ABSTRACT FAVREAU, JORIE M. The effects of food abundance, foraging rules and cognitive abilities on local animal movements. (Under the direction of Roger A. Powell.) Movement is nearly universal in the animal kingdom. Movements of animals influence not only themselves but also plant communities through processes such as seed dispersal, pollination, and herbivory. Understanding movement ecology is important for conserving biodiversity and predicting the spread of diseases and invasive species. Three factors influence nearly all movement. First, most animals move to find food. Thus, foraging dictates, in part, when and where to move. Second, animals must move by some rule even if the rule is “move at random.” Third, animals’ cognitive capabilities affect movement; even bees incorporate past experience into foraging. Although other factors such as competition and predation may affect movement, these three factors are the most basic to all movement. I simulated animal movement on landscapes with variable patch richness (amounts of food per food patch), patch density (number of patches), and variable spatial distributions of food patches. From the results of my simulations, I formulated hypotheses about the effects of food abundance on animal movement in nature. I also resolved the apparent paradox of real animals’ movements sometimes correlating positively and sometimes negatively with food abundance. I simulated variable foraging rules belonging to 3 different classes of rules (when to move, where to move, and the scale at which to assess the landscape). Simulating foraging rules demonstrated that variations in richness and density tend to have the same effects on movements, regardless of foraging rules. Still, foraging rules affect the absolute distance and frequency of movements. In my third set of simulations, I simulated a range of spatial and temporal cognitive constraints and demonstrated that omniscience is not necessarily the optimal cognitive state from an energetic standpoint. I tested my hypotheses on the effects of food abundance with data from free ranging female black bears (Ursus americanus) in Pisgah Bear Sanctuary (North Carolina, USA) and female kinkajous (Potos flavus) in Parque Nacional Soberanía (Panama), two species with low predation risk. Depending on the season, black bear movements can be explained, by food patch richness, density or both richness and density. Female kinkajou move length correlated positively with patch richness and density. The number of moves of female kinkajous correlated negatively with patch richness of all foods that kinkajous eat and selectivity. In contrast, food patch richness and density did not affect male kinkajous’ moves (length and number). Instead, male kinkajous increased foraging time on all foods that they eat as patch density decreased. Male kinkajous also decreased their selectivity on subsets of foods that they eat as patch richness increased. My results are broadly interesting because they demonstrate that the success of the habitat productivity hypotheses depends on how food is measured (patch richness or density), sex, species, and foraging and non-foraging behaviors (foraging selectivity, responses to moonlight). THE EFFECTS OF FOOD ABUNDANCE, FORAGING RULES AND COGNITIVE ABILITIES ON LOCAL ANIMAL MOVEMENTS by JORIE M. FAVREAU A dissertation submitted to the Graduate Faculty of North Carolina State University in partial fulfillment of the requirements of the Degree of Doctor of Philosophy ZOOLOGY Raleigh 2006 APPROVED BY: __________________________ _________________________ Dr. Nicholas M. Haddad Dr. George R. Hess __________________________ _________________________ Dr. Roland W. Kays Dr. Richard A. Lancia __________________________ _________________________ Dr. Michael S. Mitchell Dr. Roger A. Powell Chair of Advisory Committee ii BIOGRAPHY I chose Illinois Wesleyan University (Bloomington, Illinois) for undergraduate study because I was interested in veterinary medicine and Wesleyan has an excellent biology program. At Wesleyan I volunteered at a small zoo and realized that zoology and conservation interested me, not the medical aspect of animal biology. Despite changing my focus, I finished a Bachelor's degree in biology at Wesleyan because a solid biology background is important to understanding zoology. After graduating from Wesleyan in 1991, I monitored bald eagle nests as a temporary biologist for the Arizona Game and Fish Department. Next, I landed a job as a keeper at Brookfield Zoo in Chicago because I had interned in the hoofed stock department after my sophomore year and worked as a summer seasonal in the Children's Zoo after my junior year. I spent most of the next five years in the carnivore department. Working at Brookfield Zoo, an excellent institution with a solid commitment to the animals' well being and to conservation, was a wonderful experience because I witnessed a variety of animal behavior and became familiar with conservation efforts. Nonetheless, during my fourth year at the zoo, I desired to begin graduate school to gain skills necessary for working with conservation organizations and to return to field work. Conservation requires understanding animals’ relations to their habitats and the socioeconomic and cultural factors of the people who share that habitat. I wanted to develop those skills and management skills. DePaul University's (Chicago, Illinois) program requires students to tailor their courses to reflect their profession, thereby providing management knowledge directly applicable to their profession. I began classes in 1994 while working full time at the zoo. Class work centered on real projects at the iii zoo and other conservation organizations. In 1997, I completed my M.A. with an emphasis on wildlife conservation management. Meanwhile, I pursued a series of seasonal field jobs to learn field research techniques. In 1996, I tracked and monitored reintroduced California condors for the United States Fish and Wildlife (USFWS) California Condor Program. Then, I searched for the Po-o-uli, an endangered Hawaiian honeycreeper for the United States Geological Survey, Biological Resources Division (USGS-BRD). I ran a Monitoring Avian Productivity and Survivorship (MAPS) station in Indiana, censused hawks in the Florida Keys, and surveyed flying squirrels, voles, and Peromyscus for the United States Department of Forestry (USFS) in Alaska. In fall 1999, my husband and I moved to Albany New York where I worked for The Adirondack Council, a non-profit, non-governmental organization that is dedicated to protecting the Adirondack Park through advocacy, education, and legal action. Working for the Adirondack Council was a piece of my experiential education that I would not have sought; however, I now view that experience as an important component of my pre- PhD experience. While at the Adirondack Council, I realized it was time to continue my formal education. I was interested especially in applied wildlife management (particularly nongame species) and conservation of threatened species/ecosystems. I chose North Carolina State University (Raleigh, NC) to earn a PhD in zoology because NCSU fit the parameters that I sought. In addition, Roger Powell and his research intrigued me. I TA’d a variety of courses at NCSU such as Mammalogy and Molecular Biology, which was a good experience because it introduced me to teaching, refreshed my iv undergraduate biology knowledge, and introduced me to techniques such as polymerase chain reactions (PCR) that had not existed while I was an undergraduate. The dedication and commitment of some of the professors for whom I taught, especially Dr. Harold Heatwole and Dr. Sam Mozley, also inspired me and taught me how to teach. While a TA, I realized that an ideal career for me was to teach at a small school where learning was a primary objective. I also wanted a school that would be near my research site and places where I could frequently take students into the field. Due to family obligations, I moved to New York after finishing my coursework. I planned to not work and, instead, finish writing my dissertation during the next