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Predation Risk and Molting Decisions in the Hawaiian Crab Leptodius Sanguineus: the Ups and Downs of an Armored Exoskeleton

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Predation Risk and Molting Decisions in the Hawaiian Crab Leptodius Sanguineus: the Ups and Downs of an Armored Exoskeleton PREDATION RISK AND MOLTING DECISIONS IN THE HAWAIIAN CRAB LEPTODIUS SANGUINEUS: THE UPS AND DOWNS OF AN ARMORED EXOSKELETON Hkliine Harvey B.Sc., Universitb de Montrkal, 1988 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in the Department 0 f Biological Sciences O Hklene Harvey 1993 SIMON FRASER UNIVERSITY January 1994 All rights reserved. This work may not be reproduced in whole or in part, by photocopy or other means, without permission of the author. APPROVAL Name: Degree: Master of Science Title of Thesis: PREDATION RISK AND MOLTING DECISIONS IN THE HA WAIL4N CRAB LEPTODIUS SANGUINEUS: THE UPS AND DOWNS OF AN ARMORED EXOSKELETON Examining Committee: Chair: Dr. Alton S. Harestad, Associate Professor - - Dr. Lawrence M. ~ill,/krofe~sor,Senior Supervisor, Department of Biological Sciences, SFU I.-I Dr. Ron C. mrg,Associate Professor, Department of Biological Sciences, SFU Dr. A. ~ichard~alrher, Professor - Department ojBiology, University of Alberta .. -. - Dr. ~obe'ii'w'~1nLr. Head, Ecosystem Research, Canadian Wildlife Service Public Examiner Date Approved 6 3-q 149 G PARTIAL COPYRIGHT LICENSE I hereby grant to Simon Fraser University the right to lend my thesis, project or extended essay (the title of which is shown below) to users of the Simon Fraser University Library, and to make partial or single copies only for such users or in response to a request from the library of any other university, or other educational institution, on its own behalf or for one of its users. I further agree that permission for multiple copying of this work for scholarly purposes may be granted by me or the Dean of Graduate Studies. It is understood that copying or publication of this work for financial gain shall not be allowed without my written permission. Title of Thesis/Project/Extended Essay (name) 4 Abstract The exoskeleton of crustaceans serves many functions, including defensive armor against predators. Since the exoskeleton is softer just before, during and just after molt, this period of transition increases prey vulnerability to predators. Given this increased vulnerability, behavioral responses to minimize risk are expected. The cost of molting should also influence life-history decisions, especially when to molt and how much to grow; delaying molt when the risk of predation is high would seem an appropriate defensive adaptation. Morphological defences sensitive to risk levels could be induced during the animal's lifetime as well. Relatively few studies have addressed the trade-offs between defences and growth. In this thesis, I quantify some aspects of the vulnerability of the xanthid crab Leptodius sanguineus to a portunid crab predator (Thalamita crenata) and investigate how behavioral and morphological defences vary throughout the molt cycle. Prey size had a large impact on vulnerability of intermolt crabs. Below a predator/prey size ratio of 2.8, intermolt crabs resisted the predator from 1 to 105 min. For freshly molted crabs, this size ratio was irrelevant: most crabs were killed in less than one minute. This increased vulnerability may be reduced somewhat by changes in behavior. For example, to avoid predator detection, freshly molted Leptodius sanguineus emphasized crypsis by reducing movement in the presence of a predator: they buried into the sand more slowly and made more pauses compared to intermolt crabs. Large male crabs, when exposed to predation risk, were also more likely to molt under cover under. With increases in predation risk (detected chemically), crabs delayed their molt. Differences in foraging behavior cannot explain these results. In most cases, size increment was unaffected: crabs did not compensate for a delayed molt by a larger size increment at their next molt. Although crabs did not produce relatively larger claws under predation risk, carapace toughness did appear to increase. The carapace toughens during the 20 days following a molt. The anterior region gets harder than the posterior region, but the rate of hardening of the posterior region may be increased under predation risk. Crabs that delayed molt under the highest level of predation risk produced a tougher carapace in the anterior region than did control crabs. This study provides direct evidence that molting is plastic. Crabs may thus modify their molt cycle and exoskeletal strength in response to variable predation risk. To Patrick A Camille, Yvette Philippe et Simon Acknowledgments I landed in Vancouver on the 8th of January 1990. Within that first week at SFU, I was taught what graduate school is all about: hacker hockey. I wish long life to this practice and thank everyone