Spatial and Temporal Responses of Animals to Landscape Heterogeneity, Predation Risk, and Human Activity By Kaitlyn M. Gaynor A dissertation submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy in Environmental Science, Policy, and Management in the Graduate Division of the University of California, Berkeley Committee in charge: Professor Justin S. Brashares, Chair Professor Arthur D. Middleton Professor Eileen A. Lacey Professor Mary E. Power Spring 2019 Abstract Spatial and Temporal Responses of Animals to Landscape Heterogeneity, Predation Risk, and Human Activity by Kaitlyn M. Gaynor Doctor of Philosophy in Environmental Science, Policy, and Management University of California, Berkeley Professor Justin S. Brashares, Chair The global expansion of human activity has had profound consequences for wildlife. Research has documented the effects of widespread habitat destruction and defaunation on species and ecosystems, but the more subtle pathways through which humans alter the natural world have largely escaped quantification. Much like apex predators, humans can instill strong fear in wild animals, which may adjust their activity to avoid contact with humans. In this dissertation, my collaborators and I examine pathways through which human disturbance, predation risk, and environmental heterogeneity influence animal behavior and distribution. We review the literature and synthesize theory to develop a novel framework for studying landscapes of fear, and we apply this framework in a global meta-analysis and field studies from California and Mozambique to understand how large mammals perceive and respond to spatial and temporal patterns of risk from humans and carnivores. We consider links between risk and response in complex systems with multiple predators or multiple prey species, and we explore ecology of fear dynamics in the context of seasonality, human disturbance, and restoration. Together, this work integrates disciplines of behavioral ecology, community ecology, and landscape ecology, applying predator-prey theory to understand the role of humans in ecological communities. By elucidating behavioral pathways linking human disturbance to wildlife community dynamics, this research contributes to our understanding of wildlife ecology in human-dominated landscapes and highlights mechanisms for human-wildlife coexistence. 1 Table of Contents ACKNOWLEDGEMENTS …………………………………………………………………………. ii CHAPTER 1 Introduction ……………………………………………………………………………... 1 CHAPTER 2 Landscapes of fear: spatial patterns of risk perception and response …………………… 5 CHAPTER 3 Black-tailed deer navigate contrasting patterns of risk from hunters and mountain lions in space and time .………………………………………………………………… 24 CHAPTER 4 The influence of human disturbance on wildlife nocturnality …………………………. 54 CHAPTER 5 Effects of human settlement and roads on diel activity patterns of elephants .………… 77 CHAPTER 6 Seasonal dynamics of an ungulate assemblage in a savanna floodplain landscape.......... 94 CHAPTER 7 Concluding remarks …………………………………………………………………... 146 REFERENCES ………………………………………………………………………………….. 148 i AcknowledGements I had always imagined the dissertation to be an isolating and solitary intellectual endeavor and I am so grateful to have been proven so wrong. My experience at UC Berkeley and in the field has been enriched by the support, ideas, and company of a truly long list of remarkable people. First, I am grateful for the support of my advisor Justin Brashares, who has challenged me to apply my interests in behavioral ecology to interdisciplinary questions and taught me by example how to be not only a better scientist and scholar but a better leader, communicator, mentor, and human being. A huge thanks to the other members of my committee: to Arthur Middleton, for the great questions and collegial conversation, to Mary Power, for reminding me to find the joy in fieldwork and scientific inquiry, and to Eileen Lacey, for her thoughtful edits and insightful feedback. And to my qualifying exam committee, who brought me back to the basics and inspired my first dissertation chapter: Eileen, Mary, Claire Kremen, and Ian Wang. I am fortunate to have been surrounded by the most supportive lab group imaginable, and am grateful for the generous feedback on everything from study design to grant proposals to navigating Berkeley bureaucracy: Katie Fiorella, Tristan Nuñez, Katy Seto, Cheryl Hojnowski, Lauren Withey, Ryan Marsh, Mario Klip, Eric Dougherty, Dave Kurz, Christine Wilkinson, Phoebe Parker-Shames, Kendall Calhoun, Amy Van Scoyoc, Millie Chapman, Lindsey Rich, and Jennie Miller. A special shout-out to Briana Abrahms, my weekly goal buddy of five years and counting, and to Alex McInturff, my cohort-mate and international