p. 26]
1’ Filopodia or reticulopodia non-granular; cilia and test absent ........................................................................................... Endomyxa [p. 26]
Protozoa: Amoeboid Protozoa: Cercozoa: Filosa: Families

1 Without distinct test, sometimes with scales................................................................................................................................................... 2
1’ With test .......................................................................................................................................................................................................... 3
2(1) Filopodia extremely slender, branching, regularly granular; if present, cilia not visible with light microscope; cell typically small, often
globular (Fig. 2.8 G) .................................................................................................................................................................... Limnofilidae
2’ Filopodia numerous, extremely long and branching; two long cilia sometimes present; may be found in organic mud of hot springs............
........................................................................................................................................................................................................ Mesofilidae
3(1) Test without scales; may have spines and/or attached debris.......................................................................................................................... 4
3’ Test with secreted, siliceous scales arranged in definitive patterns................................................................................................................. 5
4(3) Test round, thin; may have spines or spicules (Fig. 2.5 S–U)............................................................................................. Chlamydophryidae
4’ Test rigid and agglutinated; cell does not fill test (Fig. 2.5 W)........................................................................................... Pseudodifflugiidae
5(3) Scales not as below.......................................................................................................................................................................................... 6
5’ Scales round to elliptical, thin, overlapping, adjacent or scattered (Fig. 2.6 B–F)....................................................................... Euglyphidae
6(5) Scales circular or oval; test usually with aperture at end of a neck bent to one side (Fig. 2.6 G–I)...................................................................
Protozoa
................................................................................................................................................................................................... Cyphoderiidae
6’ Scales long, with long axes perpendicular to aperture; aperture at end of short neck that is not bent; test ovoid, <45 μm (Fig. 2.6 K) ...........
...................................................................................................................................................................................................... Paulinellidae
Protozoa: Amoeboid Protozoa: Cercozoa: Endomyxa: Families

1 Medium to large (30 μm to >1000 μm); filopodia non-anastomosing and more or less radiate (Fig. 2.6 A) ............................ Vampyrellidae
1’ Small (<50 μm); filopodia lack microtubules and tend to project more from one face of the cell than from others (Figs. 2.5 R and 2.6 P)
....................................................................................................................................................................................................... Biomyxidae
Protozoa: Amoeboid Protozoa: Granuloreticulosea: Classes and Families

1 With a test, traditionally grouped as class Monothalamea .............................................................................................................................. 2
1’ Without a test, traditionally grouped as class Athalamea................................................................................................................................ 3
2(1) Test with one aperture ..................................................................................................................................................................................... 4
2’ Test with more than one aperture (Fig. 2.8 A) ......................................................................................................................... Amphitremidae
3(1) Test flattened on one side (Fig. 2.6 O)..................................................................................................................................... Microgromiidae
3’ Test not flattened on one side (Fig. 2.6 M)............................................................................................................................. Lieberkuehnidae
4(2) Multinucleate and highly reticulate plasmodia (Fig. 2.6 R) ................................................................................................. Reticulomyxidae
4’ With or without anastomosing pseudopodia; one or a few nuclei; body mass more or less round (Fig. 2.6 Q) ...............................................
............................................................................................................................................................................................ Chlamydomyxidae
Protozoa: Amoeboid Protozoa: Heliozoa and Pseudoheliozoa

[Note: some forms with pseudopodia are members of other protist groups, for example Diplophrys (Fig. 2.6 L) which is
a labyrinthulid.]
1 With axopodia radiating from centrosomes; axonemes visible with light microscope................................................................................... 2
1’ With thin, radiating filopodia; axonemes absent ................................................................. pseudoheliozoans (cercozoan subphylum Filosa)
......................................................................................................................................................................................................................... 6
2(1) Skeleton absent................................................................................................................................................................................................ 3
2’ With skeleton of siliceous or organic plates and/or spicules; sometimes stalked; axopodia and stalk highly contractile (Fig. 2.7 D, F, H)
................................................................................................................................................................phylum Heliozoa, order Centrohelida
3(2) With two cilia .................................................................................................................................................................................................. 4
3’ Cilia absent...................................................................................................................................................................................................... 5
4(3) Axopodia with regularly spaced, complex extrusomes; centrosome embedded in depression of centrally located nucleus (Fig. 2.8 F) .........
....................................................................................................................................................... phylum Cercozoa, family Heliomorphidae
4’ Axopodia bear moving granules; centrosome not in depression of nucleus which is offset (Fig. 2.8 K) ...................... family Acinetactidae
5(3) No centroplast or axoplast; axopods granule-studded and thicker at bases; large central nucleus surrounded by lacunar ectoplasm or several
nuclei at periphery of central area with vesicular ectoplasm (Fig. 2.7 B, C).................................................................... order Actinophryida
5’ Microtubule organizing center of dense plaques from the nuclear membrane or from centroplast; may be confused with Actinophryida
without knowledge of fine structure (origin and pattern of axopod microtubules) (Fig. 2.7 E).......................................... order Ciliophyrida
6(1) Test with mineralized perles, or plate scales .................................... orders Perlofilida and Rotosphaerida ................................................... 7
6’ Cell enclosed by a latticed organic capsule (skeleton), a rigid theca, or by irregular particles adhering to the outer surface...................... 12
7(6) Test of mineralised spherical perles; outer layer with apically pointed silica scales (Fig. 2.8 J).......................................................................
......................................................................................................................................................... order Perlofilida, family Acanthoperlidae
7’ Test of plate scales.................................................... order Rotospherida and perlofilid family Pompholyxophryidae.................................. 8
8(7) Test typically with two-tier plate scales; flagella absent ................................................. order Rotospherida ................................................ 9
8’ Body spherical, coated with a single layer of siliceous perles (Fig. 2.8 L)......................................................... family Pompholyxophryidae
9(8) Test with columnar radial spines, flared out both basally and apically......................................................................................................... 10
9’ Test without columnar spines ........................................................................................................................................................................ 11
Protozoa
10(9) Outer plate scales with single, large hole (Fig. 2.8 D).................................................................................................. family Rabdiophryidae
10’ Outer plate scales without holes (Fig. 2.8 E) ................................................................................................................ family Rabdiasteridae
11(9) With scalloped test of regular, unperforated cup-shaped scales; filopodia in groups of 1–3 (Fig. 2.8 H) ......................... family Cathrellidae
11’ Test not scalloped; filopodia extremely thin, not in groups (Fig. 2.8 C) ...................................................................... family Piaciophoridae
12(6) Cell enclosed in latticed organic capsule (skeleton); generally stalked; no centroplast; cell body spherical in adults (Fig. 2.7 A) .................
....................................................................................................................................................................................... order Desmothoracida
12’ Cell enclosed in rigid theca with many tine pores through which filopodia emerge; filopodia with long, thin extrusomes (Fig. 2.7 G)..........
............................................................................................................................................................. subclass Testosia, family Lithocollidae
Key to Classes and Subclasses of Ciliophora

