Starfish: Biology and Ecology of the Asteroidea
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STARFISH This page intentionally left blank STARFISH Biology and Ecology of the Asteroidea Edited by John M. Lawrence THE JOHNS HOPKINS UNIVERSITY PRESS | baltimore © 2013 The Johns Hopkins University Press All rights reserved. Published 2013 Printed in the United States of America on acid- free paper 9 8 7 6 5 4 3 2 1 The Johns Hopkins University Press 2715 North Charles Street Baltimore, Mary land 21218- 4363 w w w . p r e s s . j h u . e d u Library of Congress Cataloging- in- Publication Data Starfi sh : biology and ecol ogy of the Asteroidea / edited by John M. Lawrence. p. cm. Includes bibliographical references and index. ISBN 978- 1- 4214- 0787- 6 (hdbk. : alk. paper) – ISBN 1- 4214- 0787- 6 (hdbk. : alk. paper) 1. Starfi shes. I. Lawrence, John M. QL384.A8S73 2013 593.9'3–dc23 2012020461 A cata log record for this book is available from the British Library. Special discounts are available for bulk purchases of this book. For more information, please contact Special Sales at 410- 516- 6936 or [email protected]. The Johns Hopkins University Press uses environmentally friendly book materials, including recycled text paper that is composed of at least 30 percent post- consumer waste, whenever possible. Contents Contributors vii Part II • Integrative Biology Preface ix 10 Astropecten 101 Carlos Renato R. Ventura Part I • Comparative Biology 11 Luidia 109 and Ecol ogy John M. Lawrence 1 Phylogeny of the Asteroidea 3 12 Odontaster validus 120 Andrew S. Gale John S. Pearse 2 The Asteroid Arm 15 13 Acanthaster planci 132 John M. Lawrence Katharina Fabricius 3 Functional Biology of Asteroid Tube Feet 24 14 Oreaster reticulatus 142 Elise Hennebert, Michel Jangoux, Robert E. Scheibling and Patrick Flammang 15 Heliaster helianthus 153 4 Reproduction in Asteroidea 37 Juan Carlos Castilla, Sergio A. Annie Mercier and Navarrete, Tatiana Manzur, and Jean- François Hamel Mario Barahona 5 Asteroid Evolutionary Developmental Biology 16 Pisaster ochraceus 161 Carlos Robles and Ecol ogy 51 Maria Byrne 17 Asterias amurensis 174 Maria Byrne, Timothy D. O’Hara, 6 Larval Ecol ogy, Settlement, and Recruitment and John M. Lawrence of Asteroids 59 Anna Metaxas 18 Leptasterias polaris 181 Carlos F. Gaymer and 7 Ecological Role of Sea Stars from Populations John H. Himmelman to Meta- ecosystems 67 19 Coscinasterias 191 Bruce A. Menge and Eric Sanford Michael F. Barker 8 Chemistry and Ecological Role of Starfi sh 20 Echinaster 200 Secondary Metabolites 81 Richard L. Turner James B. McClintock, Charles D. Amsler, and Bill J. Baker 9 Ste roids in Asteroidea 91 References 215 Stephen A. Watts and Index 261 Kristina M. Wasson This page intentionally left blank Contributors Charles D. Amsler Juan Carlos Castilla Jean- François Hamel Department of Biology Departamento de Ecología and Society for the Exploration and University of Alabama at Centro Mileno de Conserva- Valuing of the Environment Birmingham ción Marina, Estación Costera Portugal Cove, St. Philips, Birmingham, Alabama, USA de Investigaciones Marinas, Newfoundland and Labrador, Las Cruces, Chile Canada Bill J. Baker Facultad de Ciencias Biológicas. Department of Chemistry and Universidad Católica de Chile Elise Hennebert Center for Drug Discovery Santiago, Chile Université de Mons and Innovation Laboratoire de Biologie Marine University of South Florida Katharina Fabricius Académie Universitaire Tampa, Florida, USA Australian Institute of Marine Wallonie- Bruxelles Science Mons, Belgium Mario Barahona Townsville, Queensland, Departamento de Ecología and Australia John H. Himmelman Centro Mileno de Conserva- Départment de Biologie, ción Marina, Estación Costera Patrick Flammang Université Laval de Investigaciones Marinas, Université de Mons Quebec City, Quebec, Las Cruces, Chile Laboratoire de Biologie Marine Canada Facultad de Ciencias Biológicas Académie Universitaire Universidad Católica de Chile Wallonie- Bruxelles Michel Jangoux Santiago, Chile Mons, Belgium Laboratoire de Biologie Marine Université de Mons Michael F. Barker Andrew S. Gale Académie Universitaire Department of Marine Science School of Earth and Wallonie- Bruxelles and Portobello Marine Environmental Sciences Mons, Belgium Laboratory University of Portsmouth Laboratoire de Biologie Marine University of Otago, Dunedin, Portsmouth, United Kingdom Université Libre de Bruxelles New Zealand Académie Universitaire Carlos F. Gaymer Wallonie- Bruxelles Maria Byrne Departamento de Biologia Brussels, Belgium Schools of Medical and Marina, Centro de Estudios Biological Sciences Avanzados en John M. Lawrence University of Sydney, Sydney, Zonas Áridas Department of Integrative Biology New South Wales, Universidad Católica del Norte University of South Florida Australia