The Leatherback Turtle This page intentionally left blank The Leatherback Turtle Biology and Conservation Edited by James R. Spotila and Pilar Santidrián Tomillo J OHNS H OPKINS U NIVERSITY P RESS | B ALTIMORE © 2015 Johns Hopkins University Press All rights reserved. Published 2015 Printed in the United States of America on acid- free paper 9 8 7 6 5 4 3 2 1 Johns Hopkins University Press 2715 North Charles Street Baltimore, Maryland 21218- 4363 www.press.jhu.edu Library of Congress Cataloging- in- Publication Data The leatherback turtle : biology and conservation / edited by James R. Spotila and Pilar Santidrián Tomillo. pages cm Includes bibliographical references and index. ISBN 978- 1- 4214- 1708- 0 (hardcover : alk. paper) — ISBN 1- 4214- 1708- 1 (hardcover : alk. paper) — ISBN 978- 1- 4214- 1709- 7 (electronic) — ISBN 1- 4214- 1709- X (electronic) 1. Leatherback turtle. 2. Leatherback turtle—Conservation. I. Spotila, James R., 1944– editor. II. Santidrián Tomillo, Pilar, 1974– editor. QL666.C546L43 2015 597.92'89—dc23 2014036887 A catalog 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]. 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. To Laurie Dittrich Spotila This page intentionally left blank Contents List of Contributors ix Preface xi Part I. BIOLOGY 1 Introduction: Phylogeny and Evolutionary Biology of the Leatherback Turtle 3 peter c. h. pritchard 2 Phylogeny, Phylogeography, and Populations of the Leatherback Turtle 8 peter h. dutton and kartik shanker 3 Diving Behavior and Physiology of the Leatherback Turtle 21 nathan j. robinson and frank v. paladino 4 Anatomy of the Leatherback Turtle 32 jeanette wyneken Part II. LIFE HISTORY AND REPRODUCTION 5 Reproductive Biology of the Leatherback Turtle 51 david c. rostal 6 Nesting Ecology and Reproductive Investment of the Leatherback Turtle 63 karen l. eckert, bryan p. wallace, james r. spotila, and barbara a. bell 7 Egg Development and Hatchling Output of the Leatherback Turtle 74 pilar santidrin tomillo and jennifer swiggs 8 Sex Determination and Hatchling Sex Ratios of the Leatherback Turtle 84 christopher a. binckley and james r. spotila viii Contents Part III. POPULATION STATUS AND TRENDS 9 Leatherback Turtle Populations in the Atlantic Ocean 97 marc girondot 10 Leatherback Turtle Populations in the Pacific Ocean 110 scott r. benson, ricardo f. tapilatu, nicolas pilcher, pilar santidri n tomillo, and laura sarti martnez 11 Leatherback Turtle Populations in the Indian Ocean 123 ronel nel, kartik shanker, and george hughes Part IV. FROM EGG TO ADULTHOOD 12 Leatherback Turtle Eggs and Nests, and Their Effects on Embryonic Development 135 paul r. sotherland, bryan p. wallace, and james r. spotila 13 Leatherback Turtle Physiological Ecology: Implications for Bioenergetics and Population Dynamics 149 bryan p. wallace and t. todd jones 14 Movements and Behavior of Adult and Juvenile Leatherback Turtles 162 george l. shillinger and helen bailey 15 Relation of Marine Primary Productivity to Leatherback Turtle Biology and Behavior 173 vincent s. saba, charles a. stock, and john p. dunne Part V. THE FUTURE OF THE LEATHERBACK TURTLE 16 Warming Climate: A New Threat to the Leatherback Turtle 185 james r. spotila, vincent s. saba, samir h. patel, and pilar santidrin tomillo 17 Impacts of Fisheries on the Leatherback Turtle 196 rebecca l. lewison, bryan p. wallace, and sara m. maxwell 18 Conclusion: Problems and Solutions 209 james r. spotila and pilar santidri n tomillo Index 213 Color plates follow pages 48 and 128. Contributors Helen Bailey Karen L. Eckert Chesapeake Biological Laboratory Wider Caribbean Sea Turtle Conservation Network University of Maryland Center for Environmental (WIDECAST) Science Ballwin, MO, USA Solomons, MD, USA Marc Girondot Barbara A. Bell Laboratoire Ecologie, Systematique and Evolution The Leatherback Trust Université Paris- Sud Monterey, CA, USA Paris, France Scott R. Benson George Hughes NOAA National Marine Fisheries Service 183 Amber Valley Southwest Fisheries Science Center Private Bag X30 Moss Landing Marine Laboratory Howick, South Africa Moss Landing, CA, USA T. Todd Jones Christopher A. Binckley JIMAR- UH- NOAA Department of Biology Kewalo Research Facility Arcadia University Honolulu, HI, USA Glenside, PA, USA Rebecca L. Lewison John P. Dunne Biology Department National Oceanic and Atmospheric Administration San Diego State University Office of Oceanic and Atmospheric Research San Diego, CA, USA Geophysical Fluid Dynamics Laboratory Princeton University Forrestal Campus Sara M. Maxwell Princeton, NJ, USA Department of Biological Sciences Old Dominion University