SPACE USE OF AFRICAN WILD DOGS IN RELATION TO OTHER LARGE CARNIVORES IN HLUHLUWE-IMFOLOZI PARK, SOUTH AFRICA by Angela M. Darnell A Thesis Presented to The Faculty of Humboldt State University In Partial Fulfillment Of the Requirements for the Degree Master of Science In Natural Resources: Wildlife May, 2012 ii ABSTRACT Space Use of African Wild Dogs in Relation to Other Large Carnivores in Hluhluwe- iMfolozi Park, South Africa Angela M. Darnell Interaction between species through competition is one of the principle processes shaping the structure of ecological communities. Competition can have significant effects on the behavior, distribution, and ultimately the population dynamics of species, particularly when the animals utilize similar resources. Extensive diet overlap between large African carnivores is associated with high levels of competition which is particularly evident in the interactions between lions (Panthera leo), spotted hyenas (Crocuta crocuta) and African wild dogs (Lycaon pictus). Using GPS data collected from sightings of large carnivores in Hluhluwe-iMfolozi Park, South Africa, I tested the hypothesis that wild dogs’ space utilization was influenced by other large carnivores. I found that wild dogs had different responses to their two main competitors. They avoided lions, particularly during denning seasons, through a combination of spatial and temporal avoidance. However, the dogs did not alter space use relative to hyenas, likely because pack sizes were large enough to defend their kills adequately without the necessary effort of avoidance. Information from this study is important for managing current carnivore populations, especially as reintroductions and translocations are essential tools used for the survival of endangered African wild dogs. iii ACKNOWLEDGMENTS I would like to thank my advisor, Dr. Micaela Szykman Gunther, for her knowledge and guidance of my project. I would like to thank her and Jan A. Graf for letting me use their data for this project. I would like to thank my committee members Drs. Richard Golightly and Howard Stauffer for their expertise. I would like to thank Dr. Dave Druce and the other staff of Hluhluwe-iMfolozi Park (HiP) as well as Ezemvelo KZN Wildlife for allowing me to spend time in the park gaining additional information. Funding for this project was provided by Humboldt State University Sponsored Programs Foundation and the Institute for Wildlife Studies scholarship. My time in South Africa would not have been the same without my housemate (and often chauffer) Leanne Van der Weyde. I would also like to thank Zama Zwane for driving me around the park and for his invaluable knowledge of the wild dogs in HiP. I would also like to thank Dr. Robert VanKirk for his statistical guidance. I am grateful to my mother, Darlene Darnell, for reading my drafts more times than she could count. I would like to thank my father, Rodger Darnell, for always supporting me and challenging me to aim high. Finally, I would like to thank my lab-mates (particularly Hilary Cosby for editing all my drafts), fellow graduate students, friends and family for their love and support throughout my graduate school experience. iv TABLE OF CONTENTS ABSTRACT……………………………………………..…………………….…iii ACKNOWLEDGMENTS……………………………………………………..…iv TABLE OF CONTENTS…………………………………………………………v LIST OF TABLES………………………………………………………….……vi LIST OF FIGURES………………………………………………………….…..vii LIST OF APPENDICES…………………………………………………...……vii INTRODUCTION………………………………………………………………...1 METHODS………………………………………………………………………..7 Study Area………………………………………………………………...7 Data Collection…………………………………………………………..11 Spatial Interactions without Temporal Aspect………………………..….12 Spatial Interactions with Temporal Aspect…………………………..…..14 RESULTS………………………………………………………………………..20 Spatial Interacts without Temporal Aspect………………………………20 Spatial Interactions with Temporal Aspect………………………………28 DISCUSSION……………………………………………………………………34 LITERATURE CITED…………………………………………………………..42 v LIST OF TABLES Table Page 1 Notation descriptions used in text for calculating spatial and temporal interactions between African wild dogs, lions, and spotted hyenas in Hluhluwe-iMfolozi Park, South Africa, 2002-2004…..…………………,……..17 2 Comparisons of associations by co-occurrence for African wild dogs (α) and lions (β) in Hluhluwe-iMfolozi Park, South Africa, 2002-2004……….…….….29 3 Comparisons of association by co-occurrence for African wild dogs (α) and spotted hyenas (β) in Hluhluwe-iMfolozi Park, South Africa, 2003-2004..…....30 vi LIST OF FIGURES Figure Page 1 Study area showing location of Hluhluwe-iMfolozi Park in KwaZulu- Natal Province with inset showing location of province within South Africa during study in 2002-2006….…………………………………………….