Thorp and Covich's Freshwater

Invertebrates: Keys to Palaearctic

Fauna 4th Edition James H. Thorp
Visit to download the full and correct content document:
https://ebookmass.com/product/thorp-and-covichs-freshwater-invertebrates-keys-to-p
alaearctic-fauna-4th-edition-james-h-thorp/
More products digital (pdf, epub, mobi) instant
download maybe you interests ...

Thorp and Covich's Freshwater Invertebrates: Keys to

Nearctic Fauna 4th Edition James H. Thorp

https://ebookmass.com/product/thorp-and-covichs-freshwater-
invertebrates-keys-to-nearctic-fauna-4th-edition-james-h-thorp/

Thorp and Covich's Freshwater Invertebrates: Keys to

Neotropical and Antarctic Fauna 4th Edition Cristina
Damborenea

https://ebookmass.com/product/thorp-and-covichs-freshwater-
invertebrates-keys-to-neotropical-and-antarctic-fauna-4th-
edition-cristina-damborenea/

Thorp and Covich's Freshwater Invertebrates: Keys to

Neotropical Hexapoda 4th Edition Neusa Hamada

https://ebookmass.com/product/thorp-and-covichs-freshwater-
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/
Keys to Palaearctic Fauna
 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 Neotropical Hexapoda

Edited by Neusa Hamada, James H. Thorp, and D. Christopher Rogers
Published 2018

Volume IV: Keys to Palaearctic Fauna

Edited by D. Christopher Rogers and James H. Thorp
Published 2019

Volumes in Preparation for Future Publication

Keys to Neotropical and Antarctic Fauna (2019)
Keys to Australasian Fauna (2020)
Keys to Afrotropical Fauna (2023)

Planned Future Volumes

Keys to Oriental and Oceana Hexapoda
Keys to Oriental and Oceana Fauna
Keys to Palaearctic Hexapoda
Keys to Nearctic Hexapoda

Related Publications
Field Guide to Freshwater Invertebrates of North America
James H. Thorp and D. Christopher Rogers
First Edition (2011)
Keys to Palaearctic Fauna
Thorp and Covich’s Freshwater
Invertebrates e Volume IV

Fourth Edition

Edited by

D. Christopher Rogers
Kansas Biological Survey and The Biodiversity Institute
University of Kansas, Lawrence, KS, USA

James H. Thorp
Kansas Biological Survey and Department of Ecology and Evolutionary Biology
University of Kansas, Lawrence, KS, USA
Academic Press is an imprint of Elsevier
125 London Wall, London EC2Y 5AS, United Kingdom
525 B Street, Suite 1650, San Diego, CA 92101, United States
50 Hampshire Street, 5th Floor, Cambridge, MA 02139, United States
The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, United Kingdom

Copyright Ó 2019 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.

Library of Congress Cataloging-in-Publication Data

A catalog record for this book is available from the Library of Congress

British Library Cataloguing-in-Publication Data

A catalogue record for this book is available from the British Library

ISBN: 978-0-12-385024-9

For information on all Academic Press publications visit our website at

https://www.elsevier.com/books-and-journals

Publisher: Candice Janco

Acquisitions Editor: Louisa Munro
Editorial Project Manager: Emily Thomson
Production Project Manager: Paul Prasad Chandramohan
Cover Designer: Greg Harris
Cover Photographs courtesy of (from top): Vadim Takhteev, Rafael Araujo, and Gintautas Steiblys

Typeset by TNQ Technologies

 Dedications From the Editors

To our authors who made this book possible.

D. Christopher Rogers

To my loving wife of nearly half a century who has tolerated my occasional

frustrations with my profession while showing great interest in my studies,
and to the students working in my lab over the decades who have never
failed to inspire and challenge me while reinforcing in my mind the
true value of being a professor.
James H. Thorp
This page intentionally left blank
 Contents

Contributors to Volume IV xiii Terminology and Morphology 44

About the Editors xvii Material Preparation and Preservation 45
Preface to the Fourth Edition xix Acknowledgments 45
Preface to Volume IV xxi Keys to Spongillida 45
Acknowledgments for Volume IV xxiii References 91

4. Phylum Cnidaria
1. Introduction
Thomas Jankowski, Boris Anokhin
James H. Thorp, D. Christopher Rogers
Introduction 93
Introduction to This Series, Volume, Limitations 93
and Chapter 1 Terminology and Morphology 94
Nature and Fauna of the Palaearctic Material Preparation and Preservation 96
Bioregion 2 Keys to Cnidaria 97
Components of Taxonomic Chapters 2 References 110
How to Use This Volume 2
Keys to Major Taxa of Inland Water 5. Phylum Platyhelminthes
Invertebrates of the Palaearctic 3
Carolina Noreña, Andrey Porfiriev,
References 4
Oleg Timoshkin
Introduction 113
Part I Limitations 114
Terminology and Morphology 114
2. Protozoa Abbreviations Used in Figures 116
Material Preparation and
Alan Warren, Genoveva F. Esteban
Preservation 117
Introduction 8 Keys to Platyhelminthes 118
Limitations 8 Acknowledgments 144
Terminology and Morphology 8 References 144
Material Preparation and Preservation 17
Acknowledgments 27 6. Phylum Nemertea
Keys to Protozoa 28
Malin Strand, Per Sundberg
References 42
Introduction 145
3. Phylum Porifera Limitations 145
Terminology and Morphology 145
Renata Manconi, Roberto Pronzato
Material Preparation and Preservation 146
Introduction 43 Keys to Nemertea 146
Limitations 43 References 147

vii
viii Contents

7. Phylum Gastrotricha Class Gastropoda 310

Maxim V. Vinarski
Maria Balsamo, Jean-Loup d’Hondt, Paolo Grilli
Introduction 310
Introduction 149 Limitations 311
Limitations 150 Terminology and Morphology 311
Terminology and Morphology 150 Material Preparation and Preservation 314
Material Preparation and Preservation 150 Acknowledgments 316
Keys to Gastrotricha 151 Keys to Gastropoda 323
References 218 References 343

8. Phylum Rotifera Class Bivalvia 346

Robert L. Wallace, T.W. Snell, E.J. Walsh, Rafael Araujo
S.S.S. Sarma, Hendrik Segers Introduction 346
Introduction 219 Limitations 346
Limitations 221 Terminology and Morphology 346
Terminology and Morphology 221 Material Preparation and Preservation 347
Material Preparation and Preservation 223 Keys to Bivalvia 348
Key to Rotifers 224 References 355
References 264
12. Phylum Annelida
9. Phylum Nematoda Introduction to the Phylum 360
Wilfrida Decraemer, Ursula Eisendle-Flöckner, James H. Thorp, Lawrence L. Lovell
Eyualem Abebe Introduction 360
Introduction 270 Limitations 360
Limitations 270 Terminology and Morphology 360
Terminology and Morphology 271 Material Preparation and Preservation 360
Material Preparation and Preservation 274 Keys to Annelida 364
Keys to Nematoda 276 References 364
References 298
Class Clitellata: Subclass Oligochaeta 364
10. Phylum Nematomorpha Tarmo Timm, Patrick Martin
Introduction 365
Andreas Schmidt-Rhaesa
Limitations 365
Introduction 301 Terminology and Morphology 365
Limitations 301 Material Preparation and Preservation 367
Terminology and Morphology 302 Keys to Oligochaeta 369
Material Preparation and Preservation 303 References 482
Keys to Gordiida 303
References 307 Subclass Branchiobdellida 483
Stuart R. Gelder
11. Phylum Mollusca Introduction 483
Introduction to the Phylum 310 Limitations 483
D. Christopher Rogers
Terminology and Morphology 483
Material Preparation and Preservation 484
Introduction 310 Keys to Branchiobdellida 487
Limitations 310 References 490
Material Preparation and Preservation 310
Keys to Mollusca 310
Reference 310
Contents ix

