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Jessica A. Mayhew Phd Thesis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ttention cues in apes and their role in social play behavior of western lowland gorillas (gorilla gorilla gorilla) Jessica A. Mayhew This thesis is submitted in partial fulfillment for the degree of PhD at the University of St Andrews December 3, 2012 Acknowledgements This dissertation would not have been possible without the guidance and support of many people on both sides of the Atlantic. I would like to thank my colleagues at St Andrews, particularly those in the Wolfson lab, for thoughtful discussions and sharing in both the triumphs and difficulties of the PhD process. Many thanks go to Andy Whiten and Klaus Zuberbühler for their helpful comments and critiques on this project during its initial stages. A big thank you also goes to Professor Richard Byrne and Dr. David Leavens for a positive and inspiring viva and the many insightful comments and queries about this work. Thanks to Ken Munro for the development of the “Gape” program, and to the participants in the experimental Bogart studies. Thank you to David Mackey at Plymouth State University for taking the time to answer questions about statistics. Thank you to Kirsten Pullen from the Paignton Zoo and Julie Mansell from the Belfast Zoo for contributing their gorilla groups to the white sclera study. A tremendous thank you goes to Dr. Juan-Carlos Gómez for his advice, for reading through countless manuscript revisions, and most of all, for his incredible patience throughout this entire process. I am grateful to Dr. Dario Maestripieri for his encouragement and for mentoring me in my time while at The University of Chicago. His dedication to and enthusiasm for research motivated me to pursue this PhD. A big thank you goes to Jason Watters for welcoming me to the Brookfield Zoo and introducing me to each member of their wonderful gorilla group. It was an absolute pleasure to spend my summer there. A thank you also goes to Craig Demitros for taking the time to talk gorilla with me. III My time in Chicago would not have been possible without the generosity of Jen for sharing her living room and refrigerator with me. I am eternally grateful for her friendship and love. Thank you Cass, for teaching me the significance and beauty of friendship, and to Arbi for his continuous excitement about having a monkey scientist for a friend. To Chad, mi media naranja, it has been a long journey, and I thank you for always meeting me at the airport. Thank you Charlie and Lu for being my home away from home and for being the most generous and caring people I’ve ever had the pleasure to know. Thank you to Debbie for her never-ending support and always kind and loving words, and to my Grandma for her reminders of how proud she is of me. To my parents, who have provided me with the opportunity to pursue academics without question and with the utmost support. There are not enough ways to thank you for always being excited and encouraging of my nerdy babbling. To Bry and Critter, your constant questions about primates and suggestions for new and unconventional avenues of research have always been appreciated. Thank you for dinnertime discussions that are nothing short of brilliant. And finally, to all of the animals in my life, especially Schaefer and Hails (“Spooky”), who have inspired me to keep looking for a way to get to know them better. IV Abstract Attention cues in apes and their role in social play behavior of western lowland gorillas (Gorilla gorilla gorilla) The research aims of this thesis are to investigate the attention cues available to and used by apes, especially gorillas (Gorilla gorilla), to ascertain the direction of conspecific attention during social interactions with a special reference to social play. Minimal research has been conducted on the role of attention cues – eye gaze, head, and body orientation – to regulate natural social interactions, such as social play, in non-human primates. This thesis begins with an investigation of the “cooperative eye hypothesis”, which poses that humans have evolved a unique white sclera adaptation for advertising and detecting gaze direction. Chapter 2 reports the existence of a natural white sclera variation in a proportion of gorilla eyes – contradicting the widely held assumption that white sclera is an exclusively human characteristic – and analyzes the presence of white sclera in relation to other morphological changes in the human eye. The study concludes that the morphological elongation of the eye might be a more important and unique change than the white sclera coloration. Chapter 3 experimentally explores the contribution of white sclera in both great ape and human eye gaze to the perception of gaze direction detection by human observers. This chapter concludes that although white sclera contributes to the accuracy and speed of gaze direction detection (an assumption that this thesis has put to experimental test for the first time), this merely adds to the already efficient gaze cues available in the eye areas of the ape face. Chapter 4 investigates the role of eye gaze, head, and body orientations during gorilla social play behavior, and more specifically, introduces a novel analysis of “vigilance periods” (VPs), V in which gorillas may use the interaction between attention cues to gauge the attention and intentions of play partners to successfully navigate play. The final study (Chapter 5) complements Chapter 4 and investigates the role of gorilla postures, behaviors, and movements during changes in attentional cue orientations. This chapter concludes that gorillas often engage in physical rest during VPs but maintain attentional engagement and can assemble and impart socially relevant information based on behaviors, movements, and attention orientations of their partner. Together, these studies suggest that attention orientation is conveyed and assessed by gorillas through a variety of interacting cues to navigate and modify social play interactions. VI DECLARATION .................................................................................................... II! ACKNOWLEDGEMENTS .................................................................................... III! ABSTRACT .......................................................................................................... V! FIGURES AND TABLES ..................................................................................... XI! Chapter 1: General Introduction ................................................................................................................ 14! 1. Primate vision ........................................................................................................................................ 14! 2. Gaze-following mechanisms ................................................................................................................. 16! 2.1. Detecting eyes and eye-like stimuli .......................................................................................... 17! 2.2. Additional cognitive mechanisms ............................................................................................. 19! 3. Gaze-following ...................................................................................................................................... 20! 3.1. Birds and reptiles ...................................................................................................................... 22! 3.2. Mammals .................................................................................................................................. 23! 3.3. Primates .................................................................................................................................... 24! 4. Attentional great apes ............................................................................................................................ 25! 5. Play behavior ......................................................................................................................................... 29! 5.1. Defining play ............................................................................................................................ 29! 5.2. Solitary and social play ............................................................................................................. 31! 5.3. Play functions and hypotheses .................................................................................................. 33! 5.3.1. Physical training ................................................................................................................ 33! 5.3.2. Social skills ........................................................................................................................ 34! 5.3.3. Cognitive skills ................................................................................................................... 35! 5.4. Gorilla play ............................................................................................................................... 38!
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