Animal Diversity

Diana R. Kershaw Formerly lecturer in anatomy, St Mary's Hospital Medical School

With illustrations by Brian Price Thomas

CHAPMAN & HALL London· Glasgow· New York· Tokyo· Melbourne· Madras Published by Cbapman & Hall, 2-6 Boundary Row, London SEt 8HN

Chapman & Hall, 2-6 Boundary Row, London SEI 8HN, UK Blackie Academic & Professional, Wester Cleddens Road, Bishopbriggs, Glasgow G64 2NZ, UK Chapman & Hall, 29 West 35th Street, New York NY10001, USA Chapman & Hall Japan, Thomson Publishing Japan, Hirakawacho Nemoto Building, 6F, 1-7-11 Hirakawa-cho, Tokyo 102, Japan Chapman & Hall Australia, Thomas Nelson Australia, 102 Dodds Street, South Melbourne, Victoria 3205, Australia Chapman & Hall India, R. Seshadri, 32 Second Main Road, CIT East, Madras 600 035, India

First published in 1983 by University Tutorial Press Ltd This edition 1988 Reprinted 1991, 1992 © 1983 D. R. Kershaw

ISBN-13: 978-0-412-53200-9 e-ISBN-13: 978-94-011-6035-3 DOl: 10.1007/978-94-011-6035-3

Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permission in writing of the publishers, or in the case of reprographic reproduction only in accordance with the terms ofthe licences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page. The publisher makes no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability for any errors or omissions that may be made.

A catalogue record for this book is available from the British Library To my parents Preface

This book has been written with two main purposes in mind, page. At the same time animals show immense variation the first being to give a general review of the entire animal and none is truly typical. Some idea of the immense variety kingdom, and the second to give more detailed functional of animals is given in the diversity sections, with a synopsis accounts of the anatomy of a representative of each major of the classification of each major phylum. animal group. It is intended to be used by those who are Zoology has a language of its own, which appears highly interested in animals and does not start with the assumption complicated but in most cases can, in fact, be derived simply of any great zoological knowledge. It is hoped that it will from either Latin or Greek. Translations and derivations prove particularly helpful to those studying biology or have been given of a selection of zoological terms; these zoology at 'A' level, or in the early stages of a university should be regarded as examples. The interested zoologist course. may find the use of a Greek and Latin dictionary rewarding. Most modern zoological anatomical and physiological Finally, zoology is a subject with a long history stretching research concentrates on 'bits' of animals. This text is back to the ancient world. In many modern texts the designed to help the reader to see the animal as a complete important contribution of earlier workers is forgotten in living being within its environment. For this reason both favour of concentration on more recent achievement. It is detailed description and illustrations are restricted to a few hoped that sufficient historical background has been given representative animals and where possible the structure of to remind the reader of the great value of earlier work. an animal is illustrated by a series of drawings on a single Diana R. Kershaw

The use of this book

Animal Diversity has been planned to be read in three ways, of the variation within each phylum. These sections are apart from as a unit, with the intention of providing three intended to be used together to give a more overall look at text books in one. the animal kingdom. Thirdly, major functions and organ Firstly, each major phylum includes a detailed description systems are discussed within each section under the same of one representative. These have been chosen with regard headings and in the same order. Used together, these to their availability and use in school and university syllabi provide a comparative account of the functional anatomy of and can be used together or separately to give a full account the animal kingdom. of a limited number of animals. Secondly, a review is given Acknowledgements

