UNIVERSITY OF CALIFORNIA, SAN DIEGO Hormones and blubber: An endocrinological approach for assessing life-history states in free-ranging cetacean populations A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy in Marine Biology by Nicholas M. Kellar Committee in charge: Professor William F. Perrin, Co-Chair Professor Ronald S. Burton, Co-Chair Professor Gerald L. Kooyman Professor Pamela L. Mellon Professor Greg W. Rouse 2008 3319846 Copyright 2008 by Kellar, Nicholas M. All rights reserved 3319846 2008 Copyright Nicholas M. Kellar All rights reserved The dissertation of Nicholas M. Kellar is approved, and it is acceptable in quality and form for publication on microfilm: Co-Chair Co-Chair University of California, San Diego 2008 iii DEDICATION My dissertation is dedicated to Dr. Susan Stewart Suchocki. Leading by example, she taught me the utility of the scientific method, the strength of perseverance, and the power of compassion for all. She cultivated my love for nature, the pursuit of knowledge, and pitcher dominated baseball. She has been my motivation from the start of it all. Mom, you will be missed…always. iv TABLE OF CONTENTS Signature page ……………………………………………………... iii Dedication…………………………………………………………. iv Table of contents………………………………………………….. v List of figures………………………………………………………. vi List of tables……………………………………………………… x Acknowledgements………………………………………………... xi Vita………………………………………………………………… xvi Abstract……………………………………………………………. xvii I. General summary………………………………………………….. 1 II. Chapter one: Growth and development of the Stenella attenuata corpus luteum: identifying potential characters to differentiate CLs of ovulation from those of pregnancy…… 28 III. Chapter two: Determining Pregnancy from blubber in three species of Delphinids………………………………………. 59 IV. Chapter three: Blubber testosterone: A marker of male reproductive status in short-beaked common dolphins…….. 93 V. Chapter four: Pregnancy patterns of pantropical spotted dolphins (Stenella attenuata) in the eastern tropical Pacific determined from hormonal analysis of biopsies and correlations of the patterns with the purse-seine tuna fishery………………………………………………… 136 v LIST OF FIGURES Figure 1. Computer image analysis of luteinized CL tissue……………….… 35 Figure 2. Computer image analysis of vascularized CL tissue………………. 36 Figure 3. Frequency distribution of spotted dolphin CL volume with and without observed fetuses……………………………………. 39 Figure 4. CL volume as a function of fetal length during early gestation …… 40 Figure 5. Luteinized tissue as a function of fetal length……………………… 42 Figure 6. Fraction of vascularized tissue as a function of fetal length……… 43 Figure 7. Relationship between fraction of lutienized and vascularized tissue of CL observed with and without a fetus………………………. 46 Figure 8. Blubber progesterone concentrations as a function of body length in pregnant and non-pregnant female common dolphins…… 75 Figure 9. Maternal blubber progesterone concentrations as a function of fetal length of 15 common dolphins..……………….. 76 Figure 10. Progesteorne concentrations in the outer, middle, and inner blubber layers of pregnant common dolphins……………… 78 Figure 11. Blubber progesterone concentrations at seven anatomical locations in a recently pregnant northern right-whale dolphin….. … 79 Figure 12. Geogrpahic distribution of blubber samples from fishery- killed and biopsied males common dolphins……………………….. 100 Figure 13. Compund light miscopy images showing histological vi characteristics of testes development in male shorted-beaked common dolphins…………………………………………………… 102 Figure 14. Completion curves showing parallel displacement between known standard concentrations and serial dilutions of pooled extracts of unknown concentration…………………………. 110 Figure 15. Blubber testosterone concentrations as a function of ordinal date for immature, pubertal, and sexually mature short-beaked common dolphins…………………………………………………… 113 Figure 16. Blubber testosterone concentrations as a function of right testis mass…………………………………………………………… 114 Figure 17. Testosterone concentrations of the outer, middle, and inner blubber layers in ten mature male common dolphins ………. 116 Figure 18. Blubber testosterone concentrations at nine anatomical locations in three mature male common dolphins…………..….. … 117 Figure 19. Logistic model for the p(mature) relative to blubber testosterone concentration…………………………………………. 119 Figure 20. Logistic model for the p(mature) relative to blubber testosterone concentration and ordinal date……………………….. 120 Figure 21. Blubber testosterone concentration as a function of ordinal date in biopsied male short-beaked common dolphins in the Northeastern Pacific………………………………………………… 123 Figure 22. Mean right testis mass in reference specimen as a function vii of date……………………………………………………………….. 126 Figure 23. Geographic distribution of female pantropical spotted dolphin biopsy samples that were analyzed to determine progesterone concentration………………………………………… 143 Figure 24. Frequency distribution of blubber progesterone concentrations in biopsies females pantropical spotted dolphins…………………………………………………………… 149 Figure 25. Results of randomized nearest neighbor distance analysis……. 155 Figure 26 Ripley’s K analysis of 24 biopsied pregnant pantropical spotted dolphins relative to all females sampled………………... 156 Figure 27. Prediction map from ordinary Kriging interpolation of the proportion pregnant for biopsies of female pantropical spotted dolphins…………………………………………………… 157 Figure 28. Geographic distribution of the female spotted dolphin biopsies designated as pregnant and non-pregnant as determined from blubber progesterone concentrations……………………....... 159 Figure 29. Spatial distribution of pregnant and non-pregnant females pantropical spotted dolphins in the observed historic fishery kill….. 160 Figure 30. Frequency distribution of simulated (permuted) difference of means statistic for Archer’s fishery exposure index (140-nmi ambit) of biopsied pregnant and non-pregnant female S. attenuata. ………………… 163 Figure 31. Prediction map from an ordinary Kringing interpolation of viii Archer’s fishery exposure index (140-nmi ambit) for biopsied female pantropical spotted dolphins………………………………… 164 ix
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