Magister Scientiae (Biodiversity and Conservation Biology) / Human-wildlife conflict is an ongoing issue worldwide. Within South Africa, human-carnivore conflict (HCC) as a result of carnivore depredation on small-livestock causes large-scale losses, and promotes the use of predator management tools by farmers. Despite being one of the major offenders involved in HCC, caracals, and their ecology in particular, are understudied. This is mainly due to high levels of persecution, coupled with their elusive nature. Within the Karoo region of South Africa, pastoralists make use of large-scale lethal predator controls in an attempt to remove the offenders, or non-lethal predator controls to protect livestock and deter predators. However, the effects of these various predator control techniques on caracal ecology have not been widely tested. Therefore, the aims of this thesis were, firstly, to assess caracal diet on lethal and non-lethal treatment farms in the Karoo; secondly, to investigate the drivers of caracal habitat selection on a non-lethal farm in the Karoo; and finally, to evaluate caracal activity patterns on the non-lethal farm.

Coexisting ungulate-eating carnivores--lion, spotted hyena, cheetah, leopard, African wild dog, black-backed jackal, common jackal, and six species of vulture--are examined in East Africa's Serengeti ecosystem. Niche similarities year-round, by season, and by location are described using food, habitat, time of hunting, and other variables. Intraspecific niches of cheetah sex, age, and social groups show that male coalitions differ most from the others in hunting behavior and habitat use. Tests using the carnivore data failed to support hypotheses about niche breadth variation, niche overlap variation, range of food items, and niche inclusion. Densities of the five largest Carnivora in the 35,500 square kilometer ecosystem are 0.513/sq km; including the two jackals gives a density of 1.55/sq km. Their prey--30 ungulate species--are 84 .85/sq km. Prey and predator ratios suggest that the Serengeti Plains in 1977 had a three-fourthsdecrease in relative abundance of prey to predators from wet season to dry season . Year-round the Ngorongoro Crater had a prey:predator ratio s lightly larger than that of the dry season Serengeti Plains. Literature review suggests that cleptoparasitism and direct killing are very important forms of interference competition among and within carnivore species . Evidence for exploitation competition i s scant, but is inferred because local environments are unpredictable f or carnivores. Analysis of body sizes fails to support the hypothesized ratios of 2.0 for body weigh t and 1.28 for linear dimensions . Three methods of calculating multidimensional niche metrics (product, summation, and projection) are compared. Apparently the Serengeti's carnivores coexist because of their behavioral flexibility in an unpredictable environment . Niche descriptions were of little he l p in assessing the foci for potential and real competition among carnivores . Only the direct observations of interference competition in long-term field studies identified where competitive interactions are occurring with sufficient intensity to provide a numerical response in a population. Spotted hyenas some times competitively exclude African wild dogs locally . Management f or a high abundance and diversity of carnivores probably requires maintaining high densities of prey and varied habitats. Specific recommendations are made for cheetah and African wild dog conservation.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第18842号 / 理博第4100号 / 新制||理||1589(附属図書館) / 31793 / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 幸島 司郎, 教授 伊谷 原一, 教授 村山 美穂 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DFAM

To develop effective conservation planning and mitigate biodiversity loss, standardized metrics for monitoring and assessing biodiversity are needed. This information is particularly vital in Botswana, where knowledge of many of the diverse wildlife populations is lacking. To address this knowledge gap, my dissertation research evaluated the distributions, densities, and ecology of the wildlife community in northern Botswana, with a focus on terrestrial carnivore species given their importance both ecologically and economically. My objectives were threefold: 1) estimate the distributions of the mammal community (n = 44 species) and evaluate community, group, and species-specific responses to anthropogenic and environmental variables, 2) test whether the presence of intraguild species or resource availability had a larger influence on the seasonal distributions of carnivore species, and 3) simultaneously estimate the population densities of 7 carnivore species. To accomplish these objectives, I completed a multi-year camera trap survey in a 1,154-km2 study area and analyzed the data using occupancy models (single and multi- species) and spatially explicit capture-recapture models. Estimates of species richness ranged from 8 to 27 unique species, species had a mean occurrence probability of 0.32 (95% credible interval = 0.21–0.45), and estimated densities ranged from 1.8 aardwolves (Proteles cristata) to 12.7 spotted hyenas (Crocuta crocuta) per 100 km2. The occupancy and richness of terrestrial mammals was negatively associated with human disturbance and in general, positively associated with open grasslands/floodplains. Carnivore species, specifically, tended to have greater species richness and larger population densities in open habitats than in closed. I also found carnivore distributions were positively associated with the detection rate of intraguild competitors and predators, suggesting competitor/predator avoidance did not play a large role in shaping carnivore community structure. My research highlights the pivotal role protected areas and grasslands play in conserving wildlife populations in northern Botswana. Additionally, my research helps progress camera trap analyses from single to multi-species assessments. Broader application of this multi-species approach would likely result in a better understanding of wildlife and carnivore communities which in turn, may help inform management actions aimed at addressing the loss of wildlife populations globally. / Ph. D.

