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Establishing Endocrine and Behavioral Parameters of Reproduction in Pacific Walrus (Odobenus Rosmarus Divergens)

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Establishing Endocrine and Behavioral Parameters of Reproduction in Pacific Walrus (Odobenus Rosmarus Divergens) Establishing Endocrine and Behavioral Parameters of Reproduction in Pacific Walrus (Odobenus rosmarus divergens) Lisa Triggs A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science University of Washington 2013 Committee: Glenn VanBlaricom Walt Dickhoff Karen Goodrowe Beck Kristin Laidre Program Authorized to Offer Degree: Aquatic and Fishery Sciences ©Copyright 2013 Lisa Triggs ii University of Washington Abstract Establishing Endocrine and Behavioral Parameters of Reproduction in Pacific Walrus (Odobenus rosmarus divergens) Lisa Triggs Chair of Supervisory Committee: Associate Professor Glenn R. VanBlaricom School of Aquatic and Fishery Sciences Reproduction of walrus (Odobenus rosmarus) in zoos and aquaria has met with limited success. While basic information on reproductive characteristics is available, there is limited knowledge of reproductive physiology and endocrinology for walrus. The overall goal of the research was to monitor reproductive events and seasonal changes that occur in ex situ walrus of North America by utilizing a variety of methods. To track changes in males, longitudinal hormonal analysis of saliva samples, changes in testicular size, presence or absence of spermatozoa and seasonal fluctuations in behavior were examined. Saliva samples were collected from six male walrus for periods of 12 to 36 months. Measurements of testicular length and width were collected on a weekly basis. Baseline testosterone values for each animal were established using an iterative process that eliminated elevated values. Elevations were defined as values ≥ 2 times the individual’s baseline (range 0.838-4.228 ng/ml). The mean combined baseline testicular measurement for individuals (R linear length + R linear width + L linear length + L linear width) was 54.81±2.13 cm while the mean combined increased testicular measurement (an increase of 15% or more over minimum values) was 65.38±1.71. Three of six animals displayed the typical iii rise and fall of testosterone concentrations observed in seasonal breeders in the wild and elevations were coincident with reproductive behaviors. Behaviors included visual and vocal courtship displays, pursuit of females, clasping and penetration. Sperm presence in a single walrus during two years coincided with increased testicular measurements and mean duration of spermatozoal presence was 131±3 days. Endocrine profiles varied extensively among animals, demonstrating within-animal decline in testosterone levels over time, a lack of testosterone elevations coincident with the wild walrus breeding season and a blind male with a unique derivation in his endocrine profile, hormone elevations directly opposite the wild walrus breeding season. To track female reproductive cycles, longitudinal hormonal analysis of saliva samples and seasonal fluctuations in behavior were examined. Saliva samples were collected from six female walrus at least three times a week for periods of 12 to 55 months. Salivary estradiol-17β was deemed an unreliable indicator of the follicular phase because hormone concentrations did not display a rise in estrogen concentrations prior to a rise in progesterone concentrations nor increased estrogen concentrations coincident with estrus behaviors. The most commonly reported signs of behavioral estrus were soliciting male attention, clasping and penetration. In general, behavioral observation also was not a consistently reliable method for detecting impending estrus. Salivary progesterone was found to be unsuitable for monitoring the functional capacity of the corpus luteum because profiles were difficult to interpret biologically or physiologically. Progesterone profiles did not appear to coincide with behavioral or seasonal changes. However, periods of elevated progesterone concentrations were demonstrated in two of six females and were most likely representative of consecutive pseudopregnancies (mean 204.27 iv days). An inconsistency elucidated in this study involved a blind female with a unique derivation in her endocrine profile, inconsistent with the wild walrus breeding season and varying in time frame each year. Issues with walrus reproduction elucidated by this study included: within-animal decline in testosterone levels over time, unsynchronized cycles between animals residing together and the inconsistent timing of reproductive events in blind animals. To increase breeding success of the ex situ walrus population the following changes are proposed: a) increase the number of walrus in the population or consolidate existing walrus into larger social groups; and b) separate the genders during the non-breeding season or move animals to different