PRACTICAL INVERTEBRATE ANATOMY by w. s. BULLOUGfl Ph.D., D.Sc.(Leeds} Professor of Zoology in the University of London UNDP I SF I f'AO P1SH£RIES PROJECT NO. 7G LllRARY SECOND EDITION LONDON MACMILLA... & CO LTD NEW YORK • ST MARTIN'S PRESS PRACTICAL INVERTEBRATE ANATOMY .. (... ,.::,r I f 0 FISf1.EfI!ES PROJlCr NO. "}(I LIBRARY © w. S. Bullough 1958 Fint Edition J 950 Reprinted 1951, 1954- _ Second Edition 1958 Reprinted 1960,1964.1966. 1968 MACMILLAN AND COMPANY LIMITED Litt/e Essex Street London WC 2 a/so Bombay Calcutta Madras MelbojJrne THE MACMILLAN COMPANY OF CANADA LIMITED 70 Bond Slrut Toronto 2 ST MARTIN'S PRESS INC 175 Fijth At'lmue Nero York NY roOl0 2628 PRINTED IN GREAT BRITAIN PREFACE TO THE SECOND EDITION TN 1954 de Beer * wrote: "Few ... zoologists could have thought it possible that, in the middle of the twentieth century, all text-books of zoology should require the drastic revision of the current theory of the origin and evolution of the Metazoa. Yet tha.t and nothing less is the situation resulting from the researches which for half a century Jovan Hadzi has made into the structure and development of the lower animals. They have led him to put forward a theory that multinuclear Protozoa evolved into Turbellaria Acoela, which latter (not the Coelenterata) would be the most primitive Metazoa." Close examination of Hadzi's theory, which de Beer explained for the first time in English, indicates tha.t it offers what is probably the most satisfactory explana tion of the facts as they are now known, and consequently this second revised edition has been prepared. Recently Hadzi has written an extendcd explana tion of his theory in English. t In this second edition the platyhelminths are examined before the coelenter ates, the turbellarians are considered in greater detail, and in particular a description is given of thc acoelan genus Convoluta. Within the coelenterates, the actinozoans are treated first as the most primitive class, and the terms " diploblastic" and" triploblastic" have been abandoned. The opportunity has also been taken to include two new chapters and to introduce much extra information which, during the years, has been offered by many helpful critics. In particular I am indebted to Dr. 1. Griffiths for his generous help with the descriptions of the Turbellaria and the Nemertea. W. S. BULLOUGH Birkbeck CoUege, University of London December 1957 • de Beer, G. R. (1954), "The Evolution of the Metazoa", Chapter 2 of Evolution as a P"ocess, edited by J. Huxley. London. t Hadzi, J. (1963), The Evolution of the Metazoa. London. v PREFACE TO THE FIRST EDITION THE main intention of this book is to provide brief and practical descriptions of the anatomy of those invertebrate animals which are commonly used as types in the study of zoology. Previously, such descriptions, if they existed at all, were widely scattered and not always easy either of discovery or of access. Furthermore, they were not always made from a practical point of view, or accompanied by figures which had any great practical value. To meet this situCJtion, descriptions have been made of 122 commonly studied genera. These genera are primarily selected for their availability in the British Isles and in North Am rica, but in most cases they, or closely related forms, also occur all over the world. In making the descriptions it was soon found that limits had to be imposed in order to keep the book to a reasonable size. Thus, it was decided to include only those features which can be demonstrated in whole specimens of adult animals, and, in particular, to exclude any consider ation of prepared sections and of young or larval forms. In a few instances it has been found necessary to make exceptions, but in general this rule has been closely observed. It is felt to be highly im.portant that each genus studied should, as far as possible, be appreciated in its natural systematic, ecological, and functional setting. For this reason, the details of classification have been briefly inserted into the text, the description of each animal has been prefaced by a short account of its distribution and mode of life. and notes on the function and special significance of unusual organs or structures have