Nest Site Selection in Hudsonian Godwits: Effects of Habitat and Predation Risk
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NEST SITE SELECTION IN HUDSONIAN GODWITS: EFFECTS OF HABITAT AND PREDATION RISK A Thesis Presented to the Faculty of the Graduate School of Cornell University In Partial Fulfillment of the Requirements for the Degree of Master of Science by Rose J. Swift February 2016 © 2016 Rose J. Swift ABSTRACT The choice of where to nest has enormous fitness consequences for individuals and, when scaled up, can even affect population demography. Nest site selection in birds is, therefore, guided by a wide variety of proximate and ultimate factors, including protection from predators, access to favorable microclimates, proximity to food resources, and costs/benefits stemming from interactions with neighboring con- and hetero-specifics. Nevertheless, the features associated with, cues used to identify, and availability of favorable nest sites may be changing rapidly with accelerating human development and climate change, especially in the Arctic and sub-Arctic. I investigated how habitat characteristics, predation risk, and proximity to human disturbance (i.e., roads) influenced nest site selection for the Hudsonian Godwit (Limosa haemastica). First, I identified habitat characteristics most associated with nests of Hudsonian Godwits in two breeding populations – Beluga River, Alaska and Churchill, Manitoba. I was particularly interested in habitat attributes that are (or will be) strongly impacted by climate change and human development. Second, I assessed the spatial pattern of nests in Beluga River, Alaska to evaluate if there was evidence of aggregation and then determined the extent to which nest location and fate were associated with predation risk and habitat features. Hudsonian Godwits displayed strong preferences for certain habitat features when selecting nest sites, with individuals preferring nest sites with less bare ground and more graminoid and tall shrubby cover than random locations. Nest site preferences were relatively similar in the two study populations, though birds in Churchill also preferred to nest near shallow water. Given the demonstrated preferences of nesting godwits, our study suggests that habitat degradation due to grubbing by overabundant Snow Geese (Chen caerulescens) and changing climate will reduce the suitability and availability of habitats for breeding godwits. My study also provides evidence that, despite the importance of nesting habitat attributes, additional factors must be guiding nest site selection because nests of godwits were clustered within bogs. Because the spatial aggregation of nests was not explained by either underlying patchiness in habitat or by predation risk, I suggest that clustering may be related to social cues. Therefore, understanding behavioral processes can be as important as ecological requirements for habitat selection. BIOGRAPHICAL SKETCH Rose Swift was born in the Bay Area of California. Her parents, Brian and Karen, instilled a love of the outdoors in her from an early age. Camping when she was three months old and skiing when she was three years old, age was not a limitation to explore wilderness. She took that love for the outdoors with her as she began to explore universities. She went on to attend the University of California, Davis where she graduated with a B.S. in Wildlife, Fish, and Conservation Biology with an emphasis on Wildlife Biology. Her first field position was camping at a remote lake in Alaska for three months. There she not only fell in love with the spectacular landscape, but also became engaged with the bird communities she was seeing. Thanks to her field partner, Scarlett, an old pair of binoculars, and the trusty Sibley guide, she developed a passion for birds. At UC Davis, she engaged in many research projects and developed an honors thesis with Dr. John Wingfield investigating the phenological responses of arctic breeding birds to climate change. From these experiences launched a dream – to follow arctic breeding birds from the breeding grounds south and back north again. After graduating from UC Davis, Rose went on to work many field positions where she gained a variety of ornithological skills. She worked for several seasons at Hastings Natural History Reserve on Dr. Janis Dickinson’s long-term Western Bluebird (Sialia mexicana) behavioral ecology study. While there, she began to develop a project looking at the repeatability and heritability of egg size. From there, she flew to New Zealand to assist on a project looking at the breeding biology of North Island Brown Kiwi (Apteryx mantelli) where she had the opportunity to explore the biology of the endemic Kiwi Tick (Ixodes anatis) with Dr. Sarah Jamieson. From nest-searching in Montana to shorebird banding in