f- \AJC I tC IC COMPARATIVE MORPHOLOGY OF RECENT CRUSTACEA COMPARATIVE MORPHOLOGY OF RECENT CRUSTACEA PATSY A. MCLAUGHLIN COURTESY PROFESSOR, FLORIDA INTERNATIONAL UNIVERSITY, 1977- ADJUNCT PROFESSOR, FLORIDA ATLANTIC UNIVERSITY, 1979- HR W. H. FREEMAN AND COMPANY San Francisco Sponsoring Editor: Arthur C. Bartlett Project Editor: Patricia Brewer Copyeditor: Sean Cotter Designers: Perry Smith and Sharon Smith Production Coordinator: William Murdock Illustration Coordinator: Cheryl Nufer Compositor: Graphic Typesetting Service Printer and Binder: The Maple-Vail Book Manufacturing Group Library of Congress Cataloging in Publication Data McLaughlin, Patsy A Comparative morphology of Recent Crustacea. Bibliography: p. Includes index. 1. Crustacea—Anatomy. I. Title. QL445.M25 595'.3'044 79-26066 ISBN 0-7167-1121-4 Copyright © 1980 by W. H. Freeman and Company No part of this book may be reproduced by any mechanical, photographic, or electronic process, or in the form of a phonographic recording, nor may it be stored in a retrieval system, transmitted, or otherwise copied for public or private use, without written permission from the publisher. Printed in the United States of America 987654321 Dedicated to Professor Dr. Lipke B. Holthuis and the memory of Dr. Waldo L. Schmitt (1887-1977) Two outstanding carcinologists CONTENTS INTRODUCTION 1 Basic Characters of the Superclass Crustacea 2 Classification of the Crustacea 3 CLASS CEPHALOCARIDA 4 CLASS BRANCHIOPODA 8 Order Notostraca 8 Order Anostraca 11 Order Conchostraca 16 Order Cladocera 19 CLASS OSTRACODA 23 CLASS MYSTACOCARIDA 28 CLASS COPEPODA 32 Free-Living Copepoda 33 Order Calanoida 33 Order Harpacticoida 33 Order Cyclopoida 33 Order Misophrioida 34 Parasitic Copepoda 37 Order Notodelphyoida 37 Order Monstrilloida 37 Order Siphonostomatoida 38 Order Poecilostomatoida 38 CLASS BRANCHIURA 41 CLASS CIRRIPEDIA 44 Order Acrothoracica 44 Order Thoracica 46 Suborder Lepadomorpha 47 Suborder Verrucomorpha 51 Suborder Balanomorpha 52 Order Ascothoracica 54 Order Rhizocephala 56 viii CONTENTS CLASS MALACOSTRACA 59 Subclass Phyllocarida 59 Order Leptostraca 59 Subclass Hoplocarida 63 Order Stomatopoda 63 Subclass Eumalacostraca 70 Superorder Syncarida 70 Order Anaspidacea 70 Order Stygocaridacea 74 Order Bathynellacea 75 Superorder Peracarida 79 Order Mysidacea 79 Order Thermosbaenacea 84 Order Spelaeogriphacea 87 Order Cumacea 88 Order Tanaidacea 91 Order Isopoda 95 Free-Living Isopoda 95 Suborder Valvifera 95 Suborder Anthuridea 96 Suborder Flabellifera 96 Suborder Microcerberidea 97 Suborder Asellota 98 Suborder Phreatoicidea 98 Suborder Gnathiidea 99 Suborder Oniscoidea 100 Parasitic Isopoda 106 Suborder Epicaridea 106 Order Amphipoda 109 Gammaridea and Hyperiidea 109 Suborder Gammaridea 109 S uborder Hyperiidea 110 Ingolfiellidea and Caprellidea 114 Suborder Ingolfiellidea 114 Suborder Caprellidea 114 Superorder Eucarida 117 Order Euphausiacea 117 Order Amphionidacea 123 Order Decapoda 126 Suborder Dendrobranchiata 127 Suborder Pleocyemata 132 Infraorder Stenopodidea 132 Infraorder Caridea 133 Infraorder Astacidea 137 Infraorder Austroastacidea 142 Infraorder Palinura 143 Infraorder Anomura 144 Infraorder Brachyura 151 GENERAL CRUSTACEAN REFERENCES 159 GLOSSARY OF MORPHOLOGICAL TERMS INDEX 175 PREFACE Since Caiman s monograph on crustacean morphology was published shortly after the turn of the century, our knowledge of crustacean biology has in­ creased manyfold. Recent interest in the roles played by crustaceans in the environment has taken investigations of this highly diverse group of inver­ tebrates out of the exclusive realms of basic science and fisheries and made them topics of applied research as well. As a result, the literature pertaining to crustaceans is not only prolific, but also widely scattered and often highly specialized and technical. This book summarizes much of the more recently acquired basic data and the current interpretations of existing information. Its format has been de­ veloped with two purposes in mind. The first is the presentation of the material in the form of a text in crustacean biology, complete with step-by- step directions for the study of specific groups. The second is to provide a concise reference to the major morphological and anatomical characters of the higher crustacean taxa for both specialists and nonspecialists. Many evolutionary and phylogenetic carcinological questions have not yet been resolved. Consequently not all carcinologists will agree with the clas­ sification and terminology employed in this text. With regard to the classifi­ cation, I have adopted a conservative approach, preferring to await further evidence before taking the major step of recognizing the Crustacea as a distinct phylum. With regard to