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Hormonal and Morphological Aspects of Growth and Sexual Maturation in Wild-Caught Male Vervet Monkeys (Chlorocebus Aethiops Pygerythrus) Tegan J

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Hormonal and Morphological Aspects of Growth and Sexual Maturation in Wild-Caught Male Vervet Monkeys (Chlorocebus Aethiops Pygerythrus) Tegan J University of Wisconsin Milwaukee UWM Digital Commons Theses and Dissertations December 2012 Hormonal and Morphological Aspects of Growth and Sexual Maturation in Wild-Caught Male Vervet Monkeys (Chlorocebus aethiops pygerythrus) Tegan J. Gaetano University of Wisconsin-Milwaukee Follow this and additional works at: https://dc.uwm.edu/etd Part of the Animal Sciences Commons, Developmental Biology Commons, and the Endocrinology Commons Recommended Citation Gaetano, Tegan J., "Hormonal and Morphological Aspects of Growth and Sexual Maturation in Wild-Caught Male Vervet Monkeys (Chlorocebus aethiops pygerythrus)" (2012). Theses and Dissertations. 30. https://dc.uwm.edu/etd/30 This Thesis is brought to you for free and open access by UWM Digital Commons. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of UWM Digital Commons. For more information, please contact [email protected]. HORMONAL AND MORPHOLOGICAL ASPECTS OF GROWTH AND SEXUAL MATURATION IN WILD-CAUGHT MALE VERVET MONKEYS (CHLOROCEBUS AETHIOPS PYGERYTHRUS) by Tegan J. Gaetano A Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science in Anthropology at The University of Wisconsin at Milwaukee December 2012 ABSTRACT ENDOCRINE AND MORPHOLOGICAL ASPECTS OF GROWTH AND SEXUAL MATURATION IN WILD-CAUGHT MALE VERVET MONKEYS (CHLOROCEBUS AETHIOPS PYGERYTHRUS) by Tegan J. Gaetano The University of Wisconsin at Milwaukee, 2012 Under the Supervision of Dr. Trudy R. Turner Knowledge of baseline changes in testosterone (T) and other androgens is central to both investigations of morphological, physiological, and behavioral correlates of inter-individual variation in the timing and shape of key events and transitions over the life course and questions of the evolution of species-specific schedules of maturation in primates. T represents an important determinant of spermatogenesis in male mammals and plays a central role in the expression of male sexual behavior and the development of secondary sex characteristics. This research integrates hormonal and morphometric methods to determine age-related changes in fecal testosterone (fT) metabolites and morphological markers of sexual maturation, including testicular volume, body mass (measured as BMI), and canine length, over the life course in a cross-sectional sample (n = 56) of wild-caught South African male vervet monkeys (Chlorocebus aethiops pygerythrus ). Contrary to expectations of this study, T was not significantly associated with age ( p = 0.1316 by ANOVA). However, BMI (p = 0.00022) and testis volume ( p = 4.335e-06) were strongly related to age, corresponding to the eruption of the canine teeth at adolescence (3-4 years of age). These results iii strongly suggest the existence of an adolescent growth spurt in male vervets. An interpretation of activation of the development of these maturational markers in preparation for challenges encountered at reproductive maturity, including dispersal and reproduction, is tentatively adopted here following Jolly and Phillips-Conroy (2003, 2006). However, future longitudinal observations to determine changes within individuals are necessary to provide greater confidence in this interpretation. iv ACKNOWLEDGEMENTS I would like to first thank the Department of Anthropology at the University of Wisconsin at Milwaukee (UWM) for allowing me the opportunity to pursue this research. I am grateful to so many people, but primary acknowledgement goes to my advisor, Dr. Trudy Turner, whose careful guidance and support throughout the research and writing processes were instrumental to the completion of this work. Acknowledgement also goes to the other members of my committee, Dr. Rafael Rodriguez and Dr. Benjamin Campbell, whose comments greatly improved an earlier draft of the manuscript. Special acknowledgement goes to Dr. Jennifer Danzy Cramer, who coached me every step of the way, Dr. Christopher Schmitt, who provided invaluable support in the field, Krista Millich, who did the legwork processing the samples, Dr. J. Patrick Gray, for his statistical counseling, and my parents, Kurt and Julieann Gaetano, for their unwavering love and support. I also acknowledge the many individuals who contributed in some way towards this project, from collecting samples in the field to providing emotional support at home: Elzet van Aswegen, Lindsay Barone, Micah Beller, Maryjka Blaszczyk, Riël Coetzer, Carol Daniel, Richard Gaetano, Dr. J. Paul Grobler, Michelle Heidelbach, Matt Kramer, Kathy Krueger, Hanna Legator, Dr. Lizanne Meiring, Oliver “Pez” Morton, Brian Nichols, Dewald du Plessis, Stefani