Movement Patterns, Foraging Ecology and Digestive
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MOVEMENT PATTERNS, FORAGING ECOLOGY AND DIGESTIVE PHYSIOLOGY OF BLACKTIP REEF SHARKS, CARCHARHINUS MELANOPTERUS, AT PALMYRA ATOLL: A PREDATOR DOMINATED ECOSYSTEM A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERITY OF HAWAI‘I IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY DECEMBER 2008 By Yannis P. Papastamatiou Dissertation Committee: Kim Holland, Chairperson Jeff Drazen Steve Karl Christopher Womersley Tadashi Fukami UMI Number: 3349420 INFORMATION TO USERS The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleed-through, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. ______________________________________________________________ UMI Microform 3349420 Copyright 2009 by ProQuest LLC All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. _______________________________________________________________ ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, MI 48106-1346 We certify that we have read this dissertation and that, in our opinion, it is satisfactory in scope and quality as a dissertation for the degree of Doctor of Philosophy in Zoology. DISSERTATION COMMITTEE Chairperson ii ACKNOWLEDGMENTS I want to particularly thank my chair, Dr Kim Holland for taking me into his lab, and for teaching me some of the tricks of the trade. I would like to thank Dr’s Jeff Drazen, Steve Karl, Christopher Wormersley, and Tadashi Fukami for serving on my committee and always being available for advice when I needed it. I want to thank Dr Carl Meyer for providing me with a large number of opportunities (and employment) while a graduate student. I would also like to thank Dr Joanne Leong and Jane Ball for supporting me for a number of years and giving me the freedom to go to remote locations and do research. I would like to thank my colleagues Christopher Lowe, Alan Friedlander and Jenn Caselle, without whom I would have never completed the work at Palmyra. For helping me in the field in Palmyra I would like to thank T. Clark, N. Whitney, L. Davis, L. Max, M. Sheehy, and G. Goodmanlowe. For all experiments with captive sharks I would like to thank my army of volunteers E. Aus, A. Stankus, T. Tinhan, M. Burns, J. Coloma, E. Grau, C. Clarke, W. Connel, and J. Nakaya. For advice, discussions and arguments I would like to thank my lab mates N. Whitney, T. Clarke, D. Itano, J. Dale, T. Daly Engle, and M. Hutchinson. I would also like to acknowledge J. Dale and A. Taylor for statistical advice. L. Wedding and K. Anthony designed the maps of Palmyra. Finally I would like to thank Alexi and Caroline Papastamatiou, and Lori Davis for putting up with rather a lot from me, but still always being there when I needed it. Most importantly, I would like to dedicate this dissertation in the memory of Dimitri iii Papastamatiou for pushing me to do something I knew I wanted to do, but didn’t realize it at the time. Funding was provided by the National Geographic Society (grant # 7990-06), a UH Sea Grant Project Development fund, PADI Project Aware, and a grant from the Fish Aggregating Devices as Instrumented Observatories of Pelagic Life (FADIO) under DG Research of the European Commission. All experiments were approved by the University of Hawaii Animal Care Committee. iv ABSTRACT Apex predators may have a strong regulatory function in marine ecosystems through both density and behaviorally mediated effects. Understanding the ecological impacts of apex predators is particularly important in predator dominated ecosystems where intra-specific competition may be high. While a number of techniques are available for quantifying predator movement patterns and distribution, little is known of the causative factors that regulate these behaviors. One important aspect of predator behavior is foraging, and an important regulating aspect of foraging is digestion. To advance our understanding of the interrelationship between gastric function and foraging behavior, I tested two types of data loggers for deployment in shark stomachs. One type of tag measured stomach acidity, the other the motility of the stomach wall. Both types of tags were deployed in the stomachs of captive free-swimming blacktip reef sharks to determine the effects of feeding and fasting on gastric digestive function. Gastric pH was maintained low during long periods of fasting, suggesting continuous secretion of acid. Gastric motility was higher for meals of mackerel than for similar sized meals of squid with maximum motility occurring at meal sizes of 1 % body weight. Based on diel patterns of gastric motility and pH, I predict that blacktips will feed daily and preferably forage during times of low water temperature. Palmyra Atoll is a remote, predator dominated ecosystem, and has a large population of blacktip reef sharks. Blacktips at Palmyra are smaller than those at other locations, which may be the result of food-limited growth due to intra-specific competition. Palmyra consists of two lagoons (east and west), and abundance of sharks appears to be similar in both lagoons. Active and passive tracking was used to study the v movement patterns of the sharks at Palmyra. Sharks in the west lagoon utilized small home ranges over scales of days to weeks. Adult sharks selected ledge habitats, while smaller individuals selected sand-flats, and small pups were found in very shallow waters. Fractal analysis revealed that sharks used patches that were 3 – 17 % of the spatial scale of their home range, and that sharks move with a directed walk while in patches but move randomly between patches. Sharks in the west lagoon showed strong site fidelity with some individuals being detected there for over 3 years. Sharks showed little movements between lagoons and sharks in the east lagoon had shorter residence times. Sharks in the west lagoon had higher body condition indices than those in the east lagoon and stable isotope analysis revealed that trophic structure was different between the two lagoons. Conditions differ between the two lagoons which may be driving differences in foraging success. This study reveals the importance habitat can play in the movement patterns, home range and foraging success of sharks and suggests that intra-specific competition could be a strong regulator of apex predator populations in pristine predator dominated ecosystems. Studies of the digestive system revealed that physiology may also regulate some aspects of movement patterns, although field studies will be required to test these hypotheses. vi TABLE OF CONTENTS ACKNOWLEDGEMENTS................................................................................... iii ABSTRACT........................................................................................................... iv TABLE OF CONTENTS...................................................................................... vii LIST OF TABLES................................................................................................. ix LIST OF FIGURES ...............................................................................................10 CHAPTER I: Introduction .....................................................................................12 CHAPTER II: The response of gastric pH and motility to fasting and feeding in free swimming blacktip reef sharks, Carcharhinus melanopterus 21 ABSTRACT...............................................................................................21 INTRODUCTION .....................................................................................22 MATERIALS AND METHODS...............................................................24 RESULTS ..................................................................................................31 DISCUSSION............................................................................................44 CHAPTER III: A new acoustic pH transmitter for studying the feeding habits of free ranging sharks 53 ABSTRACT...............................................................................................53 INTRODUCTION .....................................................................................53 MATERIALS AND METHODS...............................................................55 RESULTS ..................................................................................................59 DISCUSSION............................................................................................59 CHAPTER IV: Scale dependent effects of habitat on movements and path structure of blacktip reef sharks, vii Carcharhinus melanopterus, at Palmyra Atoll: a predator dominated ecosystem 67 ABSTRACT...............................................................................................67 INTRODUCTION .....................................................................................68 MATERIALS AND METHODS...............................................................70 RESULTS ..................................................................................................81 DISCUSSION............................................................................................97