terminology, although I recognize and ap­ preciate the usefulness of specialized terms in particular situations, I believe that the use of uniform terms for homologous structures reduces confusion and makes the subject matter more comprehensible to the general reader. The field of carcinology is a vast and exciting one in which our present level of scientific knowledge is still relatively limited. I hope that this book provides a basic knowledge of the Crustacea upon which future carcinologists will build. October 1979 Patsy A. McLaughlin ACKNOWLEDGMENTS I am deeply indebted to Drs. L. G. Abele, Florida State University; J. S. Garth, Allan Hancock Foundation, University of Southern California; and D. P. Henry and P. L. Illg, University of Washington, for data and specimens contributed to this study. Much valuable information has been gained by the careful critical reviews of a number of the world's finest carcinologists: Drs. J. S. Garth and R. Brusca and Ms. J. Haig, University of Southern California; Drs. T. E. Bowman, F. A. Chace, Jr., R. F. Cressey, H. H. Hobbs, Jr., B. Kensley, L. S. Kornicker, and R. B. Manning, National Museum of Natural History, Smithsonian Institution; Drs. D. P. Henry and P. L. Illg, University of Washington; Dr. A. J. Provenzano, Jr., Old Dominion University; and Dr. L. B. Holthuis, Rijksmuseum van Natuurlijke Historie, Leiden, The Nether­ lands; however, final responsibility for inaccuracies and omissions must rest with the author. I also wish to gratefully acknowledge the most valuable assistance of Mr. J. Garcia-Gomez, University of Miami, in acquiring much of the necessary literature and of Mrs. S. F. Treat, Florida International University, for her diligent proofreading of the manuscript. Special thanks also are due my husband, Mac, for his assistance, patience, and encourage­ ment. The permission granted by the following institutions, journals, publishers, and authors to adapt illustrations for which they hold the copyright is grate­ fully acknowledged: Albert Bonniers Forlag, Stockholm, representing Acta Zoologica; Akademiya Nauk SSSR, representing Paleontologischeskii Zhur- nal; A. Asher and Company, Amsterdam; Bergen Museum; Biological Soci­ ety of Washington; British Museum (Natural History); Bulletin of Marine Science; Lund University, representing Lunds Universitets Arsskrift; Cam­ bridge University Press, London, representing the Journal of the Marine Biological Association, United Kingdom; National Institute of Oceanography, Wormley, representing the Discovery Reports; Clarendon Press, Oxford, representing the Quarterly Journal of Microscopical Science; E. J. Brill, representing Crustaceana; Geological Society of America; Hermann, Paris; Vie et Milieu; John Wiley and Sons; Royal Netherlands Academy of Sciences at Amsterdam; the University of Kansas Press; Veb Gustav Fisher Verlag; and Dr. Robert Hessler, Scripps Institution of Oceanography. P.A.M. COMPARATIVE MORPHOLOGY OF RECENT CRUSTACEA INTRODUCTION The superclass Crustacea is one of the largest, most di­ morphology of the diverse groups, another is to provide verse, and most successful groups of invertebrates. The an easy reference to major structures for specialists. commercial importance of numerous species of shrimps, Every attempt has been made to combine a uniform crabs, and lobsters has long been recognized. Lim- description of morphological structures with the spe­ nologists have been aware of the contributions made by cialized interpretations of carcinologists working with the the numerous small crustacean groups to lake and stream various taxa. This has not always been possible, and ecosystems. Similarly, fishery biologists have learned when a choice has been required, it has been made with that the variety of planktonic crustaceans play a signifi­ a preference to the basic morphological patterns. For cant role in the food chains of fishes. More recently, instance, in copepods and several other major taxa, benthic biologists and ecologists have discovered that a specialists have tended to disregard thoracic somites that broad spectrum of benthic crustaceans serve vital roles in have been incorporated into the cephalon through fu­ trophic levels and energy cycles within communities. sion. Where 2 thoracic somites are fused to the cephalon, The literature on some of the taxa, such as the the adjacent somite is morphologically the 3rd thoracic copepods and decapods, is extremely extensive, highly somite; however, many specialists would call it the 1st. technical and specialized, and not readily adapted for Each of the thoracic somites (thoracomeres) basically is general usage. In contrast, literature on some of the less provided with a pair of appendages, referred to herein as familiar or lesser known groups is extremely