Subclass Hirudinida 491 15. Phylum Tardigrada

Fredric R. Govedich, William E. Moser,
Diane R. Nelson, Roberto Guidetti,
Takafumi Nakano, Aleksander Bielecki,
Lorena Rebecchi
Bonnie A. Bain, Serge Utevsky
Introduction 491 Introduction 533
Terminology and Morphology 492 Limitations 535
Material Preparation and Preservation 494 Terminology and Morphology 535
Keys to Leeches 495 Material Preparation and Preservation 540
References 507 Keys to Tardigrada 544
References 547
Subclass Acanthobdella 507
Class Aphanoneura 508 Part II
João Gil, Christopher J. Glasby, Daniel Martin
Introduction 508 16. Phylum Arthropoda: Introduction and
Limitations 508 Arachnida
Terminology and Morphology 508
Introduction to Arthropoda 551
Material Preparation and Preservation 508
Keys to Aphanoneura 508 D. Christopher Rogers
References 511 Introduction 551
Terminology and Morphology 551
Class Polychaeta 512 Keys to Arthropoda 552
João Gil, Christopher J. Glasby, Daniel Martin References 553
Introduction 512
Limitations 513 Arachnida: Acari: Trombidiformes:
Terminology and Morphology 513 Halacaridae 554
Material Preparation and Preservation 514 Ilse Bartsch
Keys to Polychaeta 515 Introduction 554
References 517 Terminology and Morphology 554
Material Preparation and Preservation 554
13. Phylum Ectoprocta Keys to Halacaridae 555
Timothy S. Wood References 562

Introduction 519 16.1. Arthropoda: Introduction to Crustacea

Limitations 519 and Hexapoda
Terminology and Morphology 520
Material Preparation and Preservation 521 Introduction to the Subphylum Crustacea 565
Keys to Ectoprocta (Bryozoa) 525 D. Christopher Rogers
References 529
Introduction 565
Terminology and Morphology 565
14. Phylum Entoprocta Keys to Crustacea 566
Timothy S. Wood References 566

Introduction 531
Terminology and Morphology 531
Material Preparation and Preservation 532
References 532
x Contents

Crustacea: Hexapoda 566 Thecostraca: Subclass Sessilia 782

R. Edward DeWalt, Vincent H. Resh Robert J. Van Syoc
Introduction 566 Introduction 782
Limitations 567 Terminology and Morphology 782
Terminology and Morphology 567 Material Preparation and Preservation 783
Material Preparation and Preservation 567 Key to Subclass Sessilia 786
Acknowledgments 570 References 787
Keys to Hexapoda 570
References 639 16.6. Phylum Arthropoda: Malacostraca
Introduction to Malacostraca 791
16.2. Arthropoda: Class Branchiopoda
D. Christopher Rogers
D. Christopher Rogers, Alexey A. Kotov,
Introduction 791
Artem Y. Sinev, Sergei M. Glagolev,
Keys to Malacostraca 791
Nikolai M. Korovchinsky, Nikolai N. Smirnov,
Eugeniya I. Bekker
Malacostraca: Cumacea 799
Introduction 644 Les Watling
Limitations 644 Introduction 799
Terminology and Morphology 645 Limitations 799
Material Preparation and Preservation 645 Terminology and Morphology 799
Keys to Branchiopoda 646 Material Preparation and Preservation 799
References 724 Keys to Cumacea 800
References 801
16.3. Arthropoda: Ostracoda
David J. Horne, Claude Meisch, Malacostraca: Bathynellacea 802
Koen Martens D. Christopher Rogers

Introduction 725 Introduction 802

Limitations 726 Limitations 802
Terminology and Morphology 726 Terminology and Morphology 802
Material Preparation and Preservation 726 Material Preparation and Preservation 802
Acknowledgments 727 Keys to Bathynellacea 803
Keys to Ostracoda 728 References 808
References 760
Malacostraca: Amphipoda 808
16.4. Arthropoda: Copepoda Boris Sket, Hiroshi Morino, Vadim Takhteev,
D. Christopher Rogers
Dong Ju Lee, Wonchoel Lee
Introduction 808
Introduction 761 Limitations 808
Limitations 761 Terminology and Morphology 809
Terminology and Morphology 761 Material Preparation and Preservation 809
Material Preparation and Preservation 762 Acknowledgments 809
Keys to Copepoda 766 Keys to Amphipoda 809
References 780 Ponto-Caspian Region Amphipoda 823
Lake Baikal Region Amphipoda 825
16.5. Arthropoda: Thecostraca References 836

Introduction to Thecostraca 781

Malacostraca: Tanaidacea 836
D. Christopher Rogers
Tom Hansknecht
Introduction 781
Introduction 836
Keys to Class Thecostraca 781
Limitations 836
Reference 782
Terminology and Morphology 836
Contents xi

Material Preparation and Preservation 836 Malacostraca: Decapoda 845

Keys to Tanaidacea 837 Sebastian Klaus, Ngan Kee Ng, Sameer Kumar Pati,
References 837 Tohru Naruse, Dirk Brandis, D. Christopher Rogers,
Darren C.J. Yeo
Malacostraca: Isopoda 837
Introduction 846
D. Christopher Rogers, Julian J. Lewis Limitations 846
Introduction 837 Terminology and Morphology 846
Limitations 838 Material Preparation and Preservation 847
Terminology and Morphology 838 Keys to Decapoda 847
Material Preparation and Preservation 838 References 865
Acknowledgments 840
Keys to Isopoda 840 Malacostraca: Mysida and Stygiomysida 866
References 844 Mikhail E. Daneliya, Victor V. Petryashov,
Risto Väinöla
Malacostraca: Thermosbanacea 844
Introduction 866
D. Christopher Rogers Limitations 867
Introduction 844 Terminology and Morphology 867
Limitations 844 Material Preparation and Preservation 868
Terminology and Morphology 844 Keys to Mysida and Stygiomysida 868
Material Preparation and Preservation 845 References 889
Keys to Thermosbaenacea 845
Reference 845
Taxonomic Index 891
This page intentionally left blank
 Contributors to Volume IV