1 Cilia present in trophont (active) stages; retractile suctorial tentacles only in family Actinobolinidae (class Litostomatea, order Haptorida).
......................................................................................................................................................................................................................... 2
1’ Suctorial tentacles present (absent in one endocommensal group); no true cytostome or cytopharynx; adults (trophonts) usually sessile,
many species ectosymbiotic, some planktonic species; cilia absent except in free-swimming dispersal larval stages; with or without lorica
...............................................................................................................................................class Phyllopharyngea, subclass Suctoria [p. 28]
[Note: Patterns of division and release of the larvae form the basis for dividing the subclass into orders. Other characters are useful in
identification, but knowledge of the full life cycle is often required before suctorians can be confidently assigned to a genus.]
2(1) Conspicuous buccal ciliature at apical pole; buccal ciliature winds clockwise toward the center when viewed from oral end; somatic cilia-
ture reduced or absent; mobile or sessile; solitary, gregarious or colonial; some species loricate; oral region can contract and withdraw in
most species; with either scopula or complex adhesive disc (holdfast) at aboral end of cell.............................................................................
......................................................................................... class Oligohymenophorea, subclasses Mobilia and Peritrichia............................. 3
2’ Not as above: without scopula or complex adhesive disc at aboral end of cell............................................................................................... 4
3(2) Trophont mobile; symbiotic on other organisms; with complex adhesive disc (holdfast) at aboral end of cell ................................................
................................................................................................................................................................................... subclass Mobilia [p. 30]
3’ Trophont only rarely motile, usually attached to substratum, which may be inanimate objects or other organisms, via a stalk, lorica or
scopula; many gregarious or colonial species ...................................................................................................... subclass Peritrichia [p. 30]
4(2) Oral area not bordered by an adoral zone of membranelles; no ventral cirri ................................................................................................. 5
4’ Oral area usually bordered by a well-developed adoral zone of membranelles (AZM) consisting of more than three membranelles (polyki-
neties); with or without ventral cirri.............................................................................................................................................................. 11
5(4) Cytostome at end of a buccal cavity or vestibulum with cilia or paroral membrane(s) associated................................................................. 6
5’ Cytostome at or near surface; buccal cavity, if present, without cilia or paroral membranes ........................................................................ 7
6(5) Alveoli well-developed and revealed as a prominent argyrome, typically reticulate; body shape variable, many reniform and flattened........
...................................................................................................................................................................................... class Colpodea [p. 31]
6’ Alveoli not well developed, no prominent argyrome; body typically ovoid to elongate ovoid elongate ovoid ................................................
................................................................................................................................................................... class Oligohymenophorea [p. 31]
7(5) Cytostome lateral or ventral ............................................................................................................................................................................ 8
7’ Cytostome at or near anterior end, or continuing down side as a slit ............................................................................................................. 9
8(7) Circular mouth located midventrally; no proboscis; large cytostome .......................................................................................................... 10

8’ Cytostome a barely visible lateral slit on the convex side of a tapering front end, or a lateral opening at the base of an anterior proboscis
(Fig. 2.9 J; see also family Spathidiidae Fig. 2.10 I, U) ...................................................................... class Karyorelictea, family Loxodidae
9(7) Oral ciliature as simple kinetids; with dorsal brosse or brush formed of specialized dikinetids bearing clavate cilia ......................................
................................................................................................................................................................................ class Litostomatea [p. 33]
9’ With circumoral ciliation composed of dikinetids; without dorsal brosse ............................................................. class Prostomatea [p. 34]
10(8) Body nearly ellipsoid, rounded in cross section, sometimes flattened ventrally; medium to large (some 100 μm); densely ciliated all over;
oral depression present ........................................................................................................................................... class Nassophorea [p. 34]
10’ Body usually flattened; ventrum ciliated, dorsum bare or with a few cilia; anterior preoral arcs of right ventral ciliary rows continuous with
more posterior parts...................................................................................................... class Phyllopharyngea, subclass Cyrtophoria [p. 34]
11(4) Somatic ventral ciliature as polykinetids (cirri) in groups or files on ventral surface................................................................................... 12
11’ Somatic ciliature as monokinetids or dikinetids, but not cirri....................................................................................................................... 14
12(11) Body typically oval to rectangular, sometime ellipsoidal, rigid and dorsoventrally flattened, often heavily ribbed; ventral cirri often con-
spicuous; pellicular alveoli well developed .................................................................................................................................................. 13
12’ Body elongate, often flexible; ventral cirri often inconspicuous; pellicular alveoli poorly developed .............................................................
Protozoa
............................................................................................................................................ class Spirotrichea, subclass Stichotrichia [p. 35]