Coquimbo, Chile Tampa, Florida, USA viii contributors Tatiana Manzur Sergio A. Navarette Robert E. Scheibling Departamento de Ecología and Departamento de Ecología and Department of Biology Centro Mileno de Conserva- Centro Mileno de Conserva- Dalhousie University ción Marina, Estación Cos- ción Marina, Estación Costera Halifax, Nova Scotia, Canada tera de Investigaciones de Investigaciones Marinas Marinas Las Cruces, Chile Richard L. Turner Las Cruces, Chile Facultad de Ciencias Biológicas Department of Biological Facultad de Ciencias Biológicas Universidad Católica de Chile Sciences Universidad Católica de Chile Santiago, Chile Florida Institute of Technology Santiago, Chile Melbourne, Florida, USA Timothy D. O’Hara James B. McClintock Museum Victoria Carlos Renato R. Ventura Department of Biology Melbourne, Victoria, Australia Museu Nacional University of Alabama at Departamento de Invertebrados Birmingham John S. Pearse Universidade Federal do Rio de Birmingham, Alabama, USA Long Marine Laboratory Janeiro University of California, Quinta da Boa Vista, São Bruce A. Menge Santa Cruz Cristóvão Department of Zoology Santa Cruz, California, USA Rio de Janeiro, Brazil Oregon State University Corvallis, Oregon, USA Carlos Robles Kristina M. Wasson Department of Biology Department of Biology Annie Mercier California State University University of Alabama at Ocean Sciences Centre at Los Angeles Birmingham Memorial University Los Angeles, California, USA Birmingham, Alabama, USA St. John’s, Newfoundland and Labrador, Canada Eric Sanford Stephen A. Watts Department of Evolution and Department of Biology Anna Metaxas Ecol ogy University of Alabama at Department of Oceanography University of California Birmingham Dalhousie University Davis, California, USA Birmingham, Alabama, USA Halifax, Nova Scotia, Bodega Marine Laboratory Canada Bodega Bay, California, USA Preface tarfi sh are icons of the sea. But they also have im mense biological Sand ecological importance. They are one of the major classes of Echinodermata, with a distinctive and varied body form. Starfi sh live in a variety of marine habitats where they are major predators that greatly aff ect their communities. With a long fossil record, starfi sh are of tremendous interest to paleontologists and evolutionary biologists. Although study of starfi sh biology began in the second half of the nineteenth century, many basic aspects of their physiology, nutrition, reproduction, and even anatomy still are not well known. Study of starfi sh ecol ogy began in the fi rst half of the twentieth century but has been limited primarily to species whose eff ect can be conspicuous, in- cluding the crown- of- thorns starfi sh on coral reefs and species that af- fect shellfi sh fi sheries. Effects of other species are not as well understood because they are not as accessible or conspicuous and they do not have an economic impact. This book provides a comprehensive understanding of the biology and ecol ogy of starfi sh. It is divided into two sections: comparative chapters that consider aspects of biology and ecol ogy across the class and integrative chapters that consider aspects of biology and ecol ogy of individual species or genera. These approaches are complementary. Species and genera were selected for the integrative chapters based on the scope of knowledge about them and the availability of experts to be authors. I am grateful to my friends and colleagues, experts in their fi elds, who have contributed to this book. It is a plea sure to thank executive editor Vincent Burke, Jennifer Malat, and Michele Callaghan at the Johns Hopkins University Press for their kind assistance in its production. This page intentionally left blank STARFISH This page intentionally left blank PART I • COMPARATIVE BIOLOGY AND ECOL OGY This page intentionally left blank 1 Phylogeny of the Asteroidea Andrew S. Gale he phylogeny of asteroids has been a controversial topic for more than 100 years. The detailed morphology of spines and Tpedicellariae formed the basis of an ordinal classifi cation by Per- rier (1884, 1894), consisting of the Paxillosida, Spinulosida, Valvatida, and Forcipulatida, which have obtained widespread use among neon- tologists. Apart from some rearrangement of taxa included variously in the Spinulosida and Valvatida (Blake 1981) and the resurrection of Perrier’s Velatida (Blake 1987; Fig. 1.1a), the classifi cation has been re- markably stable. In contrast, the interrelationships of the orders have remained obscure and controversial. Fossil asteroids have been known since the early nineteenth century. It was clear from early studies (e.g., Forbes 1848) that many Jurassic and later asteroids were surprisingly similar to extant taxa at the level of families or even genera. Concomitantly, it became increasingly clear that Paleozoic