been briefly interpolated. By these various means it is hoped that the anatomical details may assume a maximum significance and interest. r n cas further information should be required. references to more detailed descriptions are given wherever it has been possible to trace them. and at the nds of most of the chapters appendices are included which describe the methods of culture, where these arc practical, of killing, fixing, and preserving, and of any special techniques which have proved valuable. The general appendix at the end of the book includes details of the composition of the fixatives, stains, alld other solutions which are mentioned in the text. The book is centred round, and originated with, its figures which have all been newly drawn from preparations or dissections. They were first made for the use of classes in McGill University, Montreal, when it became apparent how inadequate for practical purposes were most of the current textbook vii PREFACE V1l1 illustrations. The figures are all semi-diagrammatic, and it should be unnecessary to state that they are in no way intended as substitutes for the student's personal drawings of his own specimens. They are designed to save time and bewilderment, and not to replace personal investigation and effort. Many debts of gratitude have been incurred during the writing of this manuscript, and the first of these, since they concerned the beginning of the book, are to Professor N. J. Berrill and the students in the Zoology Department at McGill University. Professor Berrill did all in his power to provide the necessary facilities for the work, most of which was completed in his department, and he also readily gave his special knowledge and critical attention to the chapters on the coelenterates and the chordates. The students, of whom Mr. P. A. Orkin may be specially mentioned, formed an eager and critical grou p with whose help the figures, descriptions, and techniques could be tested. Also at McGill University, constant encouragement and advice were received from Professor V. C. Wynne-Edwards, who made particular comm~nts on the chapter dealing with the echinoderms, and from Professor T. W. M. Cameron, who provided much material for, and was repeatedly consulted about, the descriptions of parasitic protozoans, platyhelminthes. nematodes. acanthocephalans. and arachnids. Later, in the University of Sheffield, facilities for finishing the work were afforded by Professor L. E. S. Eastham. Also in England, much information has been gratefully received from Mr. T. Kerr on the platyhelminthes, Dr. T. B. Reynoldson on the annelids, Dr. F. Segrove on the myriapoo.s. and Professor L. E. S. Eastham and Dr. H. Henson on the insects. Other help, which it is a pleasant duty to recall, came from Mrs. L. Terrill, who, as librarian in tht' Blacker Library of Zoology in McGill University, gave tireless assistance in the often difficult and tedious task of tracing important references; and from Mr. J. Hancock, who, as laboratory steward in the Sheffield Department of Zoology, has given unreservedly from his extensive knowledge of practical techniques. Finally, a special mention must be made of my wife who. in a hundred ways, has helped in the production of this book from its beginning to its end. W. S. BULLOUGH Unive'Ysity of Sheffield December 1947 CONTENTS P.G~ Preface to Second Edition v Prefact: to First Edition vii CHAPTER I. PHYLUM PROTOZOA 1.00. Class Mastigophora (Flagellata) I. 10. Subclass Phytomastigina I. II. Order Euglenoitlida- Euglrna 1.12. Order Dinoflagellida-Ceratillnl 4 1.13. Order Phytomonadida (Volvocina)- Chlamydomonas 5 1.20. Subclass Zoomastigina 7 1.21. Order Protomonadida--Trypanosoma 7 2.00. Class Sarcodina (Rhizopoda) . 9 2.01. Order Lobosa (Amoebina)-Amoeba . 9 -Entamoeba II 2.02. Order Heliozoa-A ctinosphaerium I:2 2.03. Order Foraminifera-Polystomella I" 2.04. Order Radiolaria- Sphaerozotm, 16 3.00. Class Spor02:oa 18 3.10. Subclass Telosporidia. 18 3. I l. Order Grcgarinida-Mo nocystis 18 3.12. Order Coccidia- Eimeria. 20 3.13. Order Hacmosporidia-PlasmodiIHII . 22 4.00. Class Infusoria (Ciliophora) 25 4.10. Subclass Protociliata . 