who participated over the years. Hacker hockey symbolizes the spirit of the Behavioral Ecology Research Group, people with many different interests always ready to welcome newcomers. I am glad to have been part of this group, whose members introduced me to the many cultural and scientific events of the far west: concerts, good parties and dance, pot lucks, eagle counting, interesting research, lab meetings, Les ~colo~istesand a series of great conferences. A hand of applause for all past and present members. I wish to thank my supervisor, Larry Dill, for taking me on as his student and for his patience and wonderful editorial work through reading hundreds and hundreds of pages of text and graphs! I also want to thank my committee members, Rich Palmer and Ron Ydenberg for the best meetings in the Vancouver Airport and diverse cafeterias. I much appreciated their constructive comments. NSERC, Simon Fraser University, University of Hawaii, The American Museum of Natural History, Larry Dill and the Dillery poker nights have supported my work financially. Thank you. My work with crabs started at the Bamfield Marine Station. I would like to thank all the personnel for their help and Jennifer Boldt for her generous assistance. Thanks also to Dan Soluk and Liz Boulding for sharing their interest and ideas about crabs. I am also grateful to Philip Helfrich, Ernst Reese and George Losey for sponsoring my two summers of work in Hawaii. The staff and the graduate students of the Hawaii Institute of Marine Biology responded to my pressing needs for space, material, information, and the "out of hours" boat rides between islands. Special thanks are due to Bob, Wayne, Cindy, Fanny, Bill and John. Bravo to the profs and students who participated in the 1990 HIMB summer program, for memorable lectures, scientific exchange, snorkling, diving, beaching, eating, boating, fishing, clubbing and sightseeing. Special thanks to Ernie who helped me discover Gravel Island, and big thanks to Thalamita crenata and Leptodius sanguineus for their much appreciated cooperation. I thank Beatrice Burch, Colin Lau and Jim Perrish who provided tips and information for xanthid identification. On Coconut Island, Ray was always ready, with a smile, to give me a hand with whatever crazy things I needed: finding a piece of hose or pipe, filing rods, melting lead or instructions on the use of big and small tools. It was always good to talk to Lester, a happy character whose friendship I treasured. A seasoned fisherman, Lester taught me all about World War 11, how to catch my predators, and loaned me his equipment. In addition, he took care of my palate: crabs, lobsters, banana bread and chocolate. In 1991, Maria Abate was also back for research on Coconut Island; we shared her fascination with the cleaner wrasse system, and the ups and downs of working with the "Wrassehole". Also in 1991, I found a home at The Mariculture Research and Training Center. My stay there was so refreshing. Working on the farm was a good change from scrubbing buckets, at least a change of tank size. I really enjoyed my experience and peace there and thank everyone, especially Chris Brown and BO Alexander for their hospitality, for a scenic room in mango heaven (or hell), and fish for my crabs. Back in B.C., I thank Franqois Bellavance for his statistical assistance, philosophy and teaching. I thank Alex for always having the answer and the time. I also owe thanks to my lab mates for not challenging my despotic ruling of the computer. Thanks to Darren for bike rides, souvlakis, precious help and encouragement. Thanks to Lourdes for many laughs and tears. Thanks to Karen for sharing the sensuality of sushi, and Creston. Thanks to Dave for driving all the way to Seattle on our way to Anacortes. Thanks to Karen and Dave for good feedback on Chapter 2, and for frequent hiking, skiing, dancing, and being the last ones to leave at parties. I I want to thank Camille, Yvette and Sophie for their support despite the distance, and for their patience and understanding of my absence. I also want to thank my parents, Jim, Louise, Julie and Jack for visiting and sharing wonderful times together in this beautiful part of the continent. Most of all, thank you Patrick for your help, your support, and your affection every step of I 1 the way; your love calmed every storm. vii Table of Contents Approval..................................................................................................................................... i i ... Abstract....................................................................................................................................... 111 Dedication................................................................................................................................ v Acknowledgments ...................................................................................................................v
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