13(12) Body ovoid with prominent longitudinal ridges on each side; AZM stretching almost entire body length terminating near posterior pole;
ventral ciliature comprising delicate cirri in widely spaced rows (Fig. 2.18 E) ................................................................................................
................................................................................................................... class Spirotrichea, subclass Protocruziidia, family Phacodiniidae
13’ Body ovoid to rectangular or ellipsoidal; AZM not usually stretching to posterior pole .............. class Spirotrichea, subclass Hypotrichia [p. 35]
14(11) Somatic cilia sparse or absent........................................................................................................................................................................ 15
14’ Body surface densely ciliated........................................................................................................................................................................ 16
15(14) Body flattened, rigid, often with spines; body ciliature present as short, generally obvious rows; oral ciliature relatively inconspicuous;
mainly anaerobic........................................................................................................................................................ class Plagiopylea [p. 36]
15’ Membranelles numerous in complete, or almost complete, circle at oral end; body generally conical or bell-shaped; mostly planktonic
........................................................................................................... class Spirotrichea, subclasses Choreotrichia and Oligotrichia [p. 36]
16(14) Body small to medium, usually twisted to left; oral region spiraled with paramembranelles; with hydrogenosomes instead of mitochondria;
in richly organic sediments with low, or no, oxygen.............................................................................................. class Armophorea [p. 36]
16’ Body medium to large, often elongate and contractile; somatic ciliation holotrichous; left oral polykinetids conspicuous, typically para-
membranelles encircling the anterior end clockwise before plunging into the oral cavity; one or more parorals on right side........................
............................................................................................................................................................................... class Heterotrichea [p. 36]
Protozoa: Ciliophora: Suctoria: Orders

1 Budding begins in a pouch .............................................................................................................................................................................. 2
1’ Budding and cytokinesis on surface of trophont ................................................................................................................. Exogenida [p. 28]
2(1) Exogenous budding occurring in a brood pouch; swarmers become free-swimming in pouch before emerging through birth pore; swarmer
small and ciliated ............................................................................................................................................................. Endogenida [p. 29]
2’ Cytokinesis of swarmer completed exogenously, after the emergence of everted bud on cell surface; swarmer often ellipsoidal, flattened
..................................................................................................................................................................................... Evaginogenida [p. 29]
Protozoa: Ciliophora: Suctoria: Exogenida: Families

1 Trophont sac-like or spherical, basally attached to bottom of lorica near junction with stalk........................................................................ 2
1’ Trophont not sac-like, not basally attached to lorica, or without lorica .......................................................................................................... 3
2(1) Tentacles capitate, grouped apically in single fascicle or row; swarmers ovoid with somatic kineties in U-shape around body
(Fig. 2.13 B)................................................................................................................................................................................. Paracinetidae
2’ Tentacles clavate, in a group on narrow, rounded distal end of body; swarmers flattened or vermiform, ciliated on one margin (Fig. 2.14 A)
.................................................................................................................................................................................................... Thecacnetidae
3(1) Without a rod-like tentacle or protuberance of stylotheca............................................................................................................................... 4
3’ Ectoparasitic, attached to host (usually peritrich ciliates) by rod-like tentacle or protuberance of stylotheca (Fig. 2.16 L) ............................
.............................................................................................................................................................................................. Manuelophryidae
4(3) Trophont medium to large, cylindrical, conical or trumpet-shaped; ectocommensal on crustaceans (Fig. 2.18 B) ..........................................
................................................................................................................................................................................................ Spelaeophryidae
4’ Trophont small to medium, spheroid, pyriform or goblet-shaped................................................................................................................... 5
5(4) Lorica with several radial slits in distal half through which tentacles project; sometimes stalked; free-living or ectocommensal on inverte-
brates or other ciliates (Fig. 2.13 C) .......................................................................................................................................... Metacinetidae
5’ Typically aloricate and stalked ........................................................................................................................................................................ 6
6(5) Trophont goblet-shaped or laterally flattened; attached to antennules of harpacticoid copepods (Fig. 2.13 L) .......................... Lecanophryidae
6’ Trophont spheroid or pyriform; planktonic or sessile, often attached to other ciliates as parasites (Fig. 2.14 E, G) ...................... Podophryidae
Protozoa: Ciliophora: Suctoria: Endogenida: Families