facilities. Facility collaboration and continued monitoring of reproductive functioning via endocrine, anatomical and behavioral parameters would allow evaluation of the efficacy of the proposed changes. v Acknowledgments Rarely does one have the chance to research in depth the species with which one has grown so enamored. I would like to thank Dr. Glenn VanBlaricom for allowing me that opportunity when he agreed to take me as a graduate student. Thank you for seeing the potential in me for graduate level work and accepting me into your lab. Thank you for your guidance, support and encouragement. I would like to thank Dr. Walt Dickhoff for donating his personal copies of Austin and Short’s first edition of Reproduction in Mammals to substitute for a missing UW reproduction class. Your knowledge, expertise and advice in endocrinology were much appreciated. I would like to thank Dr. Karen Goodrowe Beck for encouraging me to pursue a graduate level degree and always challenging me to be better. Your patience, time, invaluable guidance and dedication to my success both as a committee member and curator are greatly appreciated. I would like to thank Dr. Kristin Laidre for joining my committee at the last minute without hesitation. Your passion for marine mammals and research is truly an inspiration. Thank you to Dr. Holly Reed and Dr. Brian Joseph for encouraging me to pursue the sciences at a higher level. Thank you for being such great mentors, colleagues and friends. Your encouragement, support and belief in me mean more than you will ever know. A very special thank you to my husband for all of his patience, love and support (both financially and emotionally) over the last five years. You helped me maintain my sanity during these overwhelming times and stood by me when sanity was no longer possible. None of this would have been possible without you. I owe you big time! For my mother, thank you for always vi being there when I needed encouragement and helping me with all the extras. You both mean the world to me. A huge thank you to all of the walrus staff at Point Defiance Zoo & Aquarium. My zoo family was very supportive and committed to this project. A special thanks to my coworker Amanda Shaffer who participated in this project from the very beginning. Thanks for assisting my brain on days when it was filled to capacity (or catastrophe) with walrus research. I would like to thank all the staff at the many organizations, Six Flags Discovery Kingdom, New York Aquarium, Sea World San Diego and the Indianapolis Zoo, that assisted in this project. A special thanks to Lisa Oland, Alisa Keyes, Hans Walters, Nicole Pisciotta, Leah Coombs, Jessa Paschke, Holley Muraco, Mike Muraco, Mitzi Synnott and William Winhall for their extra attention and assistance in this project. Without your time and commitment, this collaborative research would not have been possible. Thank you to all the incredible walrus (ET, Basilla, Joan, Sivuqaq, Uquq, Siku, Kulusiq, Ayveq, Obie, Aurora, Nereus and Brutus) that graciously allowed us to probe their mouths in an effort to collect saliva and lay out to have their testicles measured. Your patience and participation are much appreciated. Thanks to the staff at the Brookfield Zoo. In 2006 Joan and Basilla arrived at PDZA as part of a collaborative breeding loan. This collaborative effort was what prompted this project. I am grateful to the staff at the Seattle Aquarium for the generous use of their laboratory facilities and their guidance in all things EIA. Thank you to Dr. Shawn Larson for her awesome research program. A special thank you to Angela Smith for her positive personality, expertise and vii guidance in the lab and her willingness to assist me in times of need (help the printer won’t work, help the machine is acting funny, etc.). You are a joy to work with! I would like to express my gratitude to Dr. Karen Wolf and Dr. Kadie Anderson for their editing assistance. To the veterinary staff at PDZA (including Jules), thank you for all your assistance in this project. To Kate Faust, thank you for assisting in the organization of thousands of saliva samples. To Jesse Blake, the “stats guru”, your statistical knowledge and willingness to help are greatly appreciated. A special thanks to members of the VanBlaricom Lab at the University of Washington. You have all provided me with helpful advice, support and laughter. You have made me feel part of a unique school family. Last but not least, thank-you to the Point Defiance Zoo & Aquarium Dr. Holly Reed Conservation Fund for their financial support. You made this project a reality. viii Table of Contents Abstract ...................................................................................................................................................... iii Acknowledgments.....................................................................................................................................
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