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'Patterns of Habitat Use and Segregation Among African Large Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2013 Patterns of habitat use and segregation among African large carnivores Cozzi, Gabriele Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-93646 Dissertation Originally published at: Cozzi, Gabriele. Patterns of habitat use and segregation among African large carnivores. 2013, University of Zurich, Faculty of Science. PATTERNS OF HABITAT USE AND SEGREGATION AMONG AFRICAN LARGE CARNIVORES Dissertation zur Erlangung der naturwissenschaftlichen Doktorwürde (Dr. sc. nat.) vorgelegt der Mathematisch-naturwissenschaftlichen Fakultät der Universität Zürich von Gabriele Cozzi von Vacallo (TI) Promotionskomitee: Prof. Dr. Bernhard Schmid (Leitung der Dissertation und Vorsitz) Prof. Dr. Marta Manser Dr. John W. McNutt Dr. Sarah Durant Zürich, 2013 PATTERNS OF HABITAT USE AND SEGREGATION AMONG AFRICAN LARGE CARNIVORES A CASE STUDY ON THE AFRICAN WILD DOG (LYCAON PICTUS), THE SPOTTED HYENA (CROCUTA CROCUTA) AND THE LION (PANTHERA LEO) Gabriele Cozzi Die vorliegende Dissertaion wurde am 27.11.2012 zur Begutachtung eingereicht. Promotionskomitee: Prof. Dr. Bernhard Schmid (Vorsitz) Prof. Dr. Marta Manser Dr. John W. McNutt Dr. Sarah Durant This thesis is dedicated to the memory of Christine B. Müller a mentor, a colleague, a friend Contents CONTENTS SUMMARY ........................................................................................................................................................................................ 1 ZUSAMMENFASSUNG .................................................................................................... 3 CHAPTER ONE General Introduction .......................................................................................................... 5 CHAPTER TWO Cozzi G., F. Broekhuis, J.W. McNutt, L.A. Turnbull, D.W. Macdonald & B. Schmid (2012) Fear of the dark or dinner by moonlight? Reduced temporal partitioning among Africa’s large carnivores. Ecology 93: 2590-2599 .............................................. 29 CHAPTER THREE Cozzi G., F. Broekhuis, J.W. McNutt, E.P. Willems, G. Schaepma-Strub & B. Schmid The ecology of fear among African territorial large carnivores: when being afraid of the cat becomes a dog’s life. To be submitted .............................................................. 67 CHAPTER FOUR Cozzi, G., F. Broekhuis, J.W. McNutt & B. Schmid (2013) Comparison of the effects of artificial and natural barriers on large African carnivores: Implications for inter- specific relationships and connectivity. Journal of Animal Ecology 83: 707-715 ........ 103 CHAPTER FIVE Cozzi, G., F. Broekhuis, J.W. McNutt & B. Schmid (2013) Density and habitat use of lions and spotted hyenas in northern Botswana and the influence of survey and ecological variables on call-in survey estimation. Biodiversity and Conservation 22: 2937-2956 .......................................................................................................................... 141 CHAPTER SIX General Discussion, Conservation and Management Implications ............................ 183 AKNOWLEDGEMENTS ............................................................................................... 199 CURRICULUM VITAE ................................................................................................. 202 Contents Summary SUMMARY This work investigated patterns of habitat use and segregation among species of the large African carnivore guild: the African wild dog, the spotted hyena and the lion. In particular, I studied what role temporal and spatial partitioning of activities, respectively, use of space may play in promoting coexistence. The study site was the Okavango Delta in northern Botswana, one of the few remaining places in Africa where the three species still co-occur in a pristine and undisturbed ecosystem. In Chapter Two we investigated the degree of temporal segregation between the allegedly diurnal wild dogs and the nocturnal spotted hyenas and lions. Wild dogs are commonly described as day active, and such behaviour has been described as an adaptation to evade interactions with the two other dominant species. We, however, showed a degree of temporal overlap considerably higher than previously described. Such overlap was mainly due to the unexpected nocturnal behaviour of wild dogs that was mainly influenced by moonlight availability, rather than by the activity of hyenas and lions. Our results thus raise some questions concerning the real role of spotted hyenas and lions in shaping the activity patterns of wild dogs and highlight how temporal overlap may only play a marginal role in enhancing coexistence among the