Origins, Movements, and Foraging Behavior of Hawksbill Sea
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ORIGINS, MOVEMENTS, AND FORAGING BEHAVIOR OF HAWKSBILL SEA TURTLES (ERETMOCHELYS IMBRICATA) IN PALM BEACH COUNTY WATERS, FLORIDA, USA by Lawrence D. Wood A Thesis Submitted to the Faculty of The Charles E Schmidt College of Science In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Florida Atlantic University Boca Raton, FL December 2014 Copyright by Lawrence D. Wood 2014 ii ACKNOWLEDGEMENTS The author wishes to thank the many volunteers, interns, dive boat operators, organizations, and funding sources that have made this study possible. Special thanks go to the management and staff of the Palm Beach Zoo and Conservation Society for so graciously hosting this project and providing the critical support needed for its continuation and completion. Critical funding was provided by the Sea Turtle Conservancy through the Florida Sea Turtle License Plate Grants Program, the National Save the Sea Turtle Foundation, the Bay and Paul Foundations, Mr. Robert Murtagh, Dr. Terry Maple, and others. The author is grateful for the technical support provided by Dr. Barbara Brunnick and Mr. Chris Johnson, and especially grateful to Dr. Terry Maple for his inspiration, advice, and encouragement before and while achieving this degree. iv ABSTRACT Author: Lawrence D. Wood Title: Origins, movements, and foraging behavior of hawksbill sea turtles (Eretmochelys imbricata) in Palm Beach County waters, Florida, USA Institution: Florida Atlantic University Thesis Advisor: Dr. Sarah Milton Degree: Doctor of Philosophy Year: 2014 This dissertation examined the natal origins, home-range, and in-situ foraging behavior of an aggregation of sub-adult hawksbill turtles (Eretmochelys imbricata) found off the coast of Palm Beach County, Florida. Surveys were conducted on approximately 30 linear km of reef between 15 and 30 m in depth. Tissue samples were retrieved from 112 turtles for mtDNA haplotype determination. GPS-linked satellite transmitters were deployed on six resident sub-adults, resulting in both minimum convex polygon (MCP) and 95%, 50%, and 25% kernel density estimates (KDE) of home-range size. A foraging ethogram was developed, and sequential analysis performed on thirty videos (141 total minutes) of in-situ foraging behavior. Seventeen total haplotypes were identified in this aggregation, the majority (75%) of which represented rookeries on Mexico’s Yucatan Peninsula. Other sources, from most to least important, include Barbados, Costa Rica, Puerto Rico, Antigua, and the U.S. Virgin Islands. Home range estimates ranged from v 1.1-19 km2 (mean 10.1 km2) using the MCP method, and 0.01-1.2 km2 (mean 0.49 km2) using the 95% KDE method. Consistent use of core areas within the home range, particularly at night, suggests the repeated use of familiar refuges for shelter. Five behaviors leading to prey ingestion were identified: scan, target, nudge, bite, and chew. Collectively, these behaviors occurred in decreasing frequency leading to prey ingestion, suggesting a highly discriminatory feeding strategy that focused on a narrow range of poriferan prey through extensive exploration of the benthic environment. These are the first and most detailed behavioral studies to date concerning hawksbill turtles in Florida, and have contributed important baseline information concerning the biogeography and natural history of this species in this part of its range. vi ORIGINS, MOVEMENTS, AND FORAGING BEHAVIOR OF HAWKSBILL TURTLES (ERETMOCHELYS IMBRICATA) IN PALM BEACH COUNTY WATERS, FLORIDA, USA TABLES ..................................................................................................................... viii FIGURES ........................................................................................................................ x INTRODUCTION .......................................................................................................... 1 CHAPTER 1: ORIGINS ................................................................................................ 3 Introduction ............................................................................................................... 3 Methods .................................................................................................................... 5 Results ..................................................................................................................... 10 Discussion ............................................................................................................... 12 CHAPTER 2: MOVEMENTS ..................................................................................... 18 Introduction ............................................................................................................. 18 Methods .................................................................................................................. 21 Results ..................................................................................................................... 29 Discussion ............................................................................................................... 50 CHAPTER 3: FORAGING BEHAVIOR .................................................................... 65 Introduction ............................................................................................................. 65 Methods .................................................................................................................. 68 Results ..................................................................................................................... 71 Discussion ............................................................................................................... 76 REFERENCES ............................................................................................................. 83 vii TABLES Table 1.1. Short (380-480 bp) and long (740 bp) haplotypes and their proportions for all and juvenile hawksbills only from three foraging grounds in the western Atlantic: Palm Beach (PB), Mona Island Puerto Rico, and Cayman Island. Data for Mona are from Vélez-Zuazo et al. (2008); data from Cayman Islands are from Blumenthal et al. (2009b). ..................................................... 10 Table 1. 2 Pairwise comparisons of haplotype frequencies (FST values) among three aggregations of hawksbill turtles from which 740 base-pair haplotypes are available: Palm Beach (PB), Mona Island Puerto Rico, and Cayman Islands. F-statistics from haplotype frequencies are shown below the diagonal, FST P-values above the diagonal.. ........................................................................... 11 Table 1.3. Maximum likelihood estimate for nesting beach origin of juvenile hawksbill turtles from Palm Beach County, Florida, based on 94 of 106 available sequences (12 juveniles had haplotypes that have not been reported from any nesting beach). Estimate is based on 384 bp of control region sequence and was generated using SPAM 3.7b (Alaska Department of Fish and Game, 2003). .................................................................................................... 11 Table 2.1. Search radii and cell size used for 95% kernel density estimates in Arcview 10.1® (ESRI, Inc.). ............................................................................................ 28 Table 2.2. Deployment dates, release locations, capture depths (m), substrate type, deployment duration (days) and straight carapace length (notch of the nuchal scute to the tip of either pygal scute, in cm) for six sampled hawksbills. ........................................................................................................ 29 Table 2.3. Total, diurnal (8:00 a.m. – 7:59 p.m. EST), and nocturnal (8:00 p.m. -7:59 a.m. EST) home range (HR) estimates for six hawksbill turtles. n= number of GPS coordinates; MCP= minimum convex polygon; KDE95= kernel density estimate, 95% contour. ......................................................................... 30 Table 2.4. Total, diurnal (8:00 a.m. – 7:59 p.m. EST), and nocturnal (8:00 p.m. -7:59 a.m. EST) core-use area estimates for six hawksbill turtles. MCP= minimum convex polygon; KDE50= kernel density estimate, 50% contour, KDE25= kernel density estimate, 25% contour. ............................................... 39 Table 2.5. Coordinates per three habitat zones for three turtles (#’s 21-23) residing on the Continental Reef Tract, and the area (extent) of each in km2..................... 41 viii Table 2.6. Proportionality of coordinates per habitat zone within and between individuals. W=west of the main ledge; L&C=ledge and reef crest; E= east of reef crest. Z-score and p-value are shown for each pair. Pairs that show no significant difference are shown in bold. (Fisher’s pooled 2-sample test for proportionality: Minitab 17™ statistical software) .................................... 49 Table 2.7. Estimated foraging home ranges of hawksbill turtles (Ei) in Florida and the Caribbean. MCP= Minimum Convex Polygon method; KDE= Kernel Density Estimate. .............................................................................................. 51 Table 3.1. Ethogram of hawksbill turtle foraging behavior. ............................................. 70 Table 3.2. Cumulative transition matrix. Numbers in cells represent the number of times a behavior on the y-axis transitioned to a behavior on the x-axis. For example, ‘scan’ transitioned to ‘target’ 247 times (*). ....................................