Peter H. Dutton Norfolk, VA, USA NOAA National Marine Fisheries Service Southwest Fisheries Science Center Ronel Nel La Jolla, CA, USA Department of Zoology Nelson Mandela Metropolitan University Port Elizabeth, South Africa x Contributors Frank V. Paladino Kartik Shanker Biology Department Centre for Ecological Sciences Indiana- Purdue University at Fort Wayne Indian Institute of Science Fort Wayne, IN, USA Bangalore, India Samir H. Patel George L. Shillinger Coonamessett Farm Foundation The Leatherback Trust East Falmouth, MA, USA Monterey, CA, USA Nicolas Pilcher Paul R. Sotherland Marine Research Foundation Department of Biology Kota Kinabalu Kalamazoo College Sabah, Malaysia Kalamazoo, MI, USA Peter C. H. Pritchard James R. Spotila Chelonian Research Institute Department of Biodiversity, Earth and Oviedo, FL, USA Environmental Science Drexel University Nathan J. Robinson Philadelphia, PA, USA Biology Department Indiana- Purdue University at Fort Wayne Charles A. Stock Fort Wayne, IN, USA National Oceanic and Atmospheric Administration Office of Oceanic and Atmospheric Research David C. Rostal Geophysical Fluid Dynamics Laboratory Department of Biology Princeton University Forrestal Campus Georgia Southern University Princeton, NJ, USA Statesboro, GA, USA Jennifer Swiggs Vincent S. Saba Biology Department NOAA National Marine Fisheries Service Indiana- Purdue University at Fort Wayne Northeast Fisheries Science Center Fort Wayne, IN, USA Geophysical Fluid Dynamics Laboratory Princeton University Forrestal Campus Ricardo F. Tapilatu Princeton, NJ, USA Marine Laboratory The State University of Papua (UNIPA) Pilar Santidrián Tomillo Manokwari, Papua Barat Province, Indonesia Population Ecology Group Institut Mediterrani d’Estudis Avançats Bryan P. Wallace Esporles, Mallorca, Spain Stratus Consulting Boulder, CO, USA Laura Sarti Martínez Comisión Nacional de Áreas Naturales Protegidas Jeanette Wyneken SEMARNAT Department of Biological Sciences México DF, México Florida Atlantic University Boca Raton, FL, USA Preface with their massive size and strange appearance, leatherback turtles resemble species that inhabited Earth in the distant past. They are the sole extant members of the family Dermochelyidae, which includes at least eight other extinct species. Only leatherback turtles survive. These ancient reptiles have in recent decades introduced us to fascinating bi- ological aspects of one of the oceans’ most amazing creatures. Over the last 40 years, growing interest in the species has also resulted in an exponential increase in the number of scientific publications and in the number of conservation projects established by governments, nongov- ernmental organizations, academic institutions, and sometimes by local individuals and groups. The idea of producing a book came after several scientific discussions on a nesting beach in Costa Rica. At the time, there were already books available on the biology of sea turtles in general, sea turtles by region, as well as on particular species such as loggerhead and olive ridley turtles. We realized that there was a need to compile the latest and finest in- formation on leatherback turtles. Considering the state of knowledge, high interest, and uniqueness of the species, the need for a book became evident and the time for its production also seemed right. This book constitutes a thorough analysis of the biology and conser- vation of leatherback turtles. We have brought together world experts on the subject to build a book that can be used for academic purposes, but is also of general interest for herpetologists and people in other fields. Because leatherback turtles are both fascinating to scientists and an icon of ocean conservation, we especially wanted to make a strong connection between biology and conservation. As scientists, leatherback turtles have opened doors to the past for us. As conservationists, we need to open doors to the future for them. Many things have happened over the course of editing the book since we first presented the plan to John Hopkins University Press in 2010. Laurie Spotila was an inspiration from the very beginning, providing kindness, patience, and tolerance. Unfortunately, she passed away while the book was in press. Several family members and friends have also been especially supportive during the process. Some of them are no longer with us. The book would not have been possible without the help and support of xii Preface many people who kindly offered their services. We are Witherington for their contributions to improving the especially indebted to Vincent Burke for believing in the chapters.