year. Fate intervened in the form of an offer to teach General Biology at Siena College in Loudonville NY. My teaching schedule was demanding but I was not worried as a position as a visiting instructor was a great way to get teaching experience and I figured I would finish my dissertation the next year. As it turned out, Siena College asked me to teach a second year and I didn’t finish my dissertation that year either. In fall 2005, I began teaching in a tenure track position in the Fish and Wildlife Science Program at Paul Smith’s College (Paul Smith’s, New York), the only baccalaureate granting college in the Adirondacks. At Paul Smith’s, I teach courses such as Conservation Biology and Wildlife Management. In June 2006, I defended my dissertation. v ACKNOWLEDGEMENTS I am grateful to Nathan for making personal and professional sacrifices to support me while I was in graduate school. I am grateful to Siena College for giving me the opportunity to gain teaching experience as a visiting instructor. I am immensely grateful to interim Provost Helena Sturnick, the faculty, and students at Paul Smith’s College for their faith and support as I finished my dissertation. I am grateful to the Matlab community for allowing me to lurk on their website to learn how to program! I learned quite a bit about modeling in this process, including in ways that I never expected. So, I thank Roger Powell for being a model of academic purity, Roland Kays of the New York State Museum for being a model of realism and practicality, and fate for giving me Yin and Yang as my co-chairs. I thank Richard Lancia for being a model of the wildlife biologist that I hope to become. I thank Mike Mitchell for modeling (or at least trying to model) me into an upstanding NCSU student. I thank Nick Haddad for adopting me into his lab and allowing me to be part of his working laboratory model. I thank George Hess for introducing me to biological models . . . and for being a model teacher. I thank other models of teaching: Sam Mozley, Harold Heatwole, and Jennifer Thompson. I thank Allan Weatherwax at Siena College for modeling how to balance vital research and effective teaching at a small, undergraduate institution. I learned much from each of you. And, importantly, I thank Jennifer Sevin for being a model of fun and productivity. vi TABLE OF CONTENTS List of Tables ……………………………………………………………………………..x List of Figures …………………………………………………………………………..xiii Chapter 1: Introduction to movement ecology..................................................................1 Importance and application of movement ecology.........................................................1 Overview of dissertation.................................................................................................4 Literature cited................................................................................................................7 Chapter 2: Standard movement terms and definitions.....................................................14 Need to standardize movement terms...........................................................................14 Movement Terms..........................................................................................................17 Basic movement terms..............................................................................................17 Terms describing movement duration, direction and distance.................................20 Terms describing path shape.....................................................................................23 Terms describing home range shape and size...........................................................25 Terms describing differential use of home ranges....................................................27 Types of movement...................................................................................................32 Conclusions...................................................................................................................33 Literature cited..............................................................................................................34 Chapter 3: Effects of patch richness, density and distribution on foraging movements..54 Abstract.........................................................................................................................54 Introduction...................................................................................................................55 Effects of food abundance on movement..................................................................56 Effects of food distribution on movement................................................................58 Advantages of modeling movement.........................................................................59 Objectives.................................................................................................................60 Methods.........................................................................................................................60 Landscape.................................................................................................................60 Effects of food abundance and distribution on movement.......................................61 Animal movement.....................................................................................................62 Results and discussion..................................................................................................64 Travel cost and consumption amount.......................................................................64 Patch richness when food patches were distributed uniformly and randomly..........65 Patch richness when food patches were aggregated.................................................66 Patch density when food patches were distributed randomly and uniformly...........68 Patch density when food patches were aggregated...................................................69 Effects of food distribution.......................................................................................70 Summary: factors affecting move length and number of moves..............................71 Hypotheses................................................................................................................71 Resolution of seemingly paradoxical field data........................................................72 vii Literature cited..............................................................................................................73 Chapter 4: Effects of foraging rules on move length and number of moves...................86 Abstract.........................................................................................................................86 Introduction...................................................................................................................87 Departure rules..........................................................................................................88 Patch selection rules..................................................................................................89 Scale of assessment...................................................................................................90 Absolute and relative effects of rules on movements...............................................91 Objectives.................................................................................................................91 Methods.........................................................................................................................92 Simulations: landscape and animal movement.........................................................92 Departure rules..........................................................................................................92 Cell selection rules....................................................................................................93 Assessment scale.......................................................................................................93 