In the preparation of this book I have benefited from Carthy, G. Causey, G. Chapman, M. Chinery, A. M. suggestions and ideas from many fellow zoologists, Clark, R. B. Clarke, J. A. Clegg, E. H. Colbert, J. L. colleagues, and students. Among my colleagues special Corliss, R. P. Dales, C. Darwin, B. Dawes, R. Denison, E. mention must be made of Drs A. D. Hoyes and R. A. J. Denton, T. Dobzhansky, F. H. Edgeworth, R. Fange, D. Travers, also Professor A. d'A Bellairs. My students at Fawcett, W. Fisher, E. A. Fraser, W. H. Freeman, V. Queen Mary College (University of London) helped me to Fretter, K. von Frisch, W. G. Fry, G. Fryer, C. Gans, W. create the undergraduate courses on which the book was Garstang, R. Gibson, T. Gilson, E. S. Goodrich, A. based. Some read parts of the book and convinced me it was Graham, P. P. Grasse, H. Gray, J. Gray, J. Green, P. H. worth continuing to write it; in this context I am particularly Greenwood, W. K. Gregory, A. J. Grove, J. Hadzi, H. J. grateful to N. M. A. Horn, with J. R. Clague and R. A. Hansen, E. D. Hanson, R. H. Harrison, B. Hatschek, J. W. Matthews. Hedgepeth, W. N. Hess, G. Huff, L. H. Hyman, A. D. My husband, Dr P. J. Edwards, read the entire book in Imms, H. Isseroff, A. V. Ivanov, J. B. Jennings, C. John, manuscript and ensured that the English was M. Jollie, R. P. S. Jefferies, D. Kennedy, G. A. Kerkut, M. comprehensible, also adding some ecological information. S. Laverack, W. E. LeGros Clark, C. H. Lewis, J. A. Mrs L. A. Wheatley turned my scrawled handwriting into a Mcleod, K. F. Liem, H. W. Lissman, E. E. Lund, K. H. tidy typescript. Brian Price-Thomas drew the illustrations Mann, S. M. Manton, A. J. Marshall, N. B. Marshall, L. H. and went to great pains to find adequate source material to Matthews, E. Mayr, P. A. Meglitsch, E. Meyer, R. S. this end, and Mrs A. L. Price Thomas and the Zoology Miles, H. M. Miller, J. E. Morton, J. A. Moy-Thomas, O. Department of Westfield College provided innumerable Nelsen, T. C. Nelson, G. Newell, P. F. Newell, D. Nichols, specimens and references. G. K. Noble, R. T. Orr, R. Owen, T. S. Parsons, W. P. I should like to thank the staff of University Tutorial Pycraft, W. J. Rees, F. S. Russell, W. D. Russell-Hunter, Press, particularly C. J. Baker, who encouraged me to write O. W. Richards, A. S. Romer, K. Schmidt Nielsen, A. the book, and Anne Hollifield and David Blogg for endless Sedgewick, G. C. Simpson, M. A. Sleigh, J. E. Smith, J. C. help in its editorial and artwork aspects. Smyth, R. E. Snodgrass, W. Stephenson, M. F. Sutton, It is impossible in a book of this type to credit every piece D'A W. Thompson, C. A. Villee, W. F. Walker, R. of information to its original source; indeed in many cases Warwick, T. H. Waterman, D. M. S. Watson, J. E. Webb, the original source has been lost. I am deeply indebted to P. S. Welch, T. S. Westoll, P. J. Whitehead, V. B. many zoologists, without whom the book could not have Wigglesworth, P. H. Williams, D. M. Wilson, H. V. been written, and particularly to: A. J. Alexander, R. McN Wilson, C. M. Yonge, J. Z. Young. Alexander, D. AIkins, R. D. Allen, E. P. Allis, D. T. I should like to thank Faber for permission to quote from Anderson, S. B. Barber, E. J. W. Barrington, R. D. 'Archy and Mehitabel' by Don Marquis, Methuen for Barnes, G. R. de Beer, A. d'A Bellairs, N. J. Berrill, Q. permission to quote from 'Now we are six' by A. A. Milne, Berry, A. Bidder, W. Bloom, Q. Bone, B. Bracegirdle, A. and Unwin for permission to quote 'The Fly' and 'The Brodal, T. H. Bullock, R. M. Cable, A. C. Campbell, J. D. Termite' by Ogden Nash. Contents