species. Chapter Three demonstrated that the spatial distribution of wild dogs was negatively influenced by the distribution of lions, but not by that of spotted hyenas. In areas catheterized by a low lion presence, wild dogs may take advantage of the best resources, but they were restricted in their behaviour or even excluded from particular areas above a certain lion density, and this irrespectively of other variables such as habitat type and prey availability. Our results highlighted spatial segregation as a key mechanism promoting coexistence between wild dogs and lions. Considering the dramatic rate at 1 Summary which habitat loss and fragmentation are happening, the need for space that allows species coexistence should sound like a warning bell. In Chapter Four we investigated how the species are affected by and adapt to anthropogenic habitat modification and human activities near the boundary of a protected area. Specifically, we analysed how different types of barrier influenced the distribution and spatial interaction among species and discussed their effects at the community composition level. We showed that lions were restricted in their movements by an artificial fence erected to control the movement of ungulate species, while the other carnivore species freely crossed it. In contrast, lions were not obstructed in their movements by rivers, which represented an almost completely impassable barrier for the smaller spotted hyenas and wild dogs. Reinforcing the findings of Chapter Three, Chapter Four provided further evidence that wild dogs may take advantage of areas characterized by a low lion presence, particularly for critical activities such as raising offspring. Chapter Five presents density estimates for spotted hyenas and lions in the study area. An accurate estimate was necessary because the intensity of interaction between competing species partly depends on the density at which each species occurs. We calculated a density of 15.4 adult hyenas /100 km2 and 16.2 lions /100 km2. These figures compared well with other highly productive ecosystems of southern and eastern Africa. The hyena and lion density in the Okavango Delta should therefore be considered to be between medium and high. 2 Zusammenfassung ZUSAMMENFASSUNG Die vorliegende Dissertation untersuchte Raumnutzung und Segregation zwischen verschiedenen Arten der Gilde afrikanischer Großraubtiere: der Afrikanische Wildhund, die Tüpfelhyäne und der Löwe. Insbesondere studierte ich, wie die zeitliche und räumliche Aufteilung der Aktivitäten, beziehungsweise der Habitatsnutzung, das Zusammenleben der Tiere beeinflusst und fördert. Das Untersuchungsgebiet war das Okavango Delta im Norden Botswanas, einer der wenigen verbliebenen Orte in Afrika, wo die drei Arten in einem unberührten und ungestörten Ökosystem vorkommen. In Kapitel Zwei untersuchten wir die zeitliche Trennung zwischen den angeblich tagaktiven Wildhunde und den nachtaktiven Tüpfelhyänen und Löwen. Wildhunde werden allgemein als tagaktiv beschrieben, und dieses Verhalten ist als Anpassung beschrieben worden, um Wechselwirkungen mit den anderen beiden dominanten Arten zu reduzieren. Wir konnten jedoch eine wesentlich höhere zeitliche Überlappung zwischen den drei Arten aufzeigen als zuvor beschrieben worden war. Diese Überlappung war vor allem durch das unerwartete, nächtliche Verhalten der Wildhunde bedingt. Ihre Nachtaktivität war durch die Mondlichtverfügbarkeit, jedoch nicht durch die Aktivität von Hyänen und Löwen, beeinflusst. Unsere Ergebnisse werfen somit einige Fragen über die eigentliche Rolle der Tüpfelhyänen und Löwen bei der Gestaltung der Aktivitätsmuster der Wildhunde auf. Ausserdem zeigen unsere Resultate, dass zeitliche Überlappung nur eine marginale Rolle bei der Verbesserung der Koexistenz zwischen den Arten spielen darf. Kapitel Drei zeigte, dass die räumliche Verteilung von Wildhunden durch die räumliche Verteilung von Löwen negativ beeinflusst wird, aber nicht durch jene von Tüpfelhyänen. In jenen Gebieten, welche durch eine niedrige Löwenpräsenz charakterisiert sind, können Wildhunde von den besten Ressourcen profitieren. Ab einer bestimmten Löwendichte sind sie aber in ihrem Verhalten eingeschränkt oder gar aus bestimmten 3 Zusammenfassung Bereichen ausgeschlossen – und dies unabhängig von anderen Variablen wie Vegetationstyp oder Beuteverfügbarkeit und -verteilung. Unsere Ergebnisse weisen darauf hin, dass räumliche Segregation ein wichtiges Element zur Förderung der Koexistenz zwischen Wildhunde und Löwen ist. Angesichts der dramatischen Geschwindigkeit, mit welcher Lebensraumverlust und -fragmentierung vorwärtsschreiten, sollte uns der Raumbedarf, welcher für das Zusammenleben von Arten notwendig ist, alarmieren. In Kapitel Vier
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