Quam, Dr. Murray Stokoe, Nicholas Theron, the librarians at the UWM Golda Meir library, and everyone at the University of California at Los Angeles (UCLA) Center for Neurobehavioral Genetics. v Generous funding for this research was provided in part by a grant from the National Institute of Health (NIH R01RR016300). Gratitude is also due to the government of South Africa for permission to conduct this research and everyone at the South African National Department of Water and Environmental Affairs, the Free State, Kwa-Zulu Natal, Limpopo, and Eastern Cape provincial permit offices, and DHL. Lastly, I would like to thank the University of the Free State genetics department and the University of Illinois at Urbana-Champaign anthropology department for the use of their lab space and the UWM and UCLA IACUC committees. vi TABLE OF CONTENTS I. INTRODUCTION………………………………………………... .. 1 II. BACKGROUND………………………………………………… .. 5 A. Vervet Monkey Phylogeny……………………………….. 5 B. Vervet Monkey Behavior and Ecology…………………… 11 i. Description ………………………………………..... 11 ii. Diet and Distribution ……………………………… 15 Geographic Distribution and Habitat ………..... .. 15 Diet ………………………………………………. 18 iii. Social Organization ………………………………. 20 Group Composition, Ranging, and Territoriality .. 20 Dispersal ………………………………………… 24 Intragroup Relationships ………………………… 27 iv. Life History ………………………………………... 30 C. Life History Theory……………………………………….. 35 i. Comparative Primate Life Histories ……………….. 37 ii. Male Life History Strategies ………………………. 41 iii. Life History Variables in Ontogeny ………………. 46 D. Hormones and Life History………………………………. 50 i. Testosterone and the Endocrine System …………… 50 ii. Testosterone and Male Life Histories ……………... 53 iii. Testosterone Secretion over the Life Course ……... 56 III. OBJECTIVES/HYPOTHESES…………………………………. 63 IV. METHODS………………………………………………………. 65 A. Study Sites and Populations………………………………... 66 B. Sampling Procedures………………..……………………… 68 C. Data Collection and Analysis………………………………. 70 i. Age Classifications …………………………………… 70 ii. Morphological Measurements…………………………... 72 iii. Hormonal Analysis ……………………………………… 76 vi. Data Analysis ……………………………………………. 78 V. RESULTS…………………………………………………………... 79 VI. DISCUSSION……………………………………………………... 93 VII. CONCLUSION…………………………………………………... 98 VIII. LITERATURE CITED…………………………………………... 100 vii LIST OF FIGURES Figure 1. Morphological traits of the Old World monkey subfamilies….. 7 Figure 2. Molecular phylogeny of the parvorder catarrhini……………… 11 Figure 3. Map of Africa with distribution of Ch. aethiops ………………. 17 Figure 4. Life history constants of primates and other mammals……….. 41 Figure 5. Primate life history model…………………………………….. 50 Figure 6. Map of South Africa with GPS coordinates of trapping sites... 67 Figure 7. Dental patterns of catarrhini…………………………………... 73 Figure 8. Description of morphological measurements………………..... 74 Figure 9. Formula used for calculating BMI…………………………..... 76 Figure 10. Box and beanplots of linear regressions for Schema 1…….... 85 Figure 11. Box and beanplots of linear regressions for Schema 2…….... 87 Figure 12. Box and beanplots of linear regressions for Schema 3…..….. 90 Figure 13. Scatterplots of relationships among the dependent variables... 92 viii LIST OF TABLES Table 1. Summary of pelage color variation in Ch. aethiops …………… 12 Table 2. Summary of vervet group size, home range size, and diet……. 21 Table 3. Summary of the relationship between T and SSCs……………. 57 Table 4. Timing of acquisition of SSCs for male vervets………………. 63 Table 5. Description of trapping sites…………………………………... 68 Table 6. Age categories defined according to dental eruption………….. 73 Table 7. Age breakdown of sample according to classification schema... 75 Table 8. Summary of descriptive statistics…………………………….... 80 Table 9. Summary of ANOVA for log fT and log BMI……………….... 81 Table 10. Summary of ANOVA test for pair-wise comparisons for log fT and log BMI…………………………………………………….... 82 Table 11. Summary of Kruskal-Wallis test for relative canine length and relative testis volume……………………………………………... 82 Table 12. Summary of post Kruskal-Wallis test for multiple comparisons for relative canine length and relative testis volume……………... 8 ix 1 I. INTRODUCTION The concept of life history trade-offs has been instrumental to explanations of observed variation in the timing and shape of key events, transitions, and phases in the life course by providing a framework that considers all aspects of an organism. Life history theory describes differential strategies of allocation of resources among components of fitness (e.g. energy invested in survival versus reproduction) through the organizing principle of trade-offs. Trade-offs refers to the negative relationship between life history components and can be imagined as a series of facultative decisions as regards the allocation of energy to one aspect of the life course over another.
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