….........8 2 Map of Hluhluwe-iMfolozi Park showing vegetation, major rivers and roads during study in 2002-2006…………………………………………...........10 3 Average distance (in km) between neighboring non-overlapping African wild dog and lion core use areas (± SE) during denning (n=19) and not denning (n=10) seasons in Hluhluwe-iMfolozi Park, South Africa, 2002-2004...21 4 Mean percentage overlap (± SE) of home ranges of African wild dogs with lions (n=18) and spotted hyenas (n=10) between seasons in Hluhluwe-iMfolozi Park, South Africa, 2002-2004………………………………………...…….......22 5 Home ranges of African wild dogs, lions, and spotted hyenas during the a) denning season, b) post-denning season, and c) non-denning season in 2004Hluhluwe-iMfolozi Park, South Africa……………………………….....….23 6 Mean percentage of overlap (±SE) of core use areas of African wild dogs with lions (n=18) and spotted hyenas (n=10) between seasons in Hluhluwe- iMfolozi Park, South Africa, 2002-2004………………………...………...…….25 7 Core use areas of African wild dogs, lions and spotted hyenas during the a) denning season, b) post-denning season, and c) non-denning season in 2004 in Hluhluwe-iMfolozi Park, South Africa…………...…….……………………26 8 Mean percentage volume of intersection of home ranges (± SE) of African wild dogs with lions (n=9) and spotted hyenas (n=5) between seasons using intensity of use (3-dimensional overlap) in Hluhluwe-iMfolozi Park, South Africa, 2002- 2004...…………………………………………………………………………….27 vii LIST OF APPENDICES Appendix Page A Interaction within overlapping home ranges and core use areas for African wild dogs (A) and lions (B), showing spatial attraction or avoidance by each and interaction between groups (ixn) as well as deviation of odds from random in Hluhluwe-iMfolozi Park, South Africa, 2002-2004………………………………………….…...…54 B Interaction within overlapping home ranges and core use areas for African wild dogs (A) and spotted hyenas (B), showing spatial attraction or avoidance by each and interaction between groups (ixn) as well as deviation of odds from random in Hluhluwe-iMfolozi Park, South Africa, 2002-2004………………………………………………....55 viii INTRODUCTION Interaction between species through competition is one of the principle processes shaping the structure of ecological communities (Case and Gilpin 1974). Competition can have significant effects on the behavior, distribution, and ultimately the population dynamics of species (Case and Gilpin 1974, Chesson and Rosenzweig 1991, Creel et al. 2001). Interference competition in particular can play a major role in determining the abundance and distribution of competing species, and its effects usually have different impacts on the species involved (Case and Gilpin 1974, Linnell and Strand 2000, Creel et al. 2001). When animals utilize similar resources, larger competitors may exclude smaller competitors (Johnson et al. 1996). Competition between carnivores has been considered a key ecological factor affecting carnivore species within the same guild (Caro and Stoner 2003). Past studies have focused on effects of carnivores on prey or vice versa, while ignoring the significant effects carnivores can have on each other (Fuller and Kat 1990, Fuller and Sievert 2001, Owen-Smith and Mills 2008). Carnivores of the same guild may compete for similar prey resources, often resulting in smaller species either being excluded from or actively avoiding areas with higher densities of the larger competitor when in direct competition (Johnson et al. 1996, Linnell and Strand 2000, Creel et al. 2001). In Nepal, leopards (Panthera pardus) avoided habitats where tiger (Panthera tigris) densities were high (McDougal 1988). Mills and Mills (1982) found a significant pattern of avoidance of spotted hyenas (Crocuta crocuta) by the smaller brown hyenas (Hyaena brunnea). 1 Studies have also suggested that gray wolves (Canis lupus) displace and exclude coyotes (Canis latrans) from preferred habitat (Ballard et al. 2003). These studies illustrate the widespread pattern of avoidance and exclusion of smaller carnivores with less competitive advantage due to interference competition. Displacement by larger carnivores can result in serious effects on smaller species (Creel et al. 2001, Hunter and Caro 2008). Smaller competitors can be excluded from certain habitats with high prey density, reducing hunting opportunities (Creel et al. 2001, Caro and Stoner 2003). The amount of prey available may decrease when the density of larger species is high, and rates of food stealing may increase, forcing smaller species to travel long distances to forage (Caro and Stoner 2003). These situations result in smaller species being forced to increase the time and energy expended to obtain sufficient food. Increased time spent hunting limits the energy available for other essential behaviors, particularly