Introduction 1 Order Monogenea: Polystomum 66 Order Digenea: Fasciola 67 The characteristics of living things 1 Class Cestoda 72 The differences between animals and plants 1 Subclass Eucestoda: Taenia 72 The cell 2 Synopsis of phylum Platyhelminthes 76 Cell division 5 Phylum Nemertinea 76 Animal classification 8 Phylum Mesozoa 81 The origins and interrelationships of animals 9 Phylum Gnathostomulida 82 Animals and their environment 11 Animal body cavities 84 Phylum Protozoa 14 Amoeba: a protozoan of simple structure 14 The pseudocoelomate phyla 85 A general consideration of protozoan structure 17 Euglena: a 'plant-like' protozoan 18 Super phylum Aschelminthes 85 Paramecium: a complex protozoan 20 Phylum Nematoda: Ascaris 85 Monocystis and Plasmodium: parasitic Protozoa 25 Minor pseudocoelomate phyla 90 The classification of the Protozoa 28 Phylum Rotifera 90 Protozoan diversity 28 Phylum Gastrotricha 92 Synopsis of the Protozoa 33 Phylum Kinorhyncha 93 Phylum Nematomorpha 94 Phylum Acanthocephala 95 Phylum Porifera 34 Phylum Annelida 97 The Radiata 38 Introduction to the coelomate animals and metamerism 97 Phylum Cnidaria 38 Class Polychaeta: Nereis 98 Hydra: an example of a polyp 38 Arenicola marina 104 Obelia: a representative hydrozoan 42 Polychaete diversity 107 Characteristics of the Cnidaria 45 Class Oligochaeta: Lumbricus terrestris 109 Class Scyphozoa 45 Earthworms and the soil 115 Aurelia: a representative scyphozoan 45 Class Hirudinea: Hirudo medicinalis 116 Class Anthozoa 47 Hirudinean diversity 119 Actinia: a representative anthozoan 48 Synopsis of phylum Annelida 120 Cnidarian classification and diversity 49 Synopsis of phylum Cnidaria 55 Phylum Arthropoda 121 Phylum Ctenophora 55 Introduction to the Arthropoda 121 Synopsis of the phylum Arthropoda 123 The acoelomate bilateral phyla 59 Subphylum Crustacea 125 Phylum Platyhelminthes 59 Astacus 125 Class Turbellaria 59 Crustacean diversity and classification 130 The planarians: examples of free-living platyhelminths 60 Class Branchiopoda 130 Characteristics of the Platyhelminthes 65 Class Ostracoda 132 Class Trematoda 65 Class Copepoda 132 Structure of a generalized trematode 65 Class Cirripedia 133 Trematode life cycles 66 Classes Mystacocarida and Branchiura 134 Class Malacostraca 134 Class Scaphopoda 186 Synopsis of the subphylum Crustacea 137 Class Aplacophora 186 Successful land arthropods 137 Class Cephalopoda: Sepia 187 Subphylum Uniramia 137 Cephalopod diversity 190 Class Insecta 137 Synopsis of class Cephalopoda 191 Locusta: the locust 137 Periplaneta: the cockroach 144 Metamorphosis 145 Early embryonic development: Insect diversity 147 the protostomes and Insect social organization 154 deuterosto mes 192 Synopsis of the main insect orders 156 The minor coelomate phyla 195 The myriapodous arthropoda 156 Class Chilopoda: the centipedes 156 Introduction to the minor coelomate phyla 195 Class Diplopoda: the millipedes 157 Class Symphyla 157 The minor protostome coelomates 195 Class Pauropoda 157 Phylum Priapuloidea 195 Phylum Sipunculoidea 196 The chelicerate arthropods 158 Phylum Echiuroidea 198 Class Merostomata: Limulus 158 Phylum Pogonophora 199 Class Arachnida: Araneus 160 Phylum Tardigrada 200 Arachnid diversity 163 Phylum Pentastomida 201 Class Pycnogonida 165 Synopsis of subphylum Chelicerata 166 The lophophorate phyla 202 Phylum Bryozoa Primitive arthropods 166 (also known as Ectoprocta or Polyzoa) 202 Subphylum Trilobitomorpha 166 Bryozoan diversity 204 Phylum Onychophora: Peripatus 167 Phylum Brachiopoda 204 Phylum Entoprocta 206 