Eyualem Abebe [Chapter 9] Department of Natural Survey, Champaign, IL, United States; email:
Sciences, Pharmacy and Health Professions, Elizabeth edewalt@inhs.illinois.edu
State University, Elizabeth City, NC, United States; Jean-Loup d’Hondt [Chapter 7] Département “Milieux et
email: ebabebe@mail.ecsu.edu Peuplements Aquatiques”, Muséum National
Boris Anokhin [Chapter 4] Zoological Institute RAS, Uni- d’Histoire Naturelle, Paris, France; email:
versitetskaya nab. 1, St. Petersburg, Russia; email: dhondt@mnhn.fr
cnidaria@nm.ru Ursula Eisendle-Flöckner [Chapter 9] Department of
Rafael Araujo [Chapter 11 (Bivalvia)] Museo Nacional de Cell Biology and Physiology, University of Salzburg,
Ciencias Naturales, Madrid, Spain; email: Salzburg, Austria; email: Ursula.Eisendle@sbq.ac.at
rafael@mncn.csic.es Genoveva F. Esteban [Chapter 2] Bournemouth Univer-
Bonnie A. Bain [Chapter 12 (Hirudinida)] Department of sity, Faculty of Science and Technology, Department
Biological Sciences, Southern Utah University, Cedar of Life and Environmental Sciences, Dorset, United
City, UT, United States; email: bain@dixie.edu Kingdom; email: gesteban@bournemouth.ac.uk
Maria Balsamo [Chapter 7] Department of Biomolecular Stuart R. Gelder [Chapter 12 (Branchiobdellida)] Depart-
Sciences (DISB), University of Urbino, “Carlo Bo”, ment of Science and Math, University of Maine at Pre-
Urbino, Italy; email: maria.balsamo@uniurb.it sque Isle, Presque Isle, ME, United States; email:
Ilse Bartsch [Chapter 16] Forschungsinstitut Senckenberg, stuart.gelder@umpi.edu
Hamburg, Germany; email: bartsch@meeresforschung.de João Gil [Chapter 12 (Aphanoneura; Polychaeta)]
Eugeniya I. Bekker [Chapter 16.2] A. N. Severtsov Insti- CCMAR, Universidade do Algarve, Faro, Portugal;
tute of Ecology and Evolution, Moscow, Russia; email: email: joaocfgil@gmail.com
evbekker@ya.ru Sergei M. Glagolev [Chapter 16.2] Moscow South-West
Aleksander Bielecki [Chapter 12 (Hirudinida)] Depart- High School, Moscow, Russia; email:
ment of Zoology, Faculty of Biology and Biotech- sglagolev@yandex.ru
nology, University of Warmia and Mazury in Olsztyn, Christopher J. Glasby [Chapter 12 (Aphanoneura; Poly-
Olsztyn, Poland; email: alekb@uwm.edu.pl chaeta)] Museum & Art Gallery Northern Territory,
Dirk Brandis [Chapter 16.6 (Decapoda)] Zoological Darwin, NT, Australia; email: chris.glasby@nt.gov.au
Museum, University of Kiel, Kiel, Germany; email: Fredric R. Govedich [Chapter 12 (Hirudinida)] Depart-
brandis@zoolmuseum.uni-kiel.de ment of Biological Sciences, Southern Utah University,
Mikhail E. Daneliya [Chapter 16.6 (Mysida and Stygo- Cedar City, UT, United States; email:
mysida)] Finnish Museum of Natural History, University govedich@suu.edu
of Helsinki, Helsinki, Finland; email: mikhail.daneliya@ Paolo Grilli [Chapter 7] Department of Biomolecular
helsinki.fi Sciences (DISB), University of Urbino “Carlo Bo”,
Wilfrida Decraemer [Chapter 9] Department of Biology, Urbino, Italy; email: paolo.grilli2@tin.it
Ghent University, Ghent, Belgium, Department of Roberto Guidetti [Chapter 15] Department of Life
Taxonomy & Phylogeny, Royal Belgian Institute of Sciences, University of Modena and Reggio Emilia,
Natural Sciences, Brussels, Belgium; email: Modena, Italy; email: roberto.guidetti@unimore.it
wilfrida.decraemer@ugent.be Tom Hansknecht [Chapter 16.6 (Tanaidacea)] Barry Vit-
R. Edward DeWalt [Chapter 16.1] University of Illinois, tor & Associates, Inc., Mobile, AL, United States;
Prairie Research Institute, Illinois Natural History email: bvataxa@bvaenviro.com