Cougars (Puma concolor) are a growing concern for managers of Pacific Rim National Park Reserve and Clayoquot Sound UNESCO Biosphere Reserve on the west coast of Vancouver Island, British Columbia. Since the mid-1990s, the frequency and intensity of human-cougar interactions have dramatically increased. Concurrently, these areas have become increasingly popular for human activities. The primary goal of my study was to recommend ways to reduce the potential risk of human-cougar interactions to ensure long-term conservation of cougars while minimizing risks to visitor safety. To achieve this goal, I examined the use of two noninvasive approaches. First, during 2005-2006, I compared the rate of detection, cost and time required for a detector dog, sign surveys, scented rub pads and remotely triggered cameras to detect cougars in coastal temperate rainforests. Sign surveys were the most effective method due to the availability of good tracking substrate throughout the study areas. Cameras were also practical because they could be used by less skilled personnel and had the capacity to detect several species of wildlife. Second, I demonstrated the utility of pre-existing data by analysing the spatiotemporal trends of human-cougar interactions on the West Coast Trail from 1993-2006. My results showed a moderate increase of reported human-cougar interactions (n = 157) despite a steady decline in hiker numbers across these years. I identified four areas where activities of people and cougars repeatedly overlapped (hotspots). In general, interaction locations were primarily associated with high human activity: near campsites and landscape characteristics that were associated with campsites (i.e., beaches and freshwater drainages >20 m wide). However, the distribution of hotspots suggests that the co-occurrence of human-use areas (e.g., campsites) and important travel routes (e.g., freshwater drainages and logging roads) used by cougars may increase the likelihood of interactions. These findings will allow protected area managers to proactively mitigate human-cougar conflict through visitor education and protocols that reduce people and cougars from intersecting in space and time.

Les canidés et félidés peuvent être hôtes définitifs d’Echinococcus multilocularis, Toxoplasma gondii ou Toxocara spp., parasites responsables de zoonoses transmissibles par l’alimentation. La consommation crue de fruits et légumes porteurs de leurs œufs ou oocystes peut être source de contamination humaine. Cette étude visait à évaluer et caractériser le risque d’exposition humaine lié au dépôt de fèces de chats, chiens et renards dans les terrains potagers localisés en régions d’endémie. Ce dépôt s’est avéré important dans certains potagers des Ardennes. De plus, l’ADN d’E. multilocularis et Toxocara spp. a été détecté dans 1/3 des fèces collectées et 23 % des rongeurs piégés autours des potagers ont été trouvés infectés par au moins un des parasites d’intérêt, confirmant le risque d’exposition des hôtes intermédiaires. Parallèlement, l’identification précise des facteurs responsables du dépôt de fèces de carnivores a été conduite sur 192 potagers familiaux ou professionnels des Ardennes et de la Moselle. Au total, 1016 fèces de carnivores (59% de chats, 31% de renards et 10% de chiens) ont été collectées au cours de huit sessions de prospection. Par modélisation, nous avons montré que la présence d’une clôture limite très efficacement le dépôt de fèces de renard, tandis que la présence de rongeurs ou d’arbres fruitiers à proximité le favorise. Enfin, la mise au point d’une méthode sensible a permis la détection de l’ADN d’E. multilocularis et Toxocara spp dans 42 % et 12 % des terrains potagers. Au final, l’exposition humaine aux parasites étudiés semble élevée dans certains potagers. Des mesures de prévention basées sur les résultats de l’étude sont proposées. / Canids and Felids can be definitive hosts of Echinococcus multilocularis, Toxoplasma gondii and Toxocara spp., which are food-borne parasites responsible of zoonoses. The consumption of raw fruit or vegetables carrying their eggs or oocysts can be source of human contamination. This study aimed to assess and characterize the risk of human exposure linked to the faecal deposition by cats, red foxes and dogs in kitchen gardens located in endemic areas. This deposit was found to be important in some kitchen gardens located in the Ardennes region. Furthermore, DNA of E. multilocularis and Toxocara spp were detected in 1/3 of the collected faeces and 23% of the rodents trapped in kitchen gardens proximity were found