Phylum Phoronida 207 Phylum Mollusca 169 Introduction 169 The invertebrate deuterostomes 209 The molluscan plan: a hypothetical ancestor 170 Molluscan larvae 172 Phylum Echinodermata 209 Primitive living molluscs: Class Monoplacophora: Class Asteroidea: Asterias 211 Neopilina 172 Echinoderm diversity 214 Class Polyplacophora 173 Class Ophiuroidea: Ophiothrix 215 The major mollusc classes 173 Class Echinoidea: Echinus 216 The minor echinoderm classes 218 Class Gastropoda: Helix 173 Class Holothuroidea 218 Gastropod diversity 178 Class Crinoidea 220 Subclass Prosobranchia 178 Echinoderm relationships 221 Subclass Opisthobranchia 180 Synopsis of the Echinodermata 222 Subclass Pulmonata 181 Synopsis of class Gastropoda 181 The minor deuterostome phyla 222 Phylum Chaetognatha 222 Class Bivalvia: Mytilus edulis 181 Phylum Hemichordata 224 Bivalve diversity 184 Class Enteropneusta 224 Subclass Proto branchia 184 Class Pterobranchia 226 Subclass Lamellibranchia 184 Subclass Septibranchia 185 Freshwater bivalves 185 Phylum Chordata 228 Synopsis of class Bivalvia 186 Introduction to the phylum Chordata 228 Two minor mollusc classes 186 Subphylum Urochordata: Ciona 228 Urochordate diversity 232 Chondrichthyan diversity 313 Class Ascidiacea 232 Living forms 313 Class Thaliacea 232 Fossil elasmobranchs 315 Class Larvacea 233 Synopsis of Class Elasmobranchiomorphii 316 Synopsis of subphylum Urochordata 233 Subphylum Cephalochordata: Branchiostoma 234 Class Teleostomi (Osteichthyes) 317 Synopsis of the phylum Chordata 237 Salmo 317 Telestome diversity 324 The Actinopterygii 324 Introduction to the subphylum Subclass Sarcopterygii 329 Vertebrata 237 Synopsis of the class Teleostomi 331 Introduction to the tetrapods: the pentadactyl limb 332 Animal body tissues 238 Epithelial tissue 239 Class Amphibia 333 Connective tissues 240 Rana 333 Skeletal connective tissue 242 Amphibian diversity 343 Muscular tissue 245 Order U rodela 343 Nervous tissue 246 Order Anura 345 Order Apoda 346 Vertebrate organ systems 247 The fossil Amphibia: Subclasses Labyrinthodontia The integumentary system 247 and Lepospondyli 346 The skeleto-muscular system 248 Synopsis of the class Amphibia 347 The digestive system 252 Introduction to the amniotes 347 The respiratory system 257 The circulatory system 258 Class Reptilia 347 The urinogenital system 260 Lacerta 347 The nervous system 264 Reptile diversity 355 The endocrine system 273 Living reptiles 356 Fossil reptiles 360 The early development of chordates 276 Synopsis of the class Reptilia 363 The development of amphioxus 276 Vertebrate development 280 Class Aves 363 The development of the frog 280 Columba 367 Amniote development 284 Bird diversity 374 The development of mammals 288 Synopsis of the class Aves 380 Summary of chordate embryology 292 Summary of the fate of the primary germ layers 292 Class Mammalia 380 Rattus 384 Mammalian diversity 394 Vertebrate diversity 293 Subclass Prototheria (the monotremes) 395 Subclass Theria 396 Introduction 293 Infraclass Metatheria: the marsupials 396 Infraclass Eutheria 396 Class Agnatha 294 Order Insectivora 397 Lampetra 294 Order Chiroptera 397 Agnathan diversity 300 The carnivorous mammals 398 The living Agnatha 300 The small herbivorous mammals 401 Fossil agnathans: the ostracoderms 301 The large herbivores 403 Synopsis of the class Agnatha 303 The subungulates 407 Introduction to the gnathostomes 303 Some minor mammalian orders 408 Order Primates 409 Class Elasmobranchiomorphii 304 Synopsis of the class Mammalia 412 Subclass Chondrichthyes: Scyliorhinus 304