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Foraging Ecology of Shorebirds at a Stopover Site: Niche Dynamics, Aggression and Resource Use in Delaware Bay

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Foraging Ecology of Shorebirds at a Stopover Site: Niche Dynamics, Aggression and Resource Use in Delaware Bay City University of New York (CUNY) CUNY Academic Works All Dissertations, Theses, and Capstone Projects Dissertations, Theses, and Capstone Projects 10-2014 Foraging Ecology of Shorebirds at a Stopover Site: Niche Dynamics, Aggression and Resource Use in Delaware Bay Ivana Novcic Graduate Center, City University of New York How does access to this work benefit ou?y Let us know! More information about this work at: https://academicworks.cuny.edu/gc_etds/457 Discover additional works at: https://academicworks.cuny.edu This work is made publicly available by the City University of New York (CUNY). Contact: [email protected] FORAGING ECOLOGY OF SHOREBIRDS AT A STOPOVER SITE: NICHE DYNAMICS, AGGRESSION AND RESOURCE USE IN DELAWARE BAY by IVANA NOVCIC A dissertation submitted to the Graduate Faculty in Biology in partial fulfillment of the requirements for the degree of Doctor of Philosophy, The City University of New York 2014 © 2014 IVANA NOVCIC All Rights Reserved i This manuscript has been read and accepted for the Graduate Faculty in Biology in satisfaction of the dissertation requirement for the degree of Doctor of Philosophy. ______________ ______________________________ Date Chair of Examining Committee Dr. Richard R Veit College of Staten Island ______________ _______________________________ Date Executive Officer Dr. Laurel A. Eckhardt _______________________________ Dr. Lisa L. Manne College of Staten Island _______________________________ Dr. Mark E. Hauber Hunter College _______________________________ Dr. Mark E. Siddall American Museum of Natural History _______________________________ Dr. David Mizrahi New Jersey Audubon Society Supervisory Committee The City University of New York ii Abstract Foraging ecology of shorebirds at a stopover site: niche dynamics, aggression and resource use in Delaware Bay By Ivana Novcic Advisor: Richard R. Veit Classical ecological theory predicts that generally similar species ought to partition resources in order to minimize competition amongst themselves. This basic idea becomes complex when one is dealing with species that migrate over thousands of miles and forage in a broad diversity of habitats and geographical locations. I studied a suite of migratory sandpipers, and asked whether they partitioned niches at a major migratory stopover in Delaware Bay. During migration, shorebirds form large, usually mixed-species flocks, which forage on marshes, mudflats, beaches or similar two-dimensional habitats where all individuals are distributed on the same horizontal plane. These habitats are often affected by the tidal cycle forcing birds to feed at the same time, which leads to intensified competition through both depletion and interference. Using multidimensional niche approach, I explore whether coexisting shorebirds separate by time of passage, habitat use and foraging behavior, during northbound migration, at the time shorebirds gather in large numbers to capitalize on eggs of spawning horse shoe crabs (Limulus polyphemus) (Chapter 1). I hypothesize that differential migration timing is the most important dimension for separation of species. Also, I investigate aggressive interactions of shorebirds (Chapter 2), hypothesizing that birds will exhibit more aggression toward conspecifics than to heterospecifics, as individuals of the same species, due to morphological similarity, more often compete for resources. Finally, I examine the importance of horseshoe crab eggs for study iii species through PCR of prey DNA from birds’ fecal samples with horseshoe crab specific primers that I designed for this study (Chapter 3). My research demonstrates that shorebird species mostly separate by differential timing of spring migration during stopover in Delaware Bay. Also, the study confirms higher measures of aggression between conspecifics than between heterospecifics, although the incidence of interspecific interactions was higher than previously reported for shorebirds, most likely due to interspecific dominance relationships. The presence of interspecific aggression in mixed-species foraging flocks emphasizes the importance of temporal segregation between migratory species, as such ecological segregation may reduce the opportunity for interspecific aggressive encounters, which in turn can have positive effects on birds’ time and energy budget during stopover period. In addition, this study corroborates the importance of horse shoe crab eggs for migrating shorebirds in this crucial stopover area. iv Acknowledgements I would like to thank to my advisor, Richard Veit, for his help throughout this project, as well as my dissertation committee members: Mark Hauber, Lisa Manne, David Mizrahi and Mark Siddall, for useful comments that improved my thesis. I am very grateful to William Symondson from Cardiff School of Bioscience, Cardiff University, whose expertise and patience were essential for completion of the molecular analysis of shorebirds’ diet. Also, I appreciate Silke Waap, Anziza Mahyudin and Mario Barbato for their help during my stay in Cardiff. Many thanks to Frank Burbrink for allowing the use of his lab for this research, as well as Ed Myers, Xin Chen and Alex McKelvy for their help with lab work. Also, I would like to thank to Morgan Tingley and Holly Goyert for advice on statistics in crucial moments for my project, while for help with field work I am thankful to Kim Peters, Patti Hodgetts, Nellie Tsipoura and especially to Vince Elia, whose proficiency in bird counting considerably improved the quality of collected data. From the American Museum of Natural History, I would like to thank to Sebastian Kvist and Louis Sorkin for help with identification of invertebrates, and to Lydia Garetano and Paul Sweet for access to bird collections and tissue samples. Thank you to Eugenia Naro-Maciel, John Verzani and William Wallace for suggestions that helped me solve the problems with DNA extractions, statistical analyses and invertebrate identification. In addition, I am thankful to Amanda Dey and New Jersey and Delaware Divisions of Fish and Wildlife for providing data on horseshoe crab egg abundance. This study was supported by a CUNY Doctoral Students Research grant and PSC-CUNY. v Table of Contents Introduction ............................................................................................................................... 1 Chapter 1: Niche dynamics of coexisting shorebirds at a stopover site: Foraging behavior, habitat choice and migration timing in Delaware Bay ................................................................ 7 Abstract ................................................................................................................................. 7 Introduction ........................................................................................................................... 8 Methods ................................................................................................................................ 12 Fieldwork ........................................................................................................................ 12 Foraging microhabitats .................................................................................................. 14 Foraging behavior .......................................................................................................... 15 Video analysis ................................................................................................................. 16 Data analysis .................................................................................................................. 17 Results ................................................................................................................................... 20 Migration timing .............................................................................................................. 20 Microhabitat use .............................................................................................................. 21 Foraging behavior ........................................................................................................... 22 Migration timing, microhabitat use and foraging behavior ............................................ 24 Discussion .............................................................................................................................. 25 Tables ..................................................................................................................................... 32 Figures .................................................................................................................................... 40 Supplementary material ......................................................................................................... 46 Chapter 2: Aggressive interactions in mixed-species flocks of foraging shorebirds during spring stopover ........................................................................................................................................ 47 Abstract .................................................................................................................................. 47 Introduction ............................................................................................................................ 48 Methods .................................................................................................................................. 52 Fieldwork .......................................................................................................................
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