infected with at least one of the canids or felids parasites, confirming the risk of intermediate host exposure. Concurrently, the accurate identification of factors responsible for carnivore faeces deposit was conducted from eight prospection sessions of 192 kitchen gardens, family or professional ones, located in the Ardennes and Moselle regions. A total of 1016 carnivore faeces (59% from cats, 31% from foxes and 10% from dogs) were collected. By using models to test the effect of various variables on faeces deposit, we showed that fencing efficiently limits fox faeces deposit whereas presence of rodents or fruit trees in the vicinity increases it. Finally, thanks to the development of a sensitive method, E. multilocularis and Toxocara spp. DNA was detected in 42% and 12% of the kitchen gardens. In conclusion, the human exposure to canids and felids foodborne parasites seems high in certain kitchen gardens. Prevention methods are proposed based on our results.

Le maintien des populations de grands carnivores est un élément clé dans le maintien des écosystèmes. L'étude de la dynamique des populations permet de comprendre et d'anticiper les fluctuations en taille de population et est donc un outil essentiel dans la gestion et la conservation des espèces. En Suède, la population d'ours bruns (Ursus arctos) a connu une forte croissance depuis le début du 20e siècle. Conséquemment, les quotas de chasse ont augmenté de façon fulgurante depuis le début des années 2000. Or, les effets de la chasse sur la dynamique de la population restent encore méconnus et donc, les conséquences de cette augmentation sur le taux de croissance de la population sont inconnues. La chasse devrait avoir des impacts particulièrement importants dans cette population, puisque plusieurs études suggèrent qu'une augmentation de la chasse mènerait non seulement à une augmentation de la mortalité des classes d'âge exploitées, mais aussi à une augmentation de la mortalité des oursons à travers une augmentation de l'infanticide sexuellement sélectionné (où les mâles tuent les jeunes qui ne sont pas les leurs pendant la saison de reproduction pour augmenter leurs opportunités de reproduction). Mon projet avait donc pour but de quantifier les effets directs ainsi que l'effet potentiellement indirect de la chasse à travers l'infanticide sexuellement sélectionné sur la dynamique de la population d'ours brun en Suède. Les objectifs spécifiques de mon projet étaient de : i) quantifier comment les taux démographiques et le taux de croissance de la population varient sous faible et forte pression de chasse; et de ii) déterminer l'importance des différents taux démographiques, dont la survie des oursons, sur le taux de croissance de la population. Pour ce faire, j’ai collaboré au suivi individuel à long terme d’ours bruns dans le centre-sud de la Suède effectué par le Scandinavian Brown Bear Research Project. À l'aide de ces données, j'ai construit des modèles matriciels de la population qui permettent de calculer son taux de croissance ainsi que l'importance relative des différents taux démographiques pour la croissance de la population. J'ai élaboré un modèle pour l'ensemble de ma période d'étude (1990-2011) ainsi que des modèles distincts pour deux périodes qui diffèrent par la pression de chasse : de 1990 à 2005 avec une pression de chasse relativement faible, et de 2006 à 2011 avec une forte pression de chasse. J'ai trouvé que la grande majorité des taux démographiques étaient plus faibles sous une forte pression de chasse : ceci se reflétait par un taux de croissance asymptotique légèrement en deçà de 1, indiquant ainsi un déclin de la population. La survie des oursons était plus faible sous forte pression de chasse, probablement en raison d'une augmentation de l'infanticide sexuellement sélectionné. D'ailleurs, la survie des oursons avait une influence relativement forte sur le taux de croissance de la population. Finalement, j'ai aussi pu quantifier l'impact de l'infanticide sexuellement sélectionné sur le taux de croissance de la population. J'ai trouvé que le comportement était très important, puisque s’il n'y avait pas d'infanticide sexuellement sélectionné dans la population, celle-ci aurait été en croissance même sous forte pression de chasse. Ainsi, mon projet de recherche a mis en évidence l'importance du comportement individuel et des effets indirects de la chasse pour la dynamique d'une population sauvage. Les effets indirects de l'exploitation sont bien souvent méconnus et doivent être étudiés davantage afin que nous comprenions l'impact réel sur les populations exploitées.