xiii
xiv
David J. Horne [Chapter 16.3] School of Geography,
Queen Mary, University of London, London, United
Kingdom; email: d.j.horne@qmul.ac.uk
Thomas Jankowski [Chapter 4] Teningen, Germany;
email: thomas.jankowski@posteo.de
Sebastian Klaus [Chapter 16.6 (Decapoda)] Department of
Ecology & Evolution, Goethe-University e Biologicum,
Frankfurt am Main, Germany; email: klaus@bio.
uni-frankfurt.de
Nikolai M. Korovchinsky [Chapter 16.2] A. N. Severtsov
Institute of Ecology and Evolution, Moscow, Russia;
email: nmkor@yandex.ru
Alexey A. Kotov [Chapter 16.2] A. N. Severtsov Institute
of Ecology and Evolution, Moscow, Russia; email:
alexey-a-kotov@vandex.ru
Dong Ju Lee [Chapter 16.4] Department of Life Science,
Hanyang University, Seoul, Republic of Korea; email:
velocy00@gmail.com
Wonchoel Lee [Chapter 16.4] Department of Life Science,
Hanyang University, Seoul, Republic of Korea; email:
wlee@hanyang.ac.kr
Julian J. Lewis [Chapter 16.6 (Isopoda)] Lewis & Associ-
ates LLC, Borden, IN, United States; email:
lewisbioconsult@aol.com
Lawrence L. Lovell [Chapter 12 (Introduction to the
Phylum)] Research and Collections, Natural History
Museum of Los Angeles County, Los Angeles, CA,
United States; email: lllpolytax@gmail.com
Renata Manconi [Chapter 3] Dipartimento di Medicina
Veterinaria, Università di Sassari, Sassari, Italy; email:
r.manconi@uniss.it
Koen Martens [Chapter 16.3] Royal Belgian Institute of
Natural Sciences, Freshwater Biology, Brussels,
Belgium; email: darwinula@gmail.com
Daniel Martin [Chapter 12 (Aphanoneura; Polychaeta)]
CEAB-CSIC, Blanes, Catalunya, Spain; email:
dani@ceab.csic.es
Patrick Martin [Chapter 12 (Oligochaeta)] Royal Belgian
Institute of Natural Sciences, Operational Directorate
Taxonomy & Phylogeny, Brussels, Belgium; email:
patrick.martin@sciencenaturelles.be
Claude Meisch [Chapter 16.3] Musée national d’histoire
naturelle, Luxembourg; email: claude.meisch@
education.lu
Hiroshi Morino [Chapter 16.6 (Amphipoda)] Department
of Zoology, National Museum of Nature and Science,
Tsukuba, Ibaraki, Japan; email: hiroshi.morino.talitrids
@vc.ibaraki.ac.jp
Contributors to Volume IV
William E. Moser [Chapter 12 (Hirudinida)] Department
of Invertebrate Zoology, National Museum of Natural
History, Smithsonian Institution, Suitland, MD, United
States; email: moserw@si.edu
Takafumi Nakano [Chapter 12 (Hirudinida)] Department
of Science Education, Graduate School of Education, Hir-
oshima University, Higashihiroshima, Japan; Department
of Zoology, Graduate School of Science, Kyoto Univer-
sity, Kyoto, Japan; email: tnakano@hiroshima-u.ac.jp
Tohru Naruse [Chapter 16.6 (Decapoda)] Tropical
Biosphere Research Center, University of the Ryukyus,
Taketomi, Japan; email: naruse@lab.u-ryukyu.ac.jp
Diane R. Nelson [Chapter 15] Department of Biological
Sciences, East Tennessee State University, Johnson
City, TN, United States; email: janddnelson@yahoo.com
Ngan Kee Ng [Chapter 16.6 (Decapoda)] Department of
Biological Sciences, National University of Singapore,
Republic of Singapore; email: ngankee@nus.edu.sg
Carolina Noreña [Chapter 5] Departamento Bio-
diversidad y Biologı́a Evolutiva, Museo Nacional de
Ciencias Naturales (CSIC), Madrid, España; email:
mcnnj92@mncn.csic.es
Sameer Kumar Pati [Chapter 16.6 (Decapoda)] Zoolog-
ical Survey of India, Western Regional Centre, Pune,
India; email: sameerkumarpati@gmail.com
Victor V. Petryashov [Chapter 16.6. (Mysida and Stygio-
mysida)] Zoological Institute of the Russian Academy
of Sciences, Saint Petersburg, Russia; email:
taxalab@taxonomicum.com
Andrey Porfiriev [Chapter 5] Kazan State University,
Kazan, Russia; email: andpor@rambler.ru
Roberto Pronzato [Chapter 3] Dipartimento di Scienze
della Terra, dell’Ambiente e della Vita (DISTAV), Uni-
versità di Genova, Genova, Italy; email:
pronzato@dipteris.unige.it
Lorena Rebecchi [Chapter 15] Department of Life
Sciences, University of Modena and Reggio Emilia,
Modena, Italy; email: lorena.rebecchi@unimore.it
Vincent H. Resh [Chapter 16.1 (Hexapoda)] Department
of Environmental Science, Policy, and Management,
University of California, Berkeley, CA, United States;
email: resh@berkeley.edu
D. Christopher Rogers [Chapters 1, 11, 16, 16.1, 16.2,
16.5, 16.6] Kansas Biological Survey and The Biodi-
versity Institute, University of Kansas, Lawrence, KS,
United States; email: branchiopod@gmail.com
S.S.S. Sarma [Chapter 8] Universidad Nacional Autón-
oma de México Campus Iztacala, México City,
México; email: sssarma@gmail.com
Contributors to Volume IV
Andreas Schmidt-Rhaesa [Chapter 10] Zoological
Museum, University Hamburg, Hamburg, Germany;
email: andreas.schmidt-rhaesa@uni-hamburg.de
Hendrik Segers [Chapter 8] Royal Belgian Institute of
Natural Sciences, Brussels, Belgium; email:
Hendrik.Segers@naturalsciences.be
Artem Y. Sinev [Chapter 16.2] Department of Invertebrate
Zoology, Biological Faculty, M.V. Lomonosov Mos-
cow State University, Leninskie Gory, Moscow, Russia;
email: artem.sinev@gmail.com
Boris Sket [Chapter 16.6 (Amphipoda)] Biology depart-
ment, Biotechnical Faculty, Univerza v Ljubljani,
Slovenia; email: sidorov@biosoil.ru
Nikolai N. Smirnov [Chapter 16.2] A. N. Severtsov Insti-
tute of Ecology and Evolution, Moscow, Russia; email:
smirnov08520@mail.ru
T.W. Snell [Chapter 8] School of Biology, Georgia Insti-
tute of Technology, Atlanta, GA, United States; email:
terry.snell@biology.gatech.edu
Malin Strand [Chapter 6] The Swedish Species Informa-
tion Centre, Swedish University of Agricultural
Sciences, Uppsala, Sweden; email: Malin.Strand@slu.se
Per Sundberg [Chapter 6] Department of Marine
Sciences, University of Gothenburg, Gothenburg,
Sweden; email: P.Sundberg@zool.gu.se
Vadim Takhteev [Chapter 16.6 (Amphipoda)] Department
of Invertebrate Zoology & Hydrobiology, Irkutsk State
University, Russia; email: amphipoda@yandex.ru
James H. Thorp [Chapters 1, 12] Kansas Biological
Survey and Department of Ecology and Evolutionary
Biology, University of Kansas, Lawrence, KS, United
States; email: thorp@ku.edu
Tarmo Timm [Chapter 12 (Oligochaeta)] Institute of
Agricultural and Environmental Sciences, Centre for
Limnology, Estonian University of Life Sciences,
Tartu, Estonia; email: tarmo.timm@emu.ee
xv
Oleg Timoshkin [Chapter 5] Limnological Institute, Ulan-
Batorskaya, Irkutsk, Russia; email: tim@lin.irk.ru
Serge Utevsky [Chapter 12 (Hirudinida)] Department of
Zoology and Animal Ecology, V.N. Karazin Kharkiv
National University, Kharkiv, Ukraine; email:
serge.utevsky@gmail.com
Risto Väinölä [Chapter 16.6 (Mysida and Stygiomysida)]
Finnish Museum of Natural History, University of
Helsinki, Helsinki, Finland; email: risto.vainola@
helsinki.fi
Robert J. Van Syoc [Chapter 16.5 (Sessilia)] California
Academy of Sciences, San Francisco, CA, United
States; email: bvansyoc@calacademy.org
Maxim V. Vinarski [Chapter 11 (Gastropoda)] Saint-
Petersburg State University, Saint-Petersburg, Russia;
email: radix.vinarski@gmail.com
Robert L. Wallace [Chapter 8] Department of Biology,
Ripon College, Ripon, WI, United States; email:
wallacer@ripon.edu
E.J. Walsh [Chapter 8] Department of Biological Science,
University of Texas at El Paso, El Paso, TX, United
States; email: ewalsh@utep.edu
Alan Warren [Chapter 2] Department of Life Sciences,
Natural History Museum, London, United Kingdom;
email: a.warren@nhm.ac.uk
Les Watling [Chapter 16.6 (Cumacea)] Department of
Biology, University of Hawaii at Manoa, Honolulu,
HI, United States; email: watling@hawaii.edu
Timothy S. Wood [Chapters 13-14] Department of Bio-
logical Sciences, Wright State University, Dayton,
OH, United States; email: tim.wood@wright.edu
Darren C.J. Yeo [Chapter 16.6 (Decapoda)] Department
of Biological Sciences, National University of
Singapore, Republic of Singapore; email:
darrenyeo@nus.edu.sg
This page intentionally left blank

Dr. D. Christopher Rogers is a research zoologist at the
University of Kansas with the Kansas Biological Survey
and is affiliated with the Biodiversity Institute. He received
his Ph.D. degree from the University of New England in
Armidale, NSW, Australia. Christopher specializes in
freshwater crustaceans (particularly Branchiopoda and
Malacostraca) and the invertebrate fauna of seasonally
astatic wetlands on a global scale. He has numerous peer-
reviewed publications in crustacean taxonomy and inver-
tebrate ecology, as well as published popular and scientific
field guides and identification manuals to freshwater
invertebrates. Christopher is an Associate Editor for the
Journal of Crustacean Biology and a founding member of
the Southwest Association of Freshwater Invertebrate
Taxonomists. He has been involved in aquatic invertebrate
conservation efforts all over the world.
About the Editors
Dr. James H. Thorp has been a Professor in the Department 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 Professor 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 University, Visiting
Associate Professor at Cornell, and Research Ecologist at the
University of Georgia’s Savannah River Ecology Laboratory.
He received his Baccalaureate from the University of Kansas
(KU) and both Masters and Ph.D. degrees from North Carolina
State. Those degrees focused on zoology, ecology, and marine
biology, with an emphasis on the ecology of freshwater and
marine invertebrates. Dr. Thorp has been on the editorial board
of three freshwater journals and is a former President of the
International Society for River Science. He teaches freshwater,
marine, and invertebrate courses at KU, and his Masters and
Doctoral graduate students work on various aspects of the
ecology of communities through macrosystems in rivers,
reservoirs, and wetlands. Prof. Thorp’s research interests and
background are highly diverse and span the gamut from
organismal biology to community, ecosystem, and macro-
system ecology. He works on both fundamental and applied
research topics using descriptive, experimental, and modeling
approaches in the field and lab. While his research emphasizes
aquatic invertebrates, he also studies fish ecology, especially as
related to food webs. He has published more than one 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 three volumes in the current fourth edition of
Thorp and Covich’s Freshwater Invertebrates.
xvii
This page intentionally left blank