1 With stalk......................................................................................................................................................................................................... 2
1’ Without stalk.................................................................................................................................................................................................... 5
2(1) With lorica ....................................................................................................................................................................................................... 3
2’ Without lorica.................................................................................................................................................................................................. 4
Protozoa
3(2) Trophonts laterally flattened, trapezoid, triangular or discoid; lorica often triangular; one type of tentacle arranged in two, rarely three,
fascicles or rows (Fig. 2.14 F).......................................................................................................................................................... Acinetidae
3’ Trophont trapezoid, laterally flattened; loricate; tentacles of two types, i.e., agile prehensile and regular feeding ones; ectosymbionts on
plants and invertebrates (Fig. 2.16 B)......................................................................................................................................... Acinetopsidae
4(2) Trophont globular to ellipsoidal; tentacles funnel-like; attached to cyclopoid crustaceans (Fig. 2.13 D)............................. Choanophryidae
4’ Trophont ovoid, cylindrical or triangular, often flattened; tentacles capitate; free-living (Fig. 2.14 K, L)................................. Tokophryidae
5(1) Without lorica or, if present, lorica mucoid..................................................................................................................................................... 6
5’ Trophont small, spheroid to ovoid; tentacles capitate; with lorica that is attached to substrate by basal surface; in periphyton or plankton
(Fig. 2.13 E) ............................................................................................................................................................................ Solenophryidae
6(5) Tentacles present.............................................................................................................................................................................................. 7
6’ Tentacles absent; trophont ovoid to spheroid; endoparasitic in cells and tissues of hosts such as other ciliates and invertebrates (Fig. 2.13 I)
................................................................................................................................................................................................. Endosphaeridae
7(6) Attached to substratum by basal body surface or protuberance of body......................................................................................................... 8
7’ Attached to substratum by tentacles or cinctum............................................................................................................................................ 10
8(7) Trophont ovoid, pyriform, truncate or branching............................................................................................................................................ 9
8’ Trophont flattened; some species in mucoid lorica; tentacles capitate or rod-like; ectocommensals on invertebrates and vertebrates includ-
ing the gills of fishes (Fig. 2.18 K)........................................................................................................................................... Trichophryidae
9(8) Trophont small, pyriform to ovoid; tentacles agile, very flexible; ectoparasites on crustaceans (Fig. 2.18 J) ........................... Rhynchetidae
9’ Trophont medium to large, pyriform to truncate to branching; tentacles capitate not conspicuously flexible; some free-living, some endo-
symbionts some ectosymbionts on turtles or the gills of crustaceans (Fig. 2.15 A)............................................................. Dendrosomatidae
10(7) Trophonts small to medium, ovoid to irregular; attached to host by arm-like cinctum; hypocommensals on peritrich ectosymbionts of fishes
(Fig. 2.18 C).............................................................................................................................................................................. Erastophryidae
10’ Trophonts small, globular to ellipsoid; with lorica; tentacles rod-like, one to several serving both for feeding and attachment; parasites of
other ciliates (e.g., folliculinids and suctorians) (Fig. 2.16 J) .............................................................................................. Pseudogemmidae
Protozoa: Ciliophora: Suctoria: Evaginogenida: Families

1 Tentacles present; not parasites of rotifers ...................................................................................................................................................... 2
1’ Tentacles absent; trophont small and sac-like; stalk absent; swarmer ellipsoidal, flattened, with several longitudinal kineties; parasites of
tissues of rotifers (Fig. 2.16 K) ............................................................................................................................................. Trypanococcidae
2(1) Tentacles ramified............................................................................................................................................................................................ 3
2’ Tentacles not ramified ..................................................................................................................................................................................... 4
3(2) Trophont vase-like, branched, lifted off substrate by basal protuberance; ectocommensals on gammarid crustaceans (Fig. 2.18 D) .............
.............................................................................................................................................................................................. Cometodendridae
3’ Trophont hemispherical or disc-shaped, unbranched, basal protuberance absent; ectocommensals on gammarid amphipods (Fig. 2.14 I, O)
.............................................................................................................................................................................................. Dendrocometidae
Another random document with
no related content on Scribd:
The Project Gutenberg eBook of Memorials of
Old Dorset
This ebook is for the use of anyone anywhere in the United
States and most other parts of the world at no cost and with
almost no restrictions whatsoever. You may copy it, give it away
or re-use it under the terms of the Project Gutenberg License
included with this ebook or online at www.gutenberg.org. If you
are not located in the United States, you will have to check the
laws of the country where you are located before using this
eBook.
Title: Memorials of Old Dorset
Editor: Rev. Thomas Perkins

Herbert Pentin
Release date: May 19, 2022 [eBook #68128]
Language: English
Original publication: United Kingdom: Bemrose and Sons, 1907
Credits: Tim Lindell, Les Galloway and the Online Distributed

Proofreading Team at https://www.pgdp.net (This file
was produced from images generously made available
by The Internet Archive/Canadian Libraries)
*** START OF THE PROJECT GUTENBERG EBOOK MEMORIALS

OF OLD DORSET ***
Transcriber’s Notes
Obvious typographical errors have been silently corrected. Variations in hyphenation

and accents have been standardised but all other spelling and punctuation remains
unchanged.
The repetition of the title immediately before the title page has been removed.
Ringstead and Holworth.
“Where one may walk along the undulating downs that skirt the Channel, held in
place by parapets of cliff that break down straight into the sea; where one may walk
mile after mile on natural lawn and not meet a soul—just one’s self, the birds, the
glorious scenery, and God.” (See page 109.)
From a water-colour sketch by Mr. William Pye.
Memorials of the Counties of England
General Editor: Rev. P. H. Ditchfield, M.A., F.S.A.
MEMORIALS
OF OLD DORSET
EDITED BY
THOMAS PERKINS, M.A.
Late Rector of Turnworth, Dorset
Author of
“Wimborne Minster and Christchurch Priory”