Recent studies have shown that neuronal connections occur between primary auditory and visual cortices of the primate (Falchier et al., 2002; Rockland and Ojima, 2003), and it has been suggested that these projections are involved in multisensory processing in these lower-level, core areas of cortex. The present study was conducted to determine if similar connections occur in other higher mammals such as carnivores (ferrets; Mustela putorius). Large injections of sensitive neuroanatomical tracer were placed within the core areas of auditory cortex in 3 ferrets. After transport and processing, labeled axon terminals were found not in primary visual cortex, but in area 19, or V3. Injection of tracer into V3 of 3 additional ferrets produced retrogradely labeled neurons not in the core region of auditory cortex, but along its posterior borders. These data indicate that cross-modal connections occur in the ferret cortex, but do not support the notion that they exist between the primary representations of the different sensory modalities.

A integração morfológica determina a quantidade de variação disponível em diferentes direções do espaço multivariado entre um conjunto de caracteres, e pode assim ter importantes consequências evolutivas, enviesando a direção e a taxa de evolução morfológica em determinadas direções. Assim, estudar a evolução da integração morfológica é importante para entender a evolução fenotípica de uma linhagem. Neste trabalho, utilizo abordagens de genética quantitativa comparativa para estudar a evolução da integração morfológica no crânio em carnívoros da subordem Feliformia, grupo com extensa diversidade filogenética, morfológica e ecológica para o qual análises anteriores de integração analisaram número reduzido de espécies. Para este fim, comparo matrizes de covariância fenotípicas intra-populacionais quantificando a associação entre 35 distâncias lineares no crânio de um conjunto de táxons incluindo todas as sete famílias e mais de um terço da diversidade filogenética atual da subordem. Investigo também a associação entre integração morfológica e a evolução da morfologia média, assim como a influência da história evolutiva (filogenia) e de dois aspectos da ecologia (dieta e socialidade) sobre a evolução da morfologia média e da integração. Os padrões de integração fenotípica no crânio (i.e. quais caracteres variam em conjunto com maior intensidade) são bastante estáveis na subordem, como observado para outros mamíferos, mas revelam alguma sub-estruturação, com padrões um pouco distintos em Felidae, Hyaenidae e Prionodontidae As magnitudes de integração são também relativamente estruturadas, com valores geralmente mais altos em Felidae e consistentemente mais baixos em Hyaenidae. Apesar disso, a magnitude geral de integração é evolutivamente plástica, como em outros mamíferos, havendo importantes variações internas a cada família. Encontrei uma associação geral entre a evolução da morfologia média e modificações em padrões e magnitudes de integração para a subordem como um todo e em vários grupos menos inclusivos, o que em alguns casos se manifesta como uma semelhança da integração entre táxons morfologicamente convergentes, mas filogeneticamente distantes. A evolução de padrões de integração se correlaciona à história filogenética em Feliformia como um todo e na maior parte dos grupos acima, mas não abaixo do nível de família, corroborando padrões de covariação relativamente distintos apesar de sua alta similaridade, bem como integração mais restrita internamente a cada família. A dieta está associada à filogenia, e correlaciona-se à evolução da morfologia média e dos padrões de integração na subordem como um todo antes, mas não após correção levando em conta a proximidade filogenética, indicando que alterações na dieta são parte da história evolutiva compartilhada que levou às diferenças em morfologia média e integração existentes no grupo. Os caracteres cujos padrões de covariação estão mais associados à dieta na subordem incluem partes das regiões oral e zigomática, além de algumas partes do neurocrânio. A socialidade está relacionada à evolução da morfologia craniana média, mas não aos padrões de integração gerais em Feliformia, apesar de existirem associações entre a socialidade e os padrões de resposta a seleção de alguns caracteres específicos, incluindo partes do