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Functional Morphology and Diversity Functional Morphology and Diversity. Les Watling and Martin Thiel. © Les Watling and Martin Thiel 2013. Published 2013 by Oxford University Press. The Natural History of the Crustacea Series SERIES EDITOR: Martin Thiel Editorial Advisory Board: Geoff Boxshall, Natural History Museum, London, UK Emmett Duffy, Virginia Institute of Marine Sciences, Gloucester, USA Darryl Felder, University of Louisiana, Lafayette, USA Gary Poore, Victoria Museum, Melbourne, Australia Bernard Sainte-Marie, Fisheries and Oceans Canada, Mont-Joli, Canada Gerhard Scholtz, Humboldt University Berlin, Berlin, Germany Fred Schram, Friday Harbor Marine Laboratory, Seattle, USA Les Watling, University of Hawaii, Hawaii, USA Functional Morphology and Diversity (Volume 1) Edited by Les Watling and Martin Thiel Functional Morphology and Diversity The Natural History of the Crustacea, Volume 1 EDITED BY LES WATLING AND MARTIN THIEL 1 3 Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trademark of Oxford University Press in the UK and certain other countries. Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016 © Oxford University Press 2013 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above. You must not circulate this work in any other form and you must impose this same condition on any acquirer. Library of Congress Cataloging-in-Publication Data Functional Morphology and Diversity p. cm—(The Natural History of the Crustacea series). Includes bibliographical references and index. ISBN 978-0-19-539803-8 (hardcover: alk. paper) 1. Crustacea. I. Watling, Les. II. Thiel, Martin, 1962– QL435.N38 2013 595.3—dc23 2012012251 9 8 7 6 5 4 3 2 1 Printed in the United States of America on acid-free paper PREFACE Many years ago, when M.T. came to Maine to work with L.W. on shallow subtidal crustaceans, we established some amphipod populations in aquaria at the University of Maine’s Darling Marine Center flowing seawater facility. What ensued was not just a series of remarkable behav- ioral observations but also a series of long discussions of “how do they do that?” in response to the burrowing and whip making that we observed. The two of us have maintained a lifelong interest in understanding how crustacean morphology “determines” what the animals can do and where they can live. Crustaceans encompass a bewildering array of body forms that they use to occupy almost every habitat type on Earth. In fact, the only habitat not occupied by crustaceans is the open air; that is, they cannot fly. Understanding what modifications of the crustacean body plan have allowed this diversification is the subject of this book. In its basic form, the crustacean body consists of a bilaterally symmetrical, segmented, more or less tubular body, with each segment bearing a pair of appendages. The number of body seg- ments and the design or even presence of appendages vary considerably, with the most extreme cases being found in those groups that have invaded the bodies of other animals. In this book we have asked the chapter authors to explore this diversification of form and to explain how various parts of the crustacean body work. While many authors have examined the functional morphology and anatomy of crustaceans in individual publications or book contributions, this has never been done in an integral way in one volume. We were particularly interested in understanding the design limitations of the crustacean body, for example, how living in a dense fluid medium might restrict the movement capabilities of the animal, and how that would vary depending on the animal’s size. But movement isn’t all that crustaceans do in their daily lives—they have to eat, respire, reproduce, and grow, all of which needs to be controlled so that the animal functions as a successfully coordinated whole. To set the stage, Schram (chapter 1) reviews the range of crustacean body plans, and Haug et al. (chapter 2) review what is known about appendage development in the earliest arthro- pods as they become what we would recognize now as crustaceans. Then follow four chapters that examine the exterior of the crustacean body. Williams (chapter 3) investigates the genetic control of appendage development as the animal develops. Olesen (chapter 4) surveys the functional constraints of the archetypical crustacean structure, the carapace. Dillaman et al. (chapter 5) take a detailed look at the crustacean cuticle, and Garm and Watling (chapter 6) review the structure and function of setae and other cuticular outgrowths. v Functional Morphology and Diversity. Les Watling and Martin Thiel. © Les Watling and Martin Thiel 2013. Published 2013 by Oxford University Press. vi Preface Our examination of the integrated functional aspects of the crustacean body starts with Boxshall and Jaume’s overview of crustacean antennae (chapter 7). Watling (chapter 8) inte- grates food consumption and digestion, showing that mouthpart appendage structure, foregut morphology, and digestive enzyme secretion are all related to diet. Belanger (chapter 9), Faulkes (chapter 10), Yen (chapter 11), and Boudrias (chapter 12) examine modes of locomotion in crustaceans and the functional constraints imposed on locomotory appendages by the physics of the medium in which