“Bath and Malmesbury Abbeys” “Romsey Abbey” &c.
AND
HERBERT PENTIN, M.A.
Vicar of Milton Abbey, Dorset
Vice-President, Hon. Secretary, and Editor

of the Dorset Natural History and Antiquarian Field Club
With many Illustrations
LONDON
BEMROSE & SONS LIMITED, 4 SNOW HILL, E.C.
AND DERBY
1907
[All Rights Reserved]
TO
THE RIGHT HONOURABLE
LORD EUSTACE CECIL, F.R.G.S.
PAST PRESIDENT OF THE DORSET NATURAL
HISTORY AND ANTIQUARIAN FIELD CLUB
THIS BOOK IS DEDICATED
BY HIS LORDSHIP’S
KIND
PERMISSION
PREFACE
he editing of this Dorset volume was originally
undertaken by the Rev. Thomas Perkins, the scholarly
Rector of Turnworth. But he, having formulated its plan
and written four papers therefor, besides gathering
material for most of the other chapters, was laid aside
by a very painful illness, which culminated in his unexpected death.
This is a great loss to his many friends, to the present volume, and to
the county of Dorset as a whole; for Mr. Perkins knew the county as
few men know it, his literary ability was of no mean order, and his
kindness to all with whom he was brought in contact was proverbial.
After the death of Mr. Perkins, the editing of the work was
entrusted to the Rev. Herbert Pentin, Vicar of Milton Abbey, whose
knowledge of the county and literary experience as Editor of the
Dorset Natural History and Antiquarian Field Club enabled him to
gather up the threads where his friend Mr. Perkins had been
compelled to lay them down, and to complete the work and see it
safely through the press. As General Editor of the series, I desire to
express my most grateful thanks to him for his kind and gracious
services in perfecting a work which had unfortunately been left
incomplete; and all lovers of Old Dorset and readers of this book will
greatly appreciate his good offices.
Few counties can rival Dorset either in natural beauty or historic
interest, and it deserves an honoured place among the memorials of
the counties of England. In preparing the work the Editors have
endeavoured to make the volume comprehensive, although it is of
course impossible in a single volume to exhaust all the rich store of
historical treasures which the county affords. After a general sketch
of the history of Dorset by the late Editor, the traces of the earliest
races which inhabited this county are discussed by Mr. Prideaux,
who tells of the ancient barrows in Dorset, and the details of the
Roman occupation are shown by Captain Acland. Dorset is rich in
churches, and no one was more capable to describe their chief
features than Mr. Perkins. His chapter is followed by others of more
detail, dealing with the three great minsters still standing—
Sherborne, Milton, and Wimborne, the monastic house at Ford, and
the memorial brasses of Dorset. A series of chapters on some of the
chief towns and “islands” of the county follows, supplemented by a
description of two well-known manor-houses. The literary
associations of the county and some of its witchcraft-superstitions
form the subjects of the concluding chapters. The names of the able
writers who have kindly contributed to this volume will commend
themselves to our readers. The Lord Bishop of Durham, the Rev. R.
Grosvenor Bartelot, Mr. Sidney Heath, Mr. Wildman, Mr. Prideaux,
Mr. Gill, Mrs. King Warry, and our other contributors, are among the
chief authorities upon the subjects of which they treat, and our
thanks are due to them for their services; and also to Mr. William Pye
for the beautiful coloured frontispiece, to Mr. Heath for his charming
drawings, and to those who have supplied photographs for
reproduction. We hope that this volume will find a welcome in the
library of every Dorset book-lover, and meet with the approbation of
all who revere the traditions and historical associations of the county.
P. H. Ditchfield,
General Editor.
CONTENTS
Page
Historic Dorset By the Rev. Thomas Perkins, M.A. 1
The Barrows of Dorset By C. S. Prideaux 19
The Roman Occupation of
By Captain J. E. Acland
Dorset 28
The Churches of Dorset By the Rev. Thomas Perkins, M.A. 44
The Memorial Brasses of Dorset By W. de C. Prideaux 62
Sherborne By W. B. Wildman, M.A. 75
Milton Abbey By the Rev. Herbert Pentin, M.A. 94
Wimborne Minster By the Rev. Thomas Perkins, M.A. 117
Ford Abbey By Sidney Heath 131
Dorchester By the Lord Bishop of Durham, D.D. 145
Weymouth By Sidney Heath 157
The Isle of Portland By Mrs. King Warry 177
The Isle of Purbeck By A. D. Moullin 187
Corfe Castle By Albert Bankes 200
Poole By W. K. Gill 222
Bridport By the Rev. R. Grosvenor Bartelot,
M.A. 232
Shaftesbury By the Rev. Thomas Perkins, M.A. 240
Piddletown and Athelhampton By Miss Wood Homer 257
Wolfeton House By Albert Bankes 264
The Literary Associations of
By Miss M. Jourdain
Dorset 273
Some Dorset Superstitions By Hermann Lea 292
Index 307
INDEX TO ILLUSTRATIONS
Ringstead and Holworth Frontispiece
(From a water-colour sketch by Mr. William Pye)
Page, or
Facing Page
Bronze Age Objects from Dorset Round Barrows 20
(From photographs by Mr. W. Pouncy)
Part of the Olga Road Tessellated Pavement, Dorchester 38
(From a photograph by the Rev. T. Perkins)
Tessellated Pavement at Fifehead Neville 41
St. Martin’s Church, Wareham 48
The Chapel on St. Ealdhelm’s Head 50
(From a drawing by Mr. Sidney Heath)
Brass to William Grey, Rector of Evershot 70
(From a rubbing by Mr. W. de C. Prideaux)
Sherborne Abbey 76
The Entrance to Sherborne School 86
Milton Abbey 94
The Paintings in Milton Abbey 95
Milton Abbey: Interior 96
(From a photograph by Mr. S. Gillingham)
The Tabernacle in Milton Abbey 97
”””
Abbot Middleton’s Rebus 101
St. Catherine’s Chapel, Milton Abbey 104
Holworth Burning Cliff in 1827 106
(From a coloured print by Mr. E. Vivian)
Liscombe Chapel 107
Milton Abbey in the year 1733 110
(From an engraving by Messrs. S. and N. Buck)
The Seal of the Town of Milton in America 116
Wimborne Minster 118
The Chained Library, Wimborne Minster 128
Ford Abbey 132
Details from Cloisters, Ford Abbey 134
(From drawings by Mr. Sidney Heath)
The Chapel, Ford Abbey 136
Panel from Cloisters, Ford Abbey 136
The Seal of Ford Abbey 140
High Street, Dorchester 146
Judge Jeffreys’ Lodgings, Dorchester 149
Cornhill, Dorchester 153
“Napper’s Mite,” Dorchester 155
”””
The Quay, Weymouth 158
”””
Chest in the Guildhall, Weymouth 164
”””
Sandsfoot Castle, Weymouth 166
”””
Doorway, Sandsfoot Castle 167
”””
Some Weymouth Tokens 169
”””
The Arms of Weymouth 170
”””
Old House on North Quay, Weymouth 171
An Old Chair in the Guildhall, Weymouth 172
The Old Stocks, Weymouth 176
Portland Cottages 185
”””
“Kimmeridge Coal Money” 192
(From a photograph by Mr. A. D. Moullin)
Corfe Castle 200
The Town Cellars, Poole 222
”””
Shaftesbury 240
”””
Gold Hill, Shaftesbury 248
”””
Piddletown Church 258
”””
Athelhampton Hall 262
”””
Wolfeton House 264
”””
The East Drawing Room, Wolfeton House 268
William Barnes 280
(From a photograph by Messrs. Dickinsons)
Thomas Hardy 284
Came Rectory 291
HISTORIC DORSET
By the Rev. Thomas Perkins, M.A.
HE physical features due to the geological formation of