neurocrânio e distâncias ligadas ao arco zigomático. Os mangustos (família Herpestidae) mostraram influência significativa tanto da dieta quanto da socialidade na evolução de sua morfologia craniana média e de seus padrões de integração, ressaltando a importância desses dois aspectos ecológicos na evolução fenotípica do grupo e mostrando, pela primeira vez, uma associação direta entre a integração no crânio e o comportamento social, possivelmente intermediada por aspectos relacionados ao desenvolvimento do cérebro. O presente trabalho sugere, assim, uma complexa interação de fatores afetando a evolução da morfologia craniana em Feliformia, e indica direções para trabalhos futuros, incluindo análises dos padrões de modularidade craniana no grupo e, possivelmente, análises comparativas em diferentes morfotipos hipercarnívoros abordando a morfologia média e a integração morfológica no crânio, bem como a dieta e possivelmente outros fatores / Morphological integration determines the standing variation available within a lineage in different directions of the multivariate space of a trait set, and can therefore have important evolutionary consequences, biasing the direction and rate of morphological evolution in certain directions. Consequently, studying the evolution of morphological integration is important to understand a lineage\'s phenotypic evolution. Here I employ a comparative quantitative genetic framework to study the evolution of morphological integration in the skull for the suborder Feliformia (Carnivora), a group with remarkable phylogenetic, morphological and ecological diversity for which previous analysis of integration included a limited number of species. To this purpose, I compare within-population phenotypic covariance matrices quantifying the associations between 35 skull linear measurements in a set of taxa including all seven feliform families and over one third of the suborder\'s extant phylogenetic diversity. I also assess the association between morphological integration and the evolution of average skull morphology, as well as the influence of evolutionary history (phylogeny) and two ecological aspects (diet and sociality) on the evolution of average morphology and integration. Skull integration patterns (i.e. which characters covary the most) are quite stable in the suborder, as seen among other mammals, but also have some structuring between groups, with relatively distinct patterns in Felidae, Hyaenidae and Prionodontidae. Integration magnitudes are also relatively structured, with generally higher values in Felidae and consistently lower values in Hyaenidae. Nevertheless, the overall magnitude of integration is evolutionarily labile, as in other mammals, with important variations occurring within each family. I found a general association between the evolution of average morphology and changes in integration patterns and magnitudes for the suborder as a whole and also for many less inclusive groups, which in some cases leads to similar integration between morphologically convergent, but phylogenetically distant taxa. The evolution of integration patterns is correlated to phylogenetic history in Feliformia as a whole and in most groups above, but none below the family level, corroborating relatively distinct covariation patterns in spite of their high similarity, and also a greater constraint of integration within each family. Diet is associated to phylogeny, and correlates to the evolution of average morphology and integration patterns in the suborder before, but not after correction for phylogenetic relatedness, indicating that dietary changes are part of the shared evolutionary history which shaped the differences in average morphology and integration existing in the group. The characters with covariation patterns more closely associated to diet include parts of the oral and zygomatic skull regions, as well as parts of the neurocranium. Sociality is related to the evolution of average skull morphology, but not to general integration patterns in Feliformia as a whole, although there are correlations between sociality and the selection response patterns of some specific characters, including parts of the neurocranium and measurements related to the zygomatic arch. Mongooses (family Herpestidae) had the evolution of their average skull morphology and integration patterns