Understanding the unequal distribution of life on earth is a fundamental goal of ecology and evolutionary biology. Past efforts to explain large-scale patterns in diversity have tended to focus on two broad classes of explanation, one invoking the importance of abiotic factors (i.e. climate and vegetation) and the other biotic (i.e. competition); but neither has proven entirely adequate. Parasites are a major but poorly understood component of life that may offer some answers. Yet despite widespread theoretical support and some empirical evidence, the role of parasites in explaining patterns in the diversity, distribution, and abundance of species remains largely untested in natural communities. In this thesis I use a mega-diverse elevation gradient of birds as a model system to study the role of avian malaria in explaining these macroecological patterns. In the first data chapter I tested the extent to which patterns of infection across species is predictable. I found that the effects of host ecology and environment were weakly related to infection prevalence and were not consistent across different malaria lineages. Instead, I show that hosts coexisting with many close phylogenetic relatives consistently experience higher infection than evolutionarily distinct host species. In the second chapter I tested if parasite sharing may help explain these observed relationships and show that parasite sharing among host pairs declines with the time since divergence. Spatial contiguity between host pairs was also positively associated with parasite sharing. In the third chapter I tested how infection prevalence varies across species ranges in accordance with expected variation in host abundance. I show that birds are more likely to be infected at the centre of their elevation range, where host abundance is expected to be highest. Intriguingly, I also found that the incidence of host infection is unrelated to the position within the geographic range of the parasite. In the fourth data chapter, I tested whether parasites may regulate diversity by limiting geographic ranges of their hosts through ‘apparent competition’ in which a non-lethal parasite in a primary host, may be lethal in a secondary host. In support of this, I found that more observed bird ranges end at parasite infection zones than would be expected by chance. Taken together, my results suggest that parasites may play a major role in shaping patterns in the distribution and diversity of species, over both ecological and evolutionary scales. This is likely to arise and be maintained by host parasite interactions in which distantly related hosts are less likely to be infected by local parasites than close relatives, thus promoting the build up of diversity locally. On the basis of my analyses, I conclude that across montane elevation gradients in birds, and across diversity gradients more generally, parasites are likely to play a crucial role in the origin and maintenance of high biological diversity.

Understanding the impacts of land-use change on biodiversity and the ecosystem services that it provides is of great importance given unprecedented growth of the human population. Past studies attempting to explore these impacts have described the overall structure of communities (i.e. species richness and trait diversity) across gradients of local scale degradation and fragmentation, and have sought to identify whether the loss of species following land-use change is non-random. Yet, despite a wealth of research we still lack a generalised understanding of how land-use change impacts on traits responsible for determining species sensitivity and their role within ecosystems, particularly for vertebrates. Moreover, despite the importance of niche-based processes in the assembly of communities, we have not yet elucidated whether these are important in mediating the collapse of communities in human-dominated landscapes. To fill these existing research gaps, I collated comprehensive avian species inventories from fragmented and degraded forests and compared their structure with communities existing in continuous forests. In Chapter 2, I tested whether sensitivity of species to forest fragmentation varies between the temperate zone and the tropics and whether there are key differences in the size of fragments required to maintain ecosystem processes in these regions. I found that sensitivity to fragmentation varies according to functional group and body mass, with the prevalence of insectivores and large frugivores declining in relation to fragment size, particularly in tropical fragments smaller than 100 ha. In Chapter 3, I tested whether functional diversity and the mean position of trait diversity of insectivores and frugivores, changed across a gradient of intensifying land-use change. I found a decline in the functional diversity of forest species and a shift in the mean community traits for both forest and non-forest species. In Chapter 4, I tested whether the structure of tropical bird communities are influenced by species interactions in a fragmented landscape. I found increasing over-dispersion in functional and phylogenetic trait relatedness among species with decreasing fragment size, suggesting that competitive interactions are important in the disassembly of avian communities. In Chapter 5, I modelled the impact of forest cover change on ecosystem function across the Brazilian Amazon, focusing on seed dispersal by birds. Furthermore, I tested whether ecosystem function declined linearly with decreased forest cover after accounting for differences in the underlying pools of species. I found the lowest levels of functional diversity along the southern arc of deforestation and that the dispersal of large seeds showed some resilience to declining forest cover. Taken together, my results suggest that the loss of species from communities in degraded and fragmented landscapes is strongly non-random. Insectivores and large frugivores are most sensitive to land-use change, with species located in the densest parts of trait space being most threatened by a decline in forest patch size, suggesting that species interactions regulate the collapse of avian diversity in human-modified forests. I conclude that land-use change has important implications for the provisioning of ecosystem services, including seed dispersal and the control of insect herbivores. The impact of future land-use change is likely to be mediated by the composition of the original pool of species and the amount of redundancy in the ecosystem services that they provide. I discuss the relevance of my findings to land-use management strategies and policy interventions, and in particular conclude that these should, where possible, maintain pristine forest patches above 1000 ha, improve connectivity among habitat patches, and ensure greater protection for logged and burnt forests. Future studies should focus on clarifying the link between shifts in vertebrate community structure and the functioning of forest ecosystems.

