Table Of ContentFunctional 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
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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 contri-
butions, 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 crus-
taceans 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