Those readers familiar with the first three editions of our
invertebrate book (Ecology and Classification of North
American Freshwater Invertebrates, edited by J.H. Thorp
and A.P. Covich) will note that the fourth edition has
expanded from a North American focus to worldwide
coverage of inland water invertebrates. We gave our book
series on inland water invertebrates the name Thorp and
Covich’s Freshwater Invertebrates to: (1) associate present
with past editions, unite current volumes, and link to future
editions; (2) establish a connection between the ecological
and general biology coverage in Volume I with the taxo-
nomic keys in the remaining volumes; and (3) give credit to
Professor Alan Covich for his work on the first three
editions. For the sake of brevity, we refer to the current
edition as T&C IV. Whether a fifth edition of T&C will ever
appear is certainly problematic, but who knows? At present
we are considering producing up to 12 volumes in the
fourth edition.
While I am the sole editor of the “series” at this point, Dr.
D. Christopher Rogers has been a major and highly valued
partner in developing ideas for the fourth edition and is thus
far a coeditor on the first four volumes in print (senior editor
on the fourth) as well as on the three volumes in develop-
ment. He is also likely to play a major role in many of the
remaining volumes because of his diverse and global
knowledge of freshwater invertebrates, especially in the area
of taxonomy. As we made significant progress on the first
seven volumes, we began contacting potential coeditors to
develop two more volumes for another zoogeographic
region, and negotiations with those potential editors are now
underway. However, we are still seeking experts in fields of
invertebrate taxonomy for various zoogeographic regions to
serve as highly dependable coeditors, especially those who
both work and live in the zoogeographic regions covered by
the various future volumes.
Preface to the Fourth Edition
Our concept for T&C IV included producing one book
(Vol. I in 2015) with six chapters on general environmental
issues applicable to many invertebrates, followed by 35
chapters devoted to individual taxa at various levels (order
to phylum, or even multiple phyla in the case of the
protozoa). Volume I was designed both as an independent
book on ecology and general biology of the various fresh-
water invertebrate taxa and as a companion volume for
users of the keys in the regional taxonomic volumes,
thereby reducing the amount of information duplicated in
the taxonomic volumes. The perhaps 11 taxonomic
volumes we foresee publishing in this series will contain
both keys for identifying invertebrates in specific zoogeo-
graphic regions and descriptions of detailed anatomical
features needed to employ those keys.
While the vast majority of authors in T&C IeIII were
from the USA or Canada, we attempted in the global
volumes of T&C IV to attract authors from many additional
countries on six continents. Although we largely succeeded
in this goal, we expect the fifth edition of T&Cdif it is ever
publisheddto continue increasing the proportion of
authors from outside North America as our books become
better known internationally.
Our goals for T&C IV are to improve the state of
taxonomic and ecological knowledge of inland water
invertebrates, to help protect our aquatic biodiversity, and
to encourage more students to devote their careers to
working with these fascinating organisms. These goals are
especially important because the verified and probable
losses of species in wetlands, ponds, lakes, creeks, and
rivers around the globe exceed those in most terrestrial
habitats.
James H. Thorp
xix
Another random document with
no related content on Scribd:
 Fig. 254. Osmundites Kolbei. (Leaf-scars.)

Osmundites Kolbei Seward, figs. 253–255.

This species was founded on a specimen obtained by Mr Kolbe
from the Uitenhage series of Cape Colony[788]. The fossil flora and
fauna of this series point to its correlation with the Wealden or
Neocomian strata of Europe[789]. The type-specimen consists of
several pieces of a stem (fig. 253) which reached a length of about
90 cm. On the weathered surface the remains of petiole-bases are
clearly seen and on the reverse side of the smaller piece shown in
the figure numerous sinuous roots are present in association with the
leaf-stalks. The depression c in the larger specimen may mark the
position of a branch: at a fig. 253 (enlarged in fig. 254, a) the
vascular strand of a petiole is exposed as a broad U-shaped band
and at b (fig. 254, b) the form of the petiole-bases is clearly
shown[790]. With the stem were found imperfectly preserved
impressions of fronds referred to Cladophlebis denticulata, a
common type of leaf which was found also in association with the
slightly older New Zealand stem, Osmundites Dunlopi.
 Fig. 255. Osmundites Kolbei Sew. Transverse section, from a photograph
supplied by Dr Kidston and Mr Gwynne-Vaughan. (2½ nat. size.)
An examination of the internal structure of the South African stem
by Dr Kidston and Mr Gwynne-Vaughan has revealed many
interesting features, which will be fully described in Part IV. of their
Monograph on fossil Osmundaceous stems. I am greatly indebted to
these authors for allowing me to publish the following note
contributed by Dr Kidston:—
“The section of Osmundites Kolbei Seward, shown in fig. 255,
presents the usual appearance of an Osmundaceous stock. The
parts contained in this section are the stele, inner and outer cortex
and a portion of the surrounding mantle of concrescent leaf-bases.
The whole specimen has suffered much from pressure, but if
restored to its original form the xylem ring must have been about 19
mm. in diameter. The number of xylem strands is about fifty-six and
several of them are more or less joined as in the modern genus
Todea. The tracheae are of the typical Osmundaceous type, that is
to say, the pits are actual perforations and several series of them
occur on each wall of the larger tracheae.
“The most interesting structural characteristic of Osmundites
Kolbei is not well seen in the figure owing to the compression of the
xylem ring. This consists in the occurrence of tracheae in the pith. In
fact, we have here a mixed pith, composed of parenchyma and true
tracheae, a condition which connects the Osmundaceae with a
parenchymatous medulla with those possessing a solid xylem stele
like Zalesskya and Thamnopteris and so completes the series of
transitions extending from the older and solid-steled forms to the
modern medullated members of the Osmundaceae.”

Osmundites skidegatensis, Penhallow.

This lower Cretaceous Canadian species, first described by
Penhallow[791] and more recently by Kidston and Gwynne-
Vaughan[792], is remarkable for the large size of the stem, the stele
alone having a diameter of 2·4 cm. Penhallow figures a fragment of
a leaf bearing a superficial resemblance to that of Osmunda
Claytoniana, which may be the foliage borne by Osmundites
skidegatensis. The xylem cylinder is broken by the exit of leaf-traces
into 50 or more strands varying in size and shape, and it is
noteworthy that the phloem is also interrupted as each leaf-trace is
given off. In recent species the xylem cylinder is almost always
interrupted, but the phloem retains its continuity. In the Canadian
fossil an internal band of phloem occurs between the xylem and the
pith, and this joins the external phloem at each leaf-gap. This internal
phloem finds an interesting parallel in certain recent species[793], but
in these the internal and external phloem do not meet at the foliar
gaps as they do in the extinct type. In Osmunda cinnamomea the
internal phloem occurs only at the regions of branching of the stem
stele; in the fossil it is always present.
It is clear that Osmundites skidegatensis represents the most
complex type of stem so far recognised in the Osmundaceae; it
illustrates a stage in elaboration of the primitive protostele in
advance of that reached by any existing species.
• • • • •
The primitive Osmundaceous stele was composed of solid xylem
surrounded by phloem (Thamnopteris and Zalesskya); at a later
stage the xylem cylinder lost its inner zone of wide and short
tracheae and assumed the form seen in Osmundites Kolbei, in which
the centre of the stele consists of parenchyma with some tracheae.
Another type is represented by O. Dowkeri in which the pith is
composed wholly of parenchyma and the xylem ring is continuous.
From this type, by expansion of the xylem ring and by the formation
of overlapping leaf-gaps, the form represented by Osmunda regalis
was reached. Osmunda cinnamomea, with internal phloem in the
regions of stelar branching, probably represents a further stage, as
Kidston and Gwynne-Vaughan believe, in increasing complexity due
to the introduction of phloem from without through gaps produced by
the branching of the stele. In Osmundites skidegatensis the leaf-
gaps became wider and the external phloem projected deeper into
the stele until a continuous internal phloem zone was produced. This
most elaborate type proved less successful than the simpler forms
which still survive.

Osmundites Sturii.
Impressions of fertile pinnae with narrow linear segments bearing
exannulate sporangia described by Raciborski from Lower Jurassic
rocks in Poland as Osmunda Sturii[794] may with some hesitation be
included in the list of Mesozoic Osmundaceae.