the district now called Dorset have had such an
influence on the inhabitants and their history that it
seems necessary to point out briefly what series of
stratified rocks may be seen in Dorset, and the lines of
their outcrop.
There are no igneous rocks, nor any of those classed as primary,
but, beginning with the Rhætic beds, we find every division of the
secondary formations, with the possible exception of the Lower
Greensand, represented, and in the south-eastern part of the district
several of the tertiary beds may be met with on the surface.
The dip of the strata is generally towards the east; hence the
earlier formations are found in the west. Nowhere else in England
could a traveller in a journey of a little under fifty miles—which is
about the distance from Lyme to the eastern boundary of Dorset—
cross the outcrop of so many strata. A glance at a geological map of
England will show that the Lias, starting from Lyme Regis, sweeps
along a curve slightly concave towards the west, almost due north,
until it reaches the sea again at Redcar, while the southern boundary
of the chalk starting within about ten miles of Lyme runs out
eastward to Beechy Head. Hence it is seen that the outcrops of the
various strata are wider the further away they are from Lyme Regis.
Dorset has given names to three well-known formations and to
one less well known: (1) The Portland beds, first quarried for building
stone about 1660; (2) the Purbeck beds, which supplied the Early
English church builders with marble for their ornamental shafts; (3)
Kimmeridge clay; and (4) the Punfield beds.
The great variety of the formation coming to the surface in the
area under consideration has given a striking variety to the character
of the landscape: the chalk downs of the North and centre, with their
rounded outlines; the abrupt escarpments of the greensand in the
neighbourhood of Shaftesbury; the rich grazing land of Blackmore
Vale on the Oxford clay; and the great Heath (Mr. Hardy’s Egdon)
stretching from near Dorchester out to the east across Woolwich,
Reading, and Bagshot beds, with their layers of gravel, sand, and
clay. The chalk heights are destitute of water; the streams and rivers
are those of the level valleys and plains of Oolitic clays—hence they
are slow and shallow, and are not navigable, even by small craft, far
from their mouths.
The only sides from which in early days invaders were likely to
come were the south and east; and both of these boundaries were
well protected by natural defences, the former by its wall of cliffs and
the deadly line of the Chesil beach. The only opening in the wall was
Poole Harbour, a land-locked bay, across which small craft might
indeed be rowed, but whose shores were no doubt a swamp
entangled by vegetation. Swanage Bay and Lulworth Cove could
have been easily defended. Weymouth Bay was the most vulnerable
point. Dense forests protected the eastern boundary. These natural
defences had a marked effect, as we shall see, on the history of the
people. Dorset for many centuries was an isolated district, and is so
to a certain extent now, though great changes have taken place
during the last fifty or sixty years, due to the two railways that carry
passengers from the East to Weymouth and the one that brings
them from the North to Poole and on to Bournemouth. This isolation
has conduced to the survival not only of old modes of speech, but
also of old customs, modes of thought, and superstitions.
It may be well, before speaking of this history, to state that the
county with which this volume deals should always be spoken of as
“Dorset,” never as “Dorsetshire”; for in no sense of the word is
Dorset a shire, as will be explained further on.
We find within the boundaries of the district very few traces of
Palæolithic man: the earliest inhabitants, who have left well-marked
memorials of themselves, were Iberians, a non-Aryan race, still
represented by the Basques of the Pyrenees and by certain
inhabitants of Wales. They were short of stature, swarthy of skin,
dark of hair, long-skulled. Their characteristic weapon or implement
was a stone axe, ground, not chipped, to a sharp edge; they buried
their dead in a crouching attitude in the long barrows which are still
to be seen in certain parts of Dorset, chiefly to the north-east of the
Stour Valley. When and how they came into Britain we cannot tell for
certain; it was undoubtedly after the glacial epoch, and probably at a
time when the Straits of Dover had not come into being and the
Thames was still a tributary of the Rhine. They were in what is
known as the Neolithic stage of civilisation; but in course of time,
after this country had become an island, invaders broke in upon
them, Aryans of the Celtic race, probably (as Professor Rhys thinks,
though he says he is not certain on this point) of the Goidelic branch.
These men were tall, fair-haired, blue-eyed, round-skulled, and were
in a more advanced stage of civilisation than the Iberians, using
bronze weapons, and burying their dead, sometimes after cremation,
in the round barrows that exist in such large numbers on the Dorset
downs. Their better arms and greater strength told in the warfare that
ensued: whether the earlier inhabitants were altogether destroyed, or
expelled or lived on in diminished numbers in a state of slavery, we
have no means of ascertaining. But certain it is that the Celts
became masters of the land. These men were some of those who
are called in school history books “Ancient Britons”; the Wessex folk