Output.......................................................................................................................94 Results...........................................................................................................................94 Effects of departure rules on movements..................................................................94 Effects of cell selection rules on movements............................................................95 Effects of assessment scale on movements...............................................................96 Optimal foraging rules..............................................................................................96 Discussion.....................................................................................................................97 Effects of departure rules..........................................................................................97 Effects of cell selection rules....................................................................................99 Effects of assessment rules.......................................................................................99 Local movement predictions...................................................................................100 Optimal foraging rules............................................................................................101 Foraging rules as mechanisms for “paradoxical” movements in response to changes in food abundance...................................................................................................101 Application of results..............................................................................................103 Literature cited............................................................................................................105 Chapter 5: Cognitively constrained animals can be as fit as omniscient animals.........119 Abstract.......................................................................................................................119 Introduction.................................................................................................................120 How does temporally and spatially variable information affect movement?.........120 Should natural selection favor omniscient animals?...............................................124 Methods.......................................................................................................................126 Results.........................................................................................................................128 Effects of cognitive states on move lengths............................................................128 Effects of cognitive states on number of moves.....................................................129 Effects of cognitive states on home range size and use..........................................130 Effects of cognitive states on energetics.................................................................132 Discussion...................................................................................................................133 Effects of cognitive states on move lengths............................................................134 viii Effects of cognitive states on home range size and use..........................................135 Effects of cognitive states on energetics.................................................................136 The value of omniscience.......................................................................................136 Application of results..............................................................................................139 Conclusions.................................................................................................................139 Literature cited............................................................................................................140 Chapter 6: Food patch richness and density partially explain travel rate of black bears (Ursus americanus).........................................................................................................155 Abstract.......................................................................................................................155 Introduction.................................................................................................................156 Bear food in Pisgah Bear Sanctuary.......................................................................156 Bears’ knowledge about food locations..................................................................157 Foraging rules and bear foraging behavior.............................................................158 Hypotheses to explain the “paradoxical” movement of black bears in Pisgah.......160 Methods.......................................................................................................................161 Study site.................................................................................................................161 Bear food.................................................................................................................161 Bear travel rate........................................................................................................163 Hypotheses..............................................................................................................164 Results.........................................................................................................................165 Bear path lengths and travel rates...........................................................................165 Bear food.................................................................................................................165 Testing hypotheses..................................................................................................166 Discussion...................................................................................................................167 Travel rate...............................................................................................................167 Effects of seasonal food patch richness and density on travel rate.........................167 Foraging rules and cognitive constraints as explanations for observed bear travel rates.........................................................................................................................170 Wildlife Management.............................................................................................171 Acknowledgements.....................................................................................................172 Literature cited............................................................................................................172 Chapter 7: Food abundance and moonlight illumination affect kinkajou (Potos flavus) moves..............................................................................................................................185 Abstract.......................................................................................................................185 Introduction.................................................................................................................186 Kinkajous as a model organism..............................................................................186 What dietary categories affect movement?.............................................................187 What other factors that affect kinkajou movement?...............................................188 Methods.......................................................................................................................189 Study site.................................................................................................................189 Fruit patch abundance.............................................................................................189 Kinkajou movements..............................................................................................190