they live. We end the book with a series of chapters dealing with system-level integrative functional anatomy. Bauer (chapter 13) reviews the structure and function of appendages used for groom- ing and reproduction. Wirkner and Richter (chapter 14) examine the integration of respiratory structures with internal details of the circulatory system. An overview of the internal anatomy of the reproductive system is provided by L ó pez Greco (chapter 15). And lastly, the coordina- tion of the whole body is dealt with in Sullivan and Herberholz’s review of crustacean nervous systems (chapter 16). We have asked the authors of these chapters to deal with their part of the crustacean body in isolation. Of course, we recognize that crustaceans are living creatures, and in order to live they have managed to integrate the functional aspects of their bodies that we have studied so well on their own. What we have not done is try to develop a fully integrated model of the crustacean body that accounts for all the physical aspects of the various habitats in which crustaceans live. We suspect this is possible but may be beyond the reach of any one of us. ACKNOWLEDGMENTS Our thanks go foremost to all contributors who attended to all our requests during the prep aration of this book. Their expertise and their willingness to invest time into the writing of their contributions make up the value of this book. We especially thank our editorial assistants, Ivan Hinojosa and Lucas Eastman, who expertly revamped many of the figures and who care fully read and edited all the chapters. The generous contribution from Universidad Catolica del Norte was essential for this project—we are very grateful for the continuous support which allowed us to focus on the task. The vision and foresight of the university authorities made this project possible and we hope that this and the upcoming volumes fulfil their expectations. We also thank those colleagues who read entire or parts of chapters. The publisher, Oxford University Press, gave us a lot of freedom, and in particular, we express our appreciation to Peter Prescott, Tisse Tagaki, Jeremy Lewis, and Hallie Stebbins for their help during the past few years. L.W. would like to acknowledge the role of his students in stimulating him to think about crustaceans as functionally whole units, and his friend C. Nouvian, who listened with interest to many crustacean natural history stories while we were working on other things. Finally, we thank our families for their patience and interest in this project—without their tolerance, none of this would have been possible. Editing of this book was generously supported by Universidad Catolica del Norte, Chile. CONTRIBUTORS EDITORS Michel A. Boudrias Martin Thiel Department of Marine Science and Facultad Ciencias del Mar Environmental Studies Universidad Catolica del Norte University of San Diego Larrondo 1281 Science and Technology 267 Coquimbo 5998 Alcala Park Chile San Diego, CA 92110 USA Les Watling Department of Biology Geoff Boxshall University of Hawaii at Manoa Department of Zoology 152 Edmondson Hall The Natural History Museum Honolulu, HI 96822 Cromwell Road USA London SW7 5BD UK AUTHORS Raymond T. Bauer Richard M. Dillaman Department of Biology Department of Biology and Marine Biology University of Louisiana University of North Carolina at Wilmington PO Box 42451 601 South College Road Lafayette, LA 70504 Wilmington, NC 28403-5915 USA USA Jim Belanger Zen Faulkes Department of Biology Department of Biology West Virginia University The University of Texas-Pan American PO Box 6057 1201W. University Drive Morgantown, WV 26506 Edinburg, TX 78539 USA USA Contributors ix Anders Garm Andreas Maas Department of Cell and Organism Biology Biosystematic Documentation Lund University University of Ulm Helgonavagen 3 Helmholtzstrasse 20 222362 Lund 89081 Ulm Sweden Germany Carolin Haug Shannon Modla University of Greifswald Delaware Biotechnology Institute Zoological Institute and Museum 15 Innovation Way, Suite 117 Department of Cytology and Evolutionary Newark, DE 19716 Biology USA Soldmannstr. 23 17487 Greifswald Germany Jorgen Olesen Natural History Museum of Denmark Joachim T. Haug (Zoological Museum) University of Greifswald University of Copenhagen Zoological Institute and Museum Universitetsparken 15 0 Department of Cytology and Evolutionary DK-2100 Copenhagen Biology Denmark Soldmannstr. 23 17487 Greifswald Stefan Richter Germany Universitat Rostock Institut fur Biowissenschaften Jens Herberholz Abteilung fur Allgemeine und Spezielle Department of Psychology and Zoologie Neuroscience and Cognitive Science Universitatsplatz 2 Program 18055 Rostock 2123H Bio-Psych Building Germany University of Maryland College Park, MD 20742 Robert Roer USA Department of Biology and Marine Biology Damiajaume University of North Carolina at Instituto Mediterraneo de Estudios Wilmington Avanzados IMEDEA (CSIC-UIB) 601 South College Road Miquel Marques 21 Wilmington, NC 28403-5915 07190 Esporles (Mallorca, Illes Balears) USA Spain Laura S. Lopez Greco Frederick R. Schram Facultad de Ciencias Exactas y Naturales Burke Museum University of Buenos Aires University of Washington at Pabellon 2, Intendente Giiiraldes 2160 Seattle Ciudad Universitaria C1428EGA Post Box 1567 Buenos Aires Langley, WA 98260 Argentina USA