Osmundites Dowkeri.
Under this name Carruthers[795] described a petrified stem from
Lower Eocene beds at Herne Bay, which in the structure of the stele
agrees closely with the Jurassic species O. Gibbiana and conforms
to the normal Osmundaceous type. It is possible, as Gardner and
Ettingshausen[796] suggested, that the foliage of this species may be
represented by some sterile Osmunda-like fragments recorded from
the Middle Bagshot beds of Bovey Tracey and Bournemouth as
Osmunda lignitum.

Todites.
This generic name[797] has been applied to fossil ferns exhibiting in
the structure of the sporangia and in the general habit of the fertile
fronds a close resemblance to the recent species Todea barbara (fig.
221, D, p. 286).

Todites Williamsoni (Brongniart) figs. 256, B, C, G.

1828. Pecopteris Williamsonis, Brongniart, Prodrome, p. 57; Hist.
vég. foss., p. 324, Pl. cx. figs. 1 and 2.
— P. whitbiensis, Brongniart, Hist. vég. foss. p. 321, Pl. cix. figs.
2–4.
— P. tenuis, ibid. p. 322, Pl. cx. figs. 3, 4.
1829. Pecopteris recentior, Phillips, Geol. Yorks. p. 148, Pl. viii. fig.
15.
— P. curtata, ibid. Pl. viii. fig. 12.
1833. Neuropteris recentior, Lindley and Hutton, Foss. Flora, Vol. i.
Pl. lxviii.
— Pecopteris dentata, ibid. Vol. iii., Pl. clxix.
1836. Acrostichites Williamsonis, Goeppert, foss. Farn. p. 285.
1841. Neuropteris Goeppertiana, Muenster, in Goeppert,
Gattungen foss. Pflanz. Lief. 5 and 6, p. 104, Pls. viii.–x.
1856. Pecopteris Huttoniana, Zigno, Flor. foss. Oolit. Vol. i. p. 133.
1867. Acrostichites Goeppertianus, Schenk, Foss. Flor. Grenzsch.
p. 44, Pl. v. fig. 5, Pl. vii. fig. 2.
1883. A. linnaeaefolius, Fontaine, Older Mesoz. Flora Virginia, p.
25, Pls. vi.–ix.
— A. rhombifolius, ibid. Pls. viii. xi.–xiv.
1885. Todea Williamsonis, Schenk, Palaeont. Vol. xxxi. p. 168, Pl.
iii. fig. 3.
1889. Cladophlebis virginiensis, Fontaine, Potomac Flora, p. 70, Pl.
iii. figs. 3–8; Pl. iv. figs. 1, 4.
 Fig. 256.
A. Cladophlebis denticulata.
B, B′. Todites Williamsoni (fertile).
C. T. Williamsoni (sterile pinna).
D. Discopteris Rallii.
E, E′. Kidstonia heracleensis.
F. Todeopsis primaeva.
G. Todites Williamsoni (sporangium).
[B, C, from specimens (13491; 39234) in the British Museum (B, very
slightly reduced; C, ½ nat. size); D, E, after Zeiller; F, after Renault; G,
after Raciborski.]
It is hopeless to attempt to arrive at satisfactory conclusions in
regard to the applicability of the name Todites Williamsoni to the
numerous fronds from Jurassic and Rhaetic rocks, agreeing more or
less closely with Brongniart’s type-specimen. Specimens from the
Rhaetic may not be specifically identical with those from the
Jurassic; the main point is that, whether actually identical or not, both
sets of fossils clearly represent the same general type of
Osmundaceous fern[798] and may for present purposes be included
under the same designation. The above synonymy, though by no
means complete[799], serves to illustrate the confusion which has
existed in regard to this widely spread type of Mesozoic fern.
Todites Williamsoni may be briefly described as follows:—
Frond bipinnate; long linear pinnae (20–30 cm.) of uniform breadth arise at
an acute angle, or in the lower part of a frond, almost at right angles, from a
stout rachis. Closely set pinnules attached by a broad base; slightly falcate, the
side towards the rachis strongly convex and the outer margin straight or
concave and bulged outwards towards the base of each segment, margin
usually entire, or it may be slightly lobed. Fertile pinnules similar to the sterile;
sporangia of the Osmundaceous type and often scattered over the whole lower
surface of the lamina (fig. 256, B, B′, G). Venation of the Cladophlebis type (cf.
fig. 256, A).

It is not always easy to distinguish Todites Williamsoni from

Cladophlebis denticulata, another common Jurassic fern, but in the
latter the pinnules are usually longer and relatively narrower and the
rachis is more slender (cf. fig. 256, B and 257). Schenk[800] and
Raciborski[801] have shown that the sporangia of Todites conform in
the absence of a true annulus to those of Todea (fig. 256, G) and
Osmunda. Nathorst[802] has recently figured a group of spores of
Todites Williamsoni in illustration of the use of the treatment of
carbonised impressions with nitric acid and potassium chlorate. This
species, though widely distributed in Jurassic rocks, is hardly
distinguishable from the German Rhaetic fronds figured by Schenk
from Bayreuth as Acrostichites Goeppertianus[803], or from other
fossils referred to an unnecessarily large number of species by
Fontaine[804] from Upper Triassic rocks of Virginia[805].
It would seem from the paucity of later records of Osmundaceae
that the family reached its zenith in the Jurassic era. When we pass
to the later Tertiary and more recent deposits evidence is afforded in
regard to the geographical range of Osmunda regalis. It has been
shown to occur in the Pliocene forest-bed of Norfolk[806] as well as in
Palaeolithic and Neolithic deposits[807].

Fig. 257. Cladophlebis denticulata. (From a specimen in the British

Museum from the Inferior Oolite rocks of Yorkshire. Slightly
reduced.)
A fertile frond from the Molteno (Rhaetic) beds of South Africa
referred to Cladophlebis (Todites) Roesserti (Presl)[808] represents in
all probability an Osmundaceous fern closely allied to Todites
Williamsoni. The same species is described by Zeiller[809] from
Rhaetic rocks of Tonkin and very similar types are figured by
Leuthardt[810] from Upper Triassic rocks of Basel as Pecopteris
Rutimeyeri Heer, and by Fontaine[811] from rocks of the same age in
Virginia.

Cladophlebis.
The generic name Cladophlebis was instituted by Brongniart for
Mesozoic fern fronds characterised by ultimate segments of linear or
more or less falcate form attached to the pinnae by the whole of the
base, as in the Palaeozoic genus Pecopteris, possessing a midrib
strongly marked at the base and dividing towards the distal end of
the lamina into finer branches and giving off secondary forked and
arched veins at an acute angle. The term is generally restricted to
Mesozoic fern fronds which, on account of the absence or
imperfection of fertile pinnae, cannot be safely assigned to a
particular family. In the case of the species described below, the
evidence in regard to systematic position, though not conclusive, is
sufficiently strong to justify its inclusion in the Osmundaceae.

Cladophlebis denticulata Brongniart. Figs. 256, A; 257, 258.