25 ... II. Order Opalinata-Opalina 25 4.20. Subclass Euciliata 27 4.21. Order Holotrlchida-Paramecillm 27 4.22. Order Heterotrichida-Nyctotherlls 30 ".23. Order Peritrichida- Vorticella 32 Appendix to the Protozoa 34 CHAPTER II. PHYLUM PORIFERA 1.00. Class Calcarea 1.01. Order Heterocoela-Sycon 2.00. Class Demolpongiae 2.01. Order Monaxonida-Ephydatia 2.02. Order Keratosa-Ettspongia (Sponcia) Appendix to the Porifera IX x CONTENTS CHAPTER III. PHYLUM PLATYHELMINTHES 1.00. Class Turbellaria . J .01. Order Acoela--Convolllta 1.02. Order Rhabdocoela-Mesostotna 1.03. Order Polycladida-Lrptoplana 1.04. Order Tricladidll-Dendt'ocoelum 1.05. Order Temnocephalida- Temnocephala 2.00. Class Trematoda 2.10. Subclass Heterocotylea (Monogenea) . 2.II. Order Polyopisthocotylea-Polyslomum 2.20. Subclass Malacocotylea (Digenea) 2.21. Order Prosostomata-Fasciola . -Schistosoma (Bilharzia) 3.00. Class Cestoda 3.10. Subclass Merozoa 3.11. Order Pseudophyllidea-Diphyllobothrium (Dibothriocephalus) 3.12. Order Cyclophyllidea-Taenia Appendix to the Platyhelminthes . CHAPTER IV. PHYLUM COELENTERATA 1.00. Class Actinozoa (Anthozoa) . 78 1.10. Subclass Alcyonatia 78 1.11. Order Alcyonacea-Alcyonium 79 1.12. Order Gorgonacea-Gol'gcnia 81 1.13. Order Pennatuli'cea-Pennatula 84 1.20. Subclass Zoantharia 86 1.21. Order Actiniaria-A ctinia. 86 1.22. Order Madreporaria-Ast,.angia go 2.00. Class Hydrozoa (Hydromedusae) 94 2.01. Order Gymnoblastea (Antbomedusae)- Bougainvillea 94 -Hyd,.a 97 2.02. Order Calyptoblastea (Leptomedusae)-Obelia . roo 2.03. Order Ttachylina-Gonionemus 104 2.04. Order Siphonophora-Physalia J06 3.00. Class Scyphomedusae (Scyphozoa) . log 3.01. Order Stauromedusae- Haliclysll4s J09 3.02. Order Discomedusae-Aurelia 111 Appendix to the Coelenterata u6 CHAPTER V. PHYLUM CTENOPHORA 1.00. Class Tentaculata . JI8 1.01. Order Cydippida-Pleu,.oll1'achia uB ~.oo. ClaRs Nuda J21 2.01. Order Beroida-Beroe 121 CONTENTS XI PAOE CHAPTER VI. PHYLUM NEMERTEA 1.00. Class Enopla . 124 1.or. Order Hoplonemcrtini-Amphip<wus 124 1.02. Order Bdclloncmcttini-Malacobdella 127 CHAPTER VII. PHYLUM NEMATODA o.or. Order Ascaroidca- Rhabdilis -Ascaris 0.02. Order Strongyloidea-A11cylosloma 0.03. Order Trichinclloidea- Tricllinrlla Appendix to the Nematoda CHAPTER VIII. PH1'LUM ACANTHOCEPHALA -Macracanthorhynchus (Echinorhynchus) CHAPTER IX. PHYLUM ROTIFERA 0.01. Order Monogonata- Hydalina (Epiphanes) Appendix to the Rotifera CHAPTER X. PHYLUM ENDOPROCTA - Pedicelli'la 153 CHAPTER XI. PHYLUM ECTOPROCTA 0.01. Order Gymnolaemata-Bug1l.1a . 0.02. Order Phylactolaemata- Pecti,lateJla Appendix to the Ectoprocta • HAPTER XII. PHYLUM BRACHIOPODA 0.01. Order Ecardinea-Lingula 162 0.02. Order Teaticardinea-Te,.eb,.atella 165 Appendix to the Brachiopoda 169 CHAPTER XIII. PHYLUM CHAETOGNATHA -Sagitta Appendix to the Chactognatha CHAPTER XIV. PHYLUM ANNELIDA 1.00. Class Chactopoda . 1.01. Order Polychaeta-Nereis -Arenicola -Amphit,..jte -Sabella xii CONTENTS rAOt: 1.02. Order Oligochaeta-Lumbricus 18b -EIlchJ'lralus 193 2.00. Class Hirudinea 196 - Hirudo. 196 3.00. Class Gephyrca 201 3.01. Order Echiuroidca-BOt.ellia 201 3.02. Order Sipunculoidea--Sipu'lcl~lus 203 Appendix to the Annelida 205 CHAPTER XV. PHYLUM ARTHROPODA 1.00. Class Onychophora 208 -Peripatus 208 CHAPTER XVI. PHYLUM ARTHROPODA (cotltim4ed) 2.00. Class Crustacea 212 2.10. Subclass Branchiopoda 212 2.11. Order Anostraca-ElIbranchiptlS 212 2.12. Order Notostraca- Apus . 216 2.13. Order Cladocera- Dap}mia 219 2.20. Subclass Copepoda 223 2.21. Order Eucopepoda-Cyclops 223 2.30. Subclass Cirripedia 227 2.31. Order Thoracica-Lepas . 227 2.40. Subclass Malacostraca 231 2·41. Order Lepto.tta"a-Nebalia 231 2.42• Order Mysidacea-Mysis . 234 2.-43. Order Isopoda-Oniscus 236 2·44· Order Amphipoda-Gammar'lls . 239 2·45· Order Decapoda- A slaclis 242 CHAPTER XVll. PHYLUM ARTHROPODA (continlled) 3.00. Class Myriapoda 3.01. Order Chilopoda-LithobitlS 3.02. Order Diplopoda- ltllllol . CHAPTER XVITI. PHYLUM ARTHROPODA (continued) 4.00. Class Insecta . 4.10. Subclass Apterygota ... n . Order Thysanura- Petrobius (Mucllilis) 4.12. Order Collembola- Tomocerus . -1.20. Subclass Exopterygota (Heterometabola) 4.21. Order Orthoptera- Blalta ... 22. Order Isoptera-Termes • ",.23. Order Ephemeroptcra- Epllemer"
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