significantly influenced by both diet and sociality, stressing the importance of these two ecological factors for phenotypic evolution of this group and showing for the first time a direct association between morphological integration in the skull and social behaviour, possibly mediated by aspects related to brain development. The present work thus suggests a complex interaction of factors affecting the evolution of skull morphology in Feliformia, and opens up directions for future work, including analyses of skull modularity patterns in this group and, possibly, comparative analyses of different hypercarnivore morphotypes addressing average skull morphology and morphological integration, as well as diet and possibly other factors

Abstract Illegal killing of wolves (hereinafter ’wolf poaching’) in human-inhabited areas where wolves are also legally harvested is a special case of wildlife crime. This doctoral thesis examines wolf poaching in Finland and Sweden from the ecological perspective. In the first paper, we examined the causes of mortality among collared Finnish wolves and the role of estimated poaching rates on population changes. The second paper related the likelihood of being poached to covariates expressing different dimensions of the wolf conflict at two spatial scales (territory and country level) in Finland. Third paper turns the focus to Sweden, where we examined the disappearances of adult wolves in relation to population size, legal harvest and inbreeding. The first two studies were done in collaboration with the Natural Resources Institute Finland (Luke) and the third with the Scandinavian wolf research project SKANDULV. Poaching outnumbered other causes of death. Most poaching cases were unverified. Other causes of death were legal harvest, traffic and natural mortalities. Both populations had a relatively high number of wolves with unknown fates. Inbreeding was not related to the disappearances of adult wolves in Sweden. Remoteness to human inhabitation and the detectability of the wolves from the forest roads (road crossings by wolves) increased the likelihood of poaching in Finland. Adult wolves suffered high risk of poaching in both populations. Risk was highest in early spring in Finland. Larger population size increased and the number of legally harvested wolves decreased poaching in both countries. Poaching seemed to limit the study populations despite the management efforts that used legal hunting as a tool to increase tolerance towards wolves. / Tiivistelmä Suden ja ihmisen rinnakkainelo johtaa konflikteihin, joiden lievittämiseksi susikantoja rajoitetaan usein luvallisella metsästyksellä. Suteen kohdistuu lisäksi laitonta tappamista eli salametsästystä. Tämä väitöstutkimus selvittää salametsästyksen ekologisia vaikutuksia Suomen ja Ruotsin susipopulaatioihin. Ensimmäisessä osatyössä selvitimme suomalaisten pantasusien kuolinsyitä, arvioimme salametsästyksen voimakkuutta ja sen vaikutuksia populaatiotasolla. Toisessa osatyössä tarkastelimme susikonfliktiin liittyvien ennustetekijöiden vaikutusta laittomasti tapetuksi tulemisen riskiin reviiritasolla ja koko Suomen mittakaavassa. Kaksi ensimmäistä osatyötä tehtiin Luonnonvarakeskuksen (Luke) suurpetotutkimuksessa. Kolmas osatyö tehtiin osana skandinaavista susitutkimushanketta (SKANDULV). Siinä selvitimme populaatiokoon, luvallisen pyynnin ja sukusiittoisuuden vaikutuksia aikuisten susien katoamiseen Ruotsissa. Salametsästys oli susien yleisin kuolinsyy, mutta suurin osa tapauksista jää toteen näyttämättä. Muita kuolinsyitä olivat luvallinen metsästys, liikenne ja luonnolliset kuolinsyyt. Aineistoissa oli runsaasti kohtaloltaan tuntemattomaksi jääneitä yksilöitä. Yksilöiden sukusiittoisuusaste ei ollut yhteydessä susien katoamisiin Ruotsissa. Syrjäinen sijainti ja susien havaittavuus metsätiestöltä lisäsivät laittoman tapon todennäköisyyttä Suomessa. Salametsästysriski oli korkein kevättalvella. Aikuisilla susilla oli huomattavan korkea riski tulla laittomasti tapetuksi. Tutkimuksen perusteella salametsästyksen määrää selittävät erityisesti susipopulaation kulloinenkin koko ja luvalliset pyyntimäärät. Suurempi susikanta lisäsi salametsästystä ja metsästyslupien määrä vähensi sen riskiä. Salametsästys vaikuttaa säädelleen susikantoja siitä huolimatta, että susikonfliktia on pyritty lieventämään luvallisella metsästyksellä.