California, iv she began to explore the topics she would choose to study for her graduate work. Soon she would begin to engage in conversations with professors, ultimately finding her love of birds would send her heading to the Lab of Ornithology at Cornell University. At Cornell, Rose has been involved in the Natural Resources and Lab of Ornithology communities. Under the guidance of Dr. Amanda Rodewald, she has been able to fulfill her dream to work with a long-distance migratory bird and travel to Alaska again. Rose will continue at Cornell for her Ph.D. continuing to work with Hudsonian Godwits on the breeding grounds, but she will spread her wings and begin to follow them from the non-breeding grounds in Chile north to Alaska. v For Garrett For your blood, sweat, and encouragement on this project you deserve so much. vi ACKNOWLEDGMENTS First and foremost, I wish to thank Dr. Amanda Rodewald for her supervision, support and expeditious editing. This project has not been a small one, and her attention to details, strategic thinking, and support have been essential in continuing this project. Special thanks also go to Dr. Nathan Senner for supervision, encouragement, and assistance from the beginning. I am indebted for his expertise, establishing this project, his years of data, and continued support. I also appreciate the contributions of my committee: Dr. John Fitzpatrick and Dr. Walter Koenig. I am also grateful for the statistical advice of Dr. Patrick Sullivan and the Cornell Statistical Consulting Unit. I am indebted to Brad Walker for his time and effort spent entering vegetation records and consulting with me on the floristics of the Churchill area. Additionally, I must thank Joe Barron and Katherine Hambury for their dedication to entering data. I send thanks to my lab mates, who contributed in many ways: Gemara Gifford, Ruth Bennett, Zephyr Mohr-Felson Züst, and Steven Sevillano Ríos for your input and assistance. And to all the scientists associated with the Bird Populations Studies and Conservation Sciences programs at the Lab of Ornithology for their expertise and advice. This work would never have been possible without the many field assistants on the project. To William Abbott, Amy Alstad, Hope Batcheller, Shawn Billerman, Jon DeCoste, Doug Gochfeld, Mike Harvey, Mike Hilchey, Tom Johnson, Andy Johnson, Julia Karagicheva, Jess Marion, Madi McConnell, Jay McGowan, Brittany Schultz, Glenn Seeholzer, Hannah Specht, Brad Walker, Ben Lagasse, Bret Davis, Kyle Parkinson, Justin Heseltine, James Klarevas-Irby, and Garrett MacDonald. Thank you. Your hours of work, companionship, and good humor were not overlooked. vii To the administrative staff at the Cornell Lab of Ornithology, I owe you gratitude for your assistance. In Beluga, Judy and Larry Heilman continue to provide us a happy home; the staff at ConocoPhillips has made our work run smoothly. Thanks to funding sources: the David and Lucile Packard Foundation, U.S. Fish and Wildlife Service, Faucett Family Foundation, National Science Foundation, the Athena Fund at the Cornell Lab of Ornithology, the Atkinson Sustainable Biodiversity Fund at Cornell University, Cornell Graduate School Mellon Fund, Cornell Chapter of Sigma Xi, American Museum of Natural History, Ducks Unlimited Canada, Churchill Northern Studies Centre, American Ornithologists’ Union, and Arctic Audubon Society. Finally, I wish to thank my partner Garrett, for his patience and understanding. Without his constant and unwavering support this would not have been possible and he has poured many hours into data collection, data entry, editing, and brainstorming with me. And my family: Brian, Karen, Lily, and Holly, for their support. My father deserves special thanks for reading through several edits. Thank you for instilling in me the confidence and curiosity that has gotten me here. viii TABLE OF CONTENTS BIOGRAPHICAL SKETCH iv ACKNOWLEDGEMENTS vii TABLE OF CONTENTS ix CHAPTER 1 1 LITERATURE CITED 6 CHAPTER 2 10 LITERATURE CITED 21 APPENDIX 31 CHAPTER 3 37 LITERATURE CITED 58 APPENDIX 93 ix CHAPTER ONE INTRODUCTION Breeding is a critical part of the avian annual cycle and can profoundly affect population dynamics. Choice of a nesting site, in particular, has consequences for both survival and reproductive success (Burger 1985, Martin 1992, Wiebe and Martin 1998), and nest preferences are generally assumed to have evolved to maximize fitness (Wiens 1989). For example, minimizing risk of nest predation, which is the primary cause of reproductive failure in birds, is an important driver of habitat selection, including nest sites (Martin 1993). Thus, patterns of nest placement are assumed to reflect the fitness benefits of nesting in preferred habitats (Cody 1981, Wiens 1989, Martin 1998). Species typically display strong nest site preferences because the success