1828. Pecopteris denticulata[812], Brongniart, Prodrome, p. 57; Hist.
vég. foss. p. 301, Pl. xcviii. figs. 1, 2.
— P. Phillipsii, Brongniart, Hist. p. 304, Pl. cix. fig. 1.
This species is often confused[813] with Todites Williamsoni. The
name Pecopteris whitbiensis has been used by different writers for
Jurassic fronds which are undoubtedly specifically distinct:
specimens so named by Brongniart should be referred to Todites
Williamsoni, while P. whitbiensis of Lindley and Hutton[814] is
Brongniart’s Cladophlebis denticulata. It is impossible to determine
with accuracy the numerous examples described as Pecopteris
whitbiensis, Asplenium whitbiense, Cladophlebis Albertsii (a
Wealden species[815]), Asplenium, or Cladophlebis, nebbense[816],
etc., from Jurassic and Rhaetic strata. The Cladophlebis denticulata
form of frond is one of the commonest in recent ferns; it is
represented by such species as Onoclea Struthopteris, Pteris arguta,
Sadleria sp., Gleichenia dubia, Alsophila lunulata, Cyathea dealbata,
and species of Polypodium. It is, therefore, not surprising to find
records of this Mesozoic species from many localities and horizons.
All that we can do is to point out what appear to be the most
probable cases of identity among the numerous examples of fronds
of this type from Mesozoic rocks, particularly Rhaetic and Jurassic,
in different parts of the world. The name Cladophlebis denticulata
may be employed in a comprehensive sense for fronds showing the
following characters:—
Leaf large, bipinnate, with long spreading pinnae borne on a comparatively
slender rachis. Pinnules, in nearly all cases, sterile, reaching a length of 3–
4cm., acutely pointed, finely denticulate or entire, attached by the whole of the
base (fig. 257). In the apical region the pinnules become shorter and broader.
Venation of the Cladophlebis type (fig. 256, A). Fertile pinnules rather straighter
than the sterile, characterised by linear sori parallel to the lateral veins (fig.
258).

In endeavouring to distinguish specifically between fronds showing

a general agreement in habit with C. denticulata, special attention
should be paid to venation characters, the shape of the pinnules, the
relation of the two edges of the lamina to one another, and to the
amount of curvature of the whole pinnule. Unless the material is
abundant, it is often impossible to distinguish between characters of
specific value and others which are the expression of differences in
age or of position on a large frond, to say nothing of the well-known
variability which is amply illustrated by recent ferns. It is remarkable
that very few specimens are known which throw any light on the
nature of the fertile pinnae. Fig. 258 represents an impression from
the Inferior Oolite rocks of the Yorkshire coast in which the exposed
upper surface of the pinnules shows a series of parallel ridges
following the course of the lateral veins and no doubt formed by
oblong sori on the lower surface. There can be little doubt that the
specimen figured by Lindley and Hutton and by others as Pecopteris
undans[817] is, as Nathorst suggests, a portion of a fertile frond of C.
denticulata. A fertile specimen of a frond resembling in habit C.
denticulata, which Fontaine has described from the Jurassic rocks of
Oregon as Danaeopsis Storrsii[818], exhibits, as that author points out,
a superficial resemblance to the specimen named by Lindley and
Hutton Pecopteris undans. There is, however, no adequate reason
for referring the American fragment to the Marattiaceae. In the
absence of sporangia we cannot speak confidently as to the
systematic position of this common type; but there are fairly good
grounds for the assertion that some at least of the fronds described
under this name are those of Osmundaceae. The English specimen
shown in fig. 258 is very similar to some Indian fossils figured by
Feistmantel as Asplenites macrocarpus[819], which are probably
identical with Pecopteris australis Morris[820], a fern that is
indistinguishable from Cladophlebis denticulata. Renault[821] figured a
fertile specimen of the Australian fossil as Todea australis, which
agrees very closely with that shown in fig. 258, and the sporangia
figured by the French author are of the Osmundaceous type.
Another example of a fertile specimen is afforded by a Rhaetic fern
from Franconia, Asplenites ottonis, which is probably identical with
Alethopteris Roesserti Presl [= Cladophlebis (Todites) Roesserti], a
plant closely resembling Cladophlebis denticulata. Another argument
in favour of including C. denticulata in the Osmundaceae is supplied
by the association of pinnae of this type with the petrified stem of
Osmundites Dunlopi recorded by Kidston and Gwynne-Vaughan.

Fig. 258. Fertile pinnae of Cladophlebis denticulata. (From a Yorkshire

specimen in the Sedgwick Museum, Cambridge.)
 Schizaeaceae.
Evidence bearing on the existence of this family in Carboniferous
floras is by no means decisive. The generic name Aneimites
proposed by Dawson[822] for some Devonian Canadian plants
resembling species of the recent genus Aneimia, and adopted by
White[823] for a species from the Pottsville beds of Virginia, is
misleading. The Canadian plants give no indication of the nature of
the reproductive organs, and the fronds described by White are, as
he shows, those of a Pteridosperm and bore seeds.
An examination of the suspiciously diagrammatic drawings
published by Corda[824] of the small fertile pinnules of a
Carboniferous fern from Bohemia, which he named Senftenbergia
elegans, leads us to conclude that the sporangia are almost certainly
those of a Schizaeaceous species. The small linear pinnules bear
two rows of sessile sporangia, singly as in recent Schizaeaceae and
not in sori, characterised by 4–5 rows of regular annular cells (fig.
270, A) surrounding the apex. It has already been pointed out that
the apical annulus of recent Schizaeaceae, though normally one row
deep, may consist in part at least of two rows. Zeiller[825] examined
specimens of Corda’s species and decided in favour of a
Schizaeaceous affinity; he describes the sporangia as 0·85–0·95
mm. in length, with 3 to 5 and occasionally only two rows of cells in
the apical annulus. Zeiller’s figures (fig. 270, A) confirm the
impression that Corda’s drawings are more beautiful than accurate.
Stur[826], on the other hand, who first pointed out that the type-
specimens of Senftenbergia came from the Radnitz beds of Bohemia
and not from the Coal-Measures, convinced himself that the
sporangia have no true annulus (fig. 270, E). He describes them as
characterised by a comparatively strong wall and by the presence of
a band of narrow vertical cells marking the line of dehiscence,
features which lead him to assign the plant to the Marattiales, a
group which seems to have exercised a dominating influence over
his judgment. In a later publication Zeiller[827] replies to Stur’s
criticism but adheres to his original opinion. Solms-Laubach[828],
while expressing himself in favour of Marattiaceous affinity,
recognises that Zeiller’s arguments cannot be set aside.
 The question must remain open until further evidence is
forthcoming; but it would seem that this Carboniferous type, not as
yet recognised in Britain, possessed sporangia having a distinct
resemblance to those of the Schizaeaceae, though this similarity
does not amount to proof of the existence of the family in the
Palaeozoic era.
Palaeozoic floras may be described as rich in generalised types,
types foreshadowing lines of evolution, which in the course of ages
led to a sorting and a redistribution of characters. It may be that
Senftenbergia is one of these generalised types.
• • • • •
It is not until we ascend the geological series as far as the older
Jurassic rocks that we meet with a type which can with confidence
be classed with the Schizaeaceae, as least so far as sporangial
characters are concerned. The species Klukia exilis is selected as
the best known and most widely-spread representative of Jurassic
Schizaeaceae.

Klukia exilis (Phillips)[829]. Fig. 259.

The generic name Klukia was proposed by Raciborski[830] for a
species originally described by Phillips[831] from the Inferior Oolite of
the Yorkshire coast as Pecopteris exilis. Bunbury’s[832] discovery
(supplemented by additional evidence obtained by Raciborski) of
well-preserved sporangia justified the substitution of a distinctive
designation for the provisional term Pecopteris.
 Fig. 259. Klukia exilis (Phillips). (Figs. 1–3, × 40; fig. 4, × 3; fig. 5, nat.
size.)
The species may be defined as follows:—
Frond tripinnate, of the Cladophlebis type; pinnae linear, lanceolate, attached
to the rachis at a wide angle. Ultimate segments short and linear, entire or, in
the lower part of a frond, crenulate, 5 mm. long or occasionally longer.
Sporangia 0·5 mm. in length, borne singly on the lower surface of the lamina in
a row on each side of the midrib.