The present thesis explored ethological and geographical approaches for the investigation of vector-borne parasites. In the first part, the role of associative learning on vector preferences for hosts was investigated through a comprehensive series of behavioural experiments using the vector of dengue and yellow fever diseases, the mosquito Aedes aegypti. To this end, the possibility that the mosquitoes were able to associate unconditional stimuli with particular odours and visual patterns to which they were responsive was explored, but no evidence supporting the hypothesis that associative learning abilities are present in adults of this species was found. A critical review of the literature on learning in mosquitoes conducted afterward allowed the reinterpretation of findings in the field, narrowing the scope of evidence suggesting the existence of these cognitive abilities in some species. In the second part of the thesis, the distribution and evolution of mosquito-borne viruses was investigated with the use of geo-coded environmental data and spatial statistics. Initially, the eco-climates associated with the distribution of Japanese encephalitis virus were described and modelled, allowing the production of a worldwide predictive map defining the probability of each region to develop this disease in the future. Predominating amongst those areas shown to be under high risk were the equatorial regions of South America and Africa. The methodology used to infer such patterns – non-linear discriminant analysis – was subsequently explored with a number of simulations. Overall, differences in the choice of parameters required for the analysis were shown to lead to differences in the final outputs produced, basically in those cases where the environmental range for which predictions are generated is not rigorously limited. Finally, eco-climate surrogates for the evolution of the Japanese encephalitis serocomplex were investigated, but the current environmental distances between the viruses did not seem to be associated with the events leading to their speciation.

Baseline data on a species' distribution and abundance are essential for developing practical conservation management plans. Such data are difficult to obtain for many low density cryptic carnivores. The Scottish wildcat, Felis silvestris silvestris, is no exception with <400 individuals thought to remain. Its conservation has been further complicated by extensive hybridisation and introgression with the domestic cat (F.s.catus). Hybridisation has also resulted in difficulties in discriminating between wildcats, wildcat x domestic hybrids (hybrids) and tabby coloured feral domestic cats. This has inhibited survey efforts, leading to a lack of general ecological information. Using the most recent identification tools available, extensive surveys using various methods including camera trapping were carried out across Northern Scotland in order to examine the current status of the Scottish wildcat. Current distribution indicates a more restricted range than recent studies. Wildcats are at risk of hybridisation from feral domestic cats and in particular, hybrids, throughout their current probable range. The distribution of hybrids overlaps with both feral domestic cats and wildcats, pointing to a significant threat from hybrids acting as a bridge between wildcats and feral cats. Mean density estimates of 3.5 (SD=0.7) wildcats/100 km² were comparable with those from other studies in Scotland using different survey methods. Total population size estimates ranged between 115-314 individuals depending on local densities and home range size. Population viability analysis (PVA) indicated the current population is not viable unless management actions are undertaken in the near future (Mean time to extinction = 48.2 years (SD = 9.39), probability of extinction=1, SE = 0), and that reducing mortality rates and/or supplementing populations from captive bred cats are likely to be necessary to achieve viability. Based on these data, the Scottish wildcat may be more endangered than many other species classified as Endangered and the current status of the Scottish wildcat should be reviewed.

