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Bone Growth Strategies and Skeletochronological Age Estimates of Desert Tortoise

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Bone Growth Strategies and Skeletochronological Age Estimates of Desert Tortoise Bone Growth Strategies and Skeletochronological Age Estimates of Desert Tortoise (Gopherus agassizii) Populations A Thesis Submitted to the Faculty of Drexel University by Amanda Jane Curtin in partial fulfillment of the requirements for the degree of Doctor of Philosophy May 2006 © Copyright 2006 Amanda J. Curtin. All rights reserved. ii Dedications For my grandparents, Peggy Lillian and David Croll Pollock and Crystal Pamela and Harold Desmond Curtin who always nurtured and encouraged my inquisitive nature and gave me a strong sense of my place in this world. In special remembrance of my grandmother, Peggy, who inspired my love of nature and also my grandfather, Harold, who inspired my love of knowledge and history....I would not have come this far without the determination and imagination that they fueled in me. iii Acknowledgements Special thanks to my advisor, Dr. Jim Spotila, for his friendship, support and encouragement. I’d also like to express my gratitude and appreciation to the other members of my committee: Dr Hal Avery (I could not have asked for a better teacher on desert ecology), Dr. Mike O’Connor, Dr. Peter Dodson and lastly Dr. George Zug, whose guidance, encouragement and friendship meant more than words can express. Thanks to the following people and institutions for donating carcasses used in this study: Dr. Philip Medica and the United States Geological Service (USGS), Dr. Roy Averill-Murray (Arizona Fish and Wildlife Service), Dr. William Boarman (USGS), the Desert Tortoise Conservation Center (Clark County, Nevada), and the Mojave National Preserve, California. To all the undergraduate and graduate students from Drexel University who helped with carcass collection and histological processing, I am most grateful. Also thanks to the Bioscience staff and faculty at Drexel for their support and encouragement, especially Dr. Laura Duwel, Dr. Jeremy Lee, Ms. Gerry Marekova and Ms. Brenda Jones-Bowden. Lastly to my friends and family, especially my mother, Louise, and step- father, George Loudon - without your love, strength and encouragement I would not have been able to achieve this dream. This research was funded by the Panaphil Foundation and Betz Chair Endowment. iv Table of Contents LIST OF TABLES……………………………………………………………………. viii LIST OF FIGURES………………………………………………………………….. xi ABSTRACT………………………………………………………………………….. xvi 1. GENERAL INTRODUCTION…………………………………………………... 1 The desert tortoise (Gopherus agassizii)..……………………………….……… 2 Skeletochronology….…….…………………………………………..….......... 4 General methodology…………………….………………….……………....... 6 Ranking protocol…….................................................................................. 7 Correction factor………..…………………………………….……….….. 9 General problems and considerations………………………………………… 9 2. CHAPTER 2. KNOWN-AGED DESERT TORTOISES (GOPHERUS AGASSIZII) FROM ROCK VALLEY, NEVADA VALIDATE SKELETOCHRONOLOGICAL ANALYSIS...…………………………………………………………………….. 13 Introduction…………………………………………………………………... 13 Materials and methods……………………………………………………….. 15 Skeletochronology....................................................................................... 17 Results............................................................................................................... 18 Age and longevity........................................................................................ 18 Age at sexual maturity................................................................................. 19 v Discussion............................................................................................................. 19 Age and longevity........................................................................................ 19 Age at sexual maturity..…………………………………………………... 21 Conclusions and implications for future research……....……………….……… 23 3. AGE AND GROWTH STRATEGIES OF SONORAN DESERT AND WEST MOJAVE DESERT TORTOISES (GOPHERUS AGASSIZII)............................... 40 Introduction........................................................................................................ 40 West Mojave Desert Tortoises..................................................................... 43 Sonoran Desert Tortoises............................................................................. 44 Materials and methods....................................................................................... 46 Materials...................................................................................................... 46 Skeletochronology....................................................................................... 46 Statistical analysis........................................................................................ 47 Results................................................................................................................ 48 Body size...................................................................................................... 48 Bone morphometry............................................................................................ 49 Resorption core diameters............................................................................ 51 Age and longevity........................................................................................ 52 Age at sexual maturity................................................................................. 54 Discussion.......................................................................................................... 55 Body size and bone morphometry............................................................... 55 Resorption core diameters........................................................................... 59 Longevity and age at maturity..................................................................... 62 Growth strategies, ecological implications and conclusions............................. 63 vi 4. AGE ESTIMATES AND GROWTH STRATEGIES IN EASTERN MOJAVE DESERT TORTOISE (GOPHERUS AGASSIZII) POPULATIONS...................... 101 Introduction........................................................................................................ 101 Materials and methods....................................................................................... 102 Results................................................................................................................ 104 Body size...................................................................................................... 104 Bone morphometry....................................................................................... 105 Resorption core diameters............................................................................ 105 Age and longevity......................................................................................... 106 Age at sexual maturity.................................................................................. 107 Discussion........................................................................................................... 107 Body size and bone morphometry................................................................ 107 Growth strategies, ecological implications and conclusions.............................. 109 5. CHAPTER 5. GROWTH STRATEGIES IN DESERT TORTOISES POPULATIONS: ECOLOGICAL IMPLICATIONS AND FUTURE TRENDS................................................................................................................. 142 Introduction........................................................................................................ 142 Materials and methods....................................................................................... 146 Results................................................................................................................ 147 Discussion.......................................................................................................... 150 Size, growth and productivity...................................................................... 150 Ecological implications and conclusions..................................................... 155 BIBLIOGRAPHY.......................................................................................................... 167 VITA.............................................................................................................................. 180 vii LIST OF TABLES 1. Shell measurements for Gopherus agassizii carcasses from the East Mojave Desert, Nevada……………………………………………………………………………… 25 2. Bone measurements for Gopherus agassizii carcasses from the East Mojave Desert, Nevada; CL = carapace length, H = humerus, F = femur, I = ilium, S = scapula, J = juvenile, F = female, M = male…………………………………………….……… 27 3. Periosteum diameter (PD) and growth layer (GL) ranking protocol and Correction Factor (CF) age estimates for Gopherus agassizii from the East Mojave Desert (Nevada); J = juvenile, F = female, M = male…………………………………….. 29 4. Maximum age estimates for Gopherus agassizii from the East Mojave Desert (Nevada) using the periosteum diameter (PD) and growth layer (GL) ranking protocol and correction factor (CF) skeletochronology methods. Age estimates in bold are equal or closest to the known age (where available). The value in brackets is the difference between the age estimate and known age; CL = carapace length……… 32 5. Student’s t-test analysis and mean difference (+ one standard error [SE] of the mean) between known age versus skeletochronology age estimates in Gopherus
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