A re-examination[833] of the specimen described by Bunbury

confirmed his account of the structure of the sporangia. The pinna
shown in fig. 259 is characterised by unusually small fertile pinnules
some of which bear 10 sporangia in two rows; the annulus includes
about 14 cells. Fertile specimens of this and similar forms are figured
by Raciborski[834] from Jurassic rocks of Poland, and good examples
of the English species may be seen in the Leckenby collection,
Cambridge, in the British Museum, the museums of Manchester,
Scarborough, and other places.
It is possible that specimens referred to K. exilis by Yokoyama[835]
from Wealden strata in Japan may afford evidence of the persistence
of the species beyond the Jurassic era, but in view of the close
resemblance of the sterile fronds described from Wealden strata as
Cladophlebis Brownii[2] and C. Dunkeri[836] to those of Klukia exilis,
identity can be established only by an examination of fertile
specimens. A Jurassic fern recently described by Yabe[837] from
Korea as Cladophlebis koraiensis may be identical with K. exilis and
there is little doubt as to the existence of the species in Jurassic
Caucasian strata[838].

Fig. 260. Ruffordia Goepperti. (A, C, sterile; B, fertile; slightly reduced.

Specimens from the Wealden of Sussex; British Museum; V.
2333, V. 2160, V. 2166.)

Ruffordia Goepperti (Dunk.). Fig. 260.

This Wealden fern[839] has been doubtfully assigned to the
Schizaeaceae on the ground of the resemblance of the sterile fronds
to those of some species of Aneimia, and because of the difference
between the sterile and fertile pinnae (Fig. 260). Ruffordia cannot be
regarded as a well authenticated member of the Schizaeaceae.

Fig. 261.
A, A′. Chrysodium lanzaeanum.
B, B′. Lygodium Kaulfussi.
C. Marattia Hookeri.
(After Gardner and Ettingshausen; A, B, ¾ nat. size.)

Lygodium Kaulfussi, Heer. Fig. 261, B, B′.

Fragments of forked pinnules, agreeing very closely in venation
and general appearance with recent species of Lygodium, have
been identified by Gardner and Ettingshausen[840] from English
Eocene beds and by Knowlton from the Miocene beds of the
Yellowstone Park[841] as Lygodium Kaulfussi Heer (fig. 261, B).
Despite the absence of sporangia it is probable that these fragments
are correctly referred to the Schizaeaceae. The sterile and fertile
specimens figured by Heer[842] from Tertiary beds of Switzerland
agree very closely with recent examples of Lygodium. Similar though
perhaps less convincing evidence of the existence of this family in
Europe is furnished by Saporta[843], who described two Eocene
species from France.

Gleicheniaceae.
The application by Goeppert[844] and other earlier writers of the
generic name Gleichenites to examples of Palaeozoic ferns was not
justified by any satisfactory evidence. One of Goeppert’s species,
Gleichenites neuropteroides, is identical with Neuropteris
heterophylla[845], a plant now included in the Pteridosperms.
The resemblance of sporangia and sori, whether preserved as
carbonised impressions or as petrified material, from Carboniferous
rocks, to those of recent species of Gleicheniaceae is in many cases
at least the result of misinterpretation of deceptive appearances.
Williamson[846] drew attention to the Gleichenia-like structure of some
sections of sporangia from the English Coal-Measures, but he did
not realise the ease with which sections of Marattiaceous sporangia
in different planes may be mistaken for those of annulate
(leptosporangiate) sporangia. In the regular dichotomous habit of
Carboniferous fronds described as species of Diplothmema (Stur)
and Mariopteris (Zeiller)[847] we have a close correspondence with
the leaves of Gleichenia, but the common occurrence of
dichotomous branching among ferns is sufficient reason for
regarding this feature as an untrustworthy criterion of relationship. It
is, however, interesting to find that in addition to the existence of
some Upper Carboniferous ferns with sori like those of recent
Gleichenias, the type of stelar anatomy illustrated by Gleichenia
dicarpa (fig. 237, C, p. 310) and other species is characteristic of the
primary structure of the stem of the Pteridosperm Heterangium. We
find in Carboniferous types undoubted indications of anatomical and
other features which in succeeding ages became the marks of
Gleicheniaceae.
Some Carboniferous fronds with short and small pinnules of the
Pecopteris type, bearing sori composed of a small number of
sporangia, have been assigned by Grand’Eury and other authors to
the Gleicheniaceae; the same form of sorus is met with also on
fronds with Sphenopteroid segments. The former is illustrated by
Oligocarpia Gutbieri[848] and the latter by O. Brongniarti described by
Stur and by Zeiller[849]. Zeiller has described the circular sori of
Oligocarpia (fig. 270, B) as consisting of three to ten pyriform
sporangia borne at the ends of lateral veins and possessing a
complete transverse annulus, but Stur[850] believes that the annulus-
like appearance is due to the manner of preservation of exannulate
sporangia. In this opinion Stur is supported by Solms-Laubach[851]
and by Schenk[852]. Despite an agreement between Oligocarpia and
Gleichenia, as regards the form of the sori and the number of
sporangia, it is not certain that the existence of a typical
Gleicheniaceous annulus has been proved to occur in any
Palaeozoic sporangia[853].
From Upper Triassic beds of Virginia, Fontaine has figured several
fronds for which he instituted the genus Mertensides[854]. The habit,
as he points out, is not dichotomous, but the sori are circular and are
said to be composed in some species of four to six sporangia. No
satisfactory evidence is brought forward in support of the use of a
designation implying a close relationship with recent Gleichenias
(sect. Mertensia). One of the species described by Fontaine was
originally named by Bunbury Pecopteris bullatus[855], the imperfect
type-specimen of which is now in the Museum of the Cambridge
Botany School. In the form of the frond, the thick rachis, and in the
pinnules this Triassic species resembles Todites Williamsoni, but the
resemblance does not extend to the sori. Two of Fontaine’s species
are recorded by Stur from Austria[856], but he places them in the
genus Oligocarpia and includes them in the Marattiaceae.
Leuthardt[857] figures what appears to be a Gleicheniaceous fern
from the Upper Triassic beds of Basel as Gleichenites gracilis (Heer)
showing sori composed of five sporangia (fig. 265, C) with a
horizontal annulus. A Rhaetic species Gleichenites microphyllus
Schenk[858] from Franconia agrees in the form of its small rounded
pinnules with Gleichenia, but no sporangia have so far been found.
An impression of a frond from Jurassic rocks of northern Italy
figured by Zigno as Gleichenites elegans[859] closely resembles in
habit recent species of Gleichenia; though no sporangia have been
found, the habit of the frond gives probability to Zigno’s
determination.
A Jurassic species from Poland, Gleichenites Rostafinskii, referred
by Raciborski[860] to Gleichenia, exhibits a close agreement in habit
and in the form of the soral impressions to some recent species of
Gleichenia.
As we pass upwards to Wealden and more recent rocks it
becomes clear that the Gleicheniaceae were prominent members of
late Mesozoic floras in north Europe and reached as far north as
Disco Island. In English Wealden beds portions of sterile fronds have
been found which were assigned to a new genus Leckenbya[861], but
it is probable that these specimens would be more correctly referred
to Gleichenites. Similarly fragments of Gleichenia-like pinnae with
very small rounded pinnules occur in the Wealden rocks of
Bernissart, Belgium[862], in north Germany[863], and elsewhere.
Conclusive evidence has been obtained by Prof. Bommer of the
existence of Gleichenites in Wealden beds near Brussels, where
many plant remains have been found in a wonderful state of
preservation. The specimens, which I had an opportunity of seeing
some years ago, might easily be mistaken for rather old and brown
pieces of recent plants. Some of the Belgian fragments, of which
Prof. Bommer has kindly sent me drawings and photographs, are
characterised by an arrangement of vascular tissue identical with
that in the petioles and rhizomes of some protostelic Gleichenias.
The stele of one of the Belgian rhizomes appears to be identical with
that of Gleichenia dicarpa (fig. 237, C. p. 310).