in after days called them “Welsh”—that is, “foreigners”—the word
that in their language answered to βάρβαροι and “barbari” of the
Greeks and Romans. What they called themselves we do not know.
Ptolemy speaks of them as “Durotriges,” the name by which they
were known to the Romans. Despite various conjectures, the
etymology of this word is uncertain. The land which they inhabited
was, as already pointed out, much isolated. The lofty cliffs from the
entrance to Poole Harbour to Portland formed a natural defence;
beyond this, the long line of the Chesil beach, and further west, more
cliffs right on to the mouth of the Axe. Most of the lowlands of the
interior were occupied by impenetrable forests, and the slow-running
rivers, which even now in rainy seasons overflow their banks, and
must then, when the rainfall was much heavier than now, have
spread out into swamps, rendered unnavigable by their thick tangle
of vegetation. The inhabitants dwelt on the sloping sides of the
downs, getting the water they needed from the valleys, and retiring
for safety to the almost innumerable encampments that crowned the
crests of the hills, many of which remain easily to be distinguished to
this day. Nowhere else in England in an equal area can so many
Celtic earthworks be found as in Dorset. The Romans came in due
course, landing we know not where, and established themselves in
certain towns not far from the coasts.
The Celts were not slain or driven out of their land, but lived on
together with the Romans, gradually advancing in civilisation under
Roman influence. They had already adopted the Christian religion:
they belonged to the old British Church, which lived on in the south-
west of England even through that period when the Teutonic
invaders—Jutes, Angles, Saxons—devastated the south-east, east,
north, and central parts of the island, and utterly drove westward
before them the Celtic Christians into Wales and the south-west of
Scotland. Dorset remained for some time untouched, for though the
Romans had cleared some of the forests before them, and had cut
roads through others, establishing at intervals along them military
stations, and strengthening and occupying many of the Celtic camps,
yet the vast forest—“Selwood,” as the English called it—defended
Dorset from any attack of the West Saxons, who had settled further
to the east. Once, and once only, if we venture, with Professor
Freeman, to identify Badbury Rings, near Wimborne, on the Roman
Road, with the Mons Badonicus of Gildas, the Saxons, under Cerdic,
in 516, invaded the land of the Durotriges, coming along the Roman
Road which leads from Salisbury to Dorchester, through the gap in
the forest at Woodyates, but found that mighty triple ramparted
stronghold held by Celtic Arthur and his knights, round whom so
much that is legendary has gathered, but who probably were not
altogether mythical. In the fight that followed, the Christian Celt was
victorious, and the Saxon invader was driven in flight back to his own
territory beyond Selwood. Some place Mons Badonicus in the very
north of England, or even in Scotland, and say that the battle was
fought between the Northumbrians and the North Welsh: if this view
is correct, we may say that no serious attack was made on the Celts
of Dorset from the east. According to Mr. Wildman’s theory, as stated
in his Life of St. Ealdhelm—which theory has a great air of probability
about it—the Wessex folk, under Cenwealh, son of Cynegils, the first
Christian King of the West Saxons, won two victories: one at
Bradford-on-Avon in 652, and one at the “Hills” in 658. Thus North
Dorset was overcome, and gradually the West Saxons passed on
westward through Somerset, until in 682 Centwine, according to the
English Chronicle, drove the Welsh into the sea. William of
Malmesbury calls them “Norht Walæs,” or North Welsh, but this is
absurd: Mr. Wildman thinks “Norht” may be a mistake for “Dorn,” or
“Thorn,” and that the Celts of Dorset are meant, and that the sea
mentioned is the English Channel. From this time the fate of the
Durotriges was sealed: their land became part of the great West
Saxon kingdom. Well indeed was it for them that they had remained
independent until after the time when their conquerors had ceased to
worship Woden and Thunder and had given in their allegiance to the
White Christ; for had these men still been worshippers of the old
fierce gods, the Celts would have fared much worse. Now, instead of
being exterminated, they were allowed to dwell among the West
Saxon settlers, in an inferior position, but yet protected by the West
Saxon laws, as we see from those of Ine who reigned over the West
Saxons from 688 to 728. The Wessex settlers in Dorset were called
by themselves “Dornsæte,” or “Dorsæte,” whence comes the name
of Dorset. It will be seen then, that Dorset is what Professor
Freeman calls a “ga”—the land in which a certain tribe settled—and
differs entirely from those divisions made after the Mercian land had
been won back from the Danes, when shires were formed by
shearing up the newly recovered land, not into its former divisions
which the Danish conquest had obliterated, but into convenient
portions, each called after the name of the chief town within its
borders, such as Oxfordshire from Oxford, Leicestershire from
Leicester. The Danes did for a time get possession of the larger part
of Wessex, but it was only for a time: the boundaries of Dorset were
not wiped out, and there was no need to make any fresh division. So
when we use the name Dorset for the county we use the very name