travel companion. GORONGOSA NATIONAL PARK, MOZAMBIQUE My research in Gorongosa National Park, Mozambique would not have been possible without the support and encouragement of a fantastic group of people. Many thanks to the assistants and interns who joined me in the field and helped me with my Portuguese: Castiano Lencastro, Diolinda Mundoza Semente, and Gabriela Curtiz. A special thanks to my first assistant, Mateus Dapitaia, for his endless patience and for welcoming me into his family. I am indebted to the fiscais of Gorongosa National Park, who kept me safe from all dangers big and small, especially Beca (who can navigate to every camera in the park better than my GPS) and Sergio (who stuck with me even after an elephant crushed our truck). A huge thanks to Marc Stalmans for welcoming me into the park, advising me on my research, facilitating logistics, and connecting me with colleagues. And to Jason Denlinger, who solved every single one of my annoying problems with a smile. Thanks to the entire E.O. Wilson Lab team, including Piotr Naskrecki, Tongai Castigo, and Tara Massad. Thanks to Mike Marchington, for making sure thousands of batteries found their way to the park from South Africa, and Anne Marchington, for making life in Chitengo not only livable but enjoyable. I’m grateful for the support of Mateus Mutemba and Pedro Muagura, who have prioritized research in the park. Finally, thanks to Greg Carr—your passion, curiosity, and dedication are an inspiration. To the entire Gorongosa Restoration Project team: keep up the great work. It was intimidating to show up at a remote field site on my own, and I am grateful to Rob Pringle and his group for welcoming me with open arms and advocating for me and supporting me throughout my years in the park. I am fortunate to be an honorary member of the Princeton crew—all the benefits of the collegiality and friendship without having to collect antelope poop all of the time. Many thanks to Jen Guyton, Josh Daskin, Arjun Potter, Justine Atkins, Matt Hutchinson, Paola Branco, Ryan Long, Elyce Gosselin, and Katie Grabowski for sharing your ii science, your space, and your stories. I am also grateful for the company and collaboration of Tara Easter, my camera trapping comrade and workshop co-instructor. And cheers to everybody with whom I’ve enjoyed a beer at Chikalango: Matt Jordan, Jacques Nels, Zak Pohlen, Callie Gesmundo, Teague Scott, Cat Chamberlain, Megan Flaherty, Max Prager, Dominique DeMille, Casey Ortbahn, and countless others who have passed through over the years. Thanks also to the team at WildCam Gorongosa and HHMI BioInteractive, including Bridget Conneely, who have brought the wildlife of Gorongosa into living rooms and classrooms all over the world while facilitating the processing of camera trap data through citizen science. I’m looking forward to continued collaboration. Finally, my Gorongosa camera trap data only exist thanks to the help of an amazing group of undergraduate women, who sorted through thousands of photos of baboons, waterbuck, and warthogs (and thousands more of grass): Alison Ke, Tammy Gu, Ava Wu, Mia Silverberg, Hadley Rosen, Madeleine Levy, Claire Jurgensen, Edith Lai. Thank you for your patience, your thought-provoking questions, and your dedication. Finally, thanks to all of the funders who supported my fieldwork in Mozambique: National Science Foundation, Rufford Foundation, Explorers Club Mamont Scholars Program, Animal Behavior Society, IdeaWild, UC Center for African Studies Rocca Fellowship, UC Institute for International Studies, Sigma Xi Berkeley Chapter, and Badreyyah Alireza. HOPLAND RESEARCH AND EXTENSION CENTER, CALIFORNIA I was spoiled to have two amazing field sites for my dissertation research, and am grateful to Alex and Justin for welcoming me as a collaborator at the UC Hopland Research and Extension Center in Mendocino. It has been a joy to get acquainted with the wildlife in my new backyard, and to realize that I don’t have to fly to Africa to find a stunningly beautiful and complex savanna system. Thanks to the entire staff of HREC for consistently going above and beyond to make us feel at home and to help our fieldwork go smoothly, especially former Director Kim Rodrigues and Interim Director John Bailey. A huge shout-out to the field team: Alison Smith for her superhuman ability to juggle so many things at once, and Troy McWilliams for making captures look easy and always having the inside scoop on the lives of the HREC deer. Thanks also to Hannah Bird, Amber Shrum, Tom Seward, Zane Kagley, and Shane Feirer. It has been my pleasure and privilege to work with our amazing field managers Jessie Roughgarden, Lara Brenner, and especially Janelle Dorcy, and with our lab technicians Bryan Bach, Ashling