Get Thorp and Covich's Freshwater Invertebrates: Keys To Nearctic Fauna 4th Edition James H. Thorp Free All Chapters

Uploaded by

Copyright:

Available Formats

Get Thorp and Covich's Freshwater Invertebrates: Keys To Nearctic Fauna 4th Edition James H. Thorp Free All Chapters

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Get Thorp and Covich's Freshwater Invertebrates: Keys To Nearctic Fauna 4th Edition James H. Thorp Free All Chapters

Uploaded by

Copyright:

Available Formats

Full download test bank at ebook ebookmass.

Thorp and Covich's Freshwater

CLICK LINK TO DOWLOAD

Thorp and Covich's Freshwater Invertebrates: Keys to

Thorp and Covich's Freshwater Invertebrates: Keys to

Thorp and Covich's Freshwater Invertebrates: Keys to

Introduction to Econometrics 3rd Edition James H. Stock

Business and Professional Communication: KEYS for

Probability and Statistics (4th Edition) Morris H.

Invertebrates: Fourth Edition Richard C. Brusca

Chaos and Nonlinear Psychology : Keys to Creativity in

A Global Series of Books on the Identification,

Volume I: Ecology and General Biology

Volume II: Keys to Nearctic Fauna

Volume III: Keys to Palaearctic Fauna

Volumes in Preparation and Under Contract

Possible Future Volumes of the Fourth Edition

AMSTERDAM • BOSTON • HEIDELBERG • LONDON • NEW YORK • OXFORD • PARIS

Copyright © 2016 Elsevier Inc. All rights reserved.

British Library Cataloguing-in-Publication Data

Library of Congress Cataloging-in-Publication Data

For information on all Academic Press publications

Publisher: Cathleen Sether

Typeset by TNQ Books and Journals

To Henry B. Ward, George C. Whipple, W. Thomas Edmondson, and

field and lab. While his research emphasizes aquatic inver-

Dr. James H. Thorp has been a Professor in the Depart-

Thorp and Covich’s Freshwater Invertebrates. http://dx.doi.org/10.1016/B978-0-12-385028-7.00001-9

Freshwater Invertebrate Kingdoms and Phyla

INTRODUCTION amoebozoans, opisthokonts, rhizarians, and excavates

Thorp and Covich’s Freshwater Invertebrates. http://dx.doi.org/10.1016/B978-0-12-385028-7.00002-0

species, which is sometimes a prerequisite for adequate

illustrations used as examples here are of one or more spe-

FIGURE 2.1 Examples of the main functional groups of protozoa.

(A) (B) (C)

(E) (F) (G)

(C) (D) (E)

Amoebae simultaneously, as in Amoeba (Figs. 2.1 B and 2.4 F), or

(G) (H) (I)

(J) (K) (L) (M) (N)

FIGURE 2.6 (A) Vampyrella lateritia;

FIGURE 2.7 (A) Clathrulina elegans;

(F) (G) (H) (I) (J)

(R) (S) (T)

(B) (C) (D) (E)

(A) (G) (H) (I) (J)

(Q) (R) (V) (W)

(D) (E) (F)

(R) (S) (T) (V)

wheat or rice, which will promote the growth of bacteria and

(H) (I) (J)

(M) (N) (O)

Key to Major Groups of Flagellated Protozoa

5(4) Cells lacking the characteristics described below ........................................................................................................................................... 6

Key to Major Groups of Amoeboid Protozoa

Protozoa: Amoeboid Protozoa: Heterolobosea: Schizopyrenida: Families

Protozoa: Amoeboid Protozoa: Lobosea: Families

Protozoa: Amoeboid Protozoa: Cercozoa: Subphyla

Protozoa: Amoeboid Protozoa: Cercozoa: Filosa: Families

5(4) Cells lacking the characteristics described below ........................................................................................................................................... 6

8(7) Circular mouth located midventrally; no proboscis; large cytostome .......................................................................................................... 10