This project used a variety of genetic markers to investigate the evolution and population genetics of hydrothermal vent fauna that were recovered from the Scotia Sea, in the Atlantic sector of the Southern Ocean. The origins of one of these species, an undescribed species of Kiwa sp. found on the East Scotia Ridge (ESR) and its constituent family Kiwaidae, a group of vent and seep-associated decapod squat lobsters (infraorder Anomura) was investigated using a concatenated nine-gene dataset and key divergences were dated using fossil calibrations. These results confirm earlier research showing Kiwaidae reside in the superfamily Chirostyloidea, but form a monophyletic clade with the non-chemosynthetic family Chirostylidae and not Eumunididae. Chirostyloid families diverged in the Cretaceous, although extant Kiwaidae radiated in the Eocene, consistent with many other chemosynthetic taxa that appear recently derived. The basal tree position of Pacific species (and the Alaska location of a likely stem-lineage kiwaid fossil) suggests kiwaids originated in the East Pacific. Within a Southern Hemisphere clade, the divergence between the southeastern Pacific K. hirsuta and a non-Pacific lineage (Kiwa sp. ESR and Southwest Indian Ridge kiwaids) is no earlier than 25.9 Ma, consistent with a spread from the Pacific into the Scotia Sea and beyond via now-extinct active ridge connections or mediated by a Miocene onset of the Antarctic Circumpolar Current (ACC) through a newly-opened Drake Passage. This project also investigated the population genetics of three undescribed species found at two vent fields ~ 440 km apart at either end of the ESR: Kiwa sp., a peltospirid gastropod and Lepetodrilus sp. limpets. Lepetodrilus sp. was also found at the Kemp Caldera, a submerged part of the South Sandwich Islands (SSI). Analyses of cytochrome c oxidase subunit 1 (COI) as well as microsatellite loci developed from Roche 454 sequence libraries revealed no differentiation along the ESR for all three species consistent with panmixia, or the dominance of non-equilibrium processes between vent field colonies within a metapopulation, possibly enhanced further by cold-induced arrested larval development. Despite apparent connectivity along the ESR, both COI and microsatellites revealed differentiation between ESR limpets and Kemp Caldera limpets ~ 95 km to the east, possibly owing to the hydrographic isolation of the caldera. Both COI and microsatellite diversity patterns were consistent with recent (< 1 Ma) demographic expansions for all three species (although the influence of selection sweeps on COI cannot be discounted); a pattern observed worldwide at vent communities and may reflect demographic instability over time as a consequence of the stochastic birth and death of vent colonies within a metapopulation. Different COI bottleneck ages between the three species (excluding the influence of possible selection) as well as the absence of kiwaids and peltospirids at Kemp, have been attributed to differences in life history, in particular larval morphology and presumed dispersal strategy. These results highlight the role of larval dispersal of vent fauna along active spreading ridges, both in maintaining vent metapopulations across vent colonies prone to stochastic birth and extinction in the short term, but also in the spread of taxa globally and the formation of biogeographic provinces. The likelihood that the three species presented here exist at vents east of the ESR and SSI, prompts further exploration along ridges in the South Atlantic, in order to investigate the effect of the ACC in enhancing gene flow and delineating biogeographic provinces.

Social behaviour is shaped by complex relationships between evolutionary and ecological processes interacting at different scales. Benefits gained from social associations can range from predator dilution to collective sensing, but little is known about how these can be influenced by social structure and phenotypic composition. In this thesis, I investigated how individual decision-making affects phenotypic social structure, and how this mediates social behaviour through emergent properties of collective group behaviour. First, using mixed-species flocks as a model system, I showed individual tits (Paridae, chapter 2) and thornbills (Acanthizae, chapter 3) varied significantly in their social positions. Within-species variation in network position was as large as between-species variation, sug- gesting that prescribing functional roles at the species level may not sufficiently account for potential differences in fitness operating at the individual level. Rather, this suggested that structure may be driven by phenotypic traits, underpinning network structure (chapter 4). Next, I used an extensive data set of foraging records to explore factors determining the composition, of flocks of great tits (Parus major, chapter 5). For example, assortment by dispersal phenotype (immigration status) was the result of spatial disaggregation, and I showed that this may facilitate social selection for breeding territories (chapter 6). Finally, I investigated how decision-making shaped mixed-species social structure. I found that tits used a common strategy for managing pressures of predation and starvation by shifting from exploration to exploitation at different times of the day (chapter 7). I then found that a very simple interaction rule successfully replicated mixed-species group structure (chapter 8). Strikingly, the same rule was applied to both conspecifics and het- erospecifics, potentially playing an important role in the maintenance of flock structure. Through experimental manipulation of ecological conditions, I found that heightened per- ceived predation resulted in stronger social attraction overall, whereas increased competition led to a reduction in attraction to conspecifics (chapter 9). Simulations suggested this could be one potential mechanism underpinning fission-fusion dynamics in these species. Together, the results in this thesis form a framework linking social behaviour to individ- ual fitness where natural selection is shaped by the social environment. This approach may prove useful for testing whether following common social rules reduces variance in benefits accrued by individuals, and how within-species variation in social behaviour can impact emergent properties of groups.

The Effects of Tropical Forest Management on Biodiversity and Ecosystem Functioning Eleanor M. Slade 1. Between 35 % and 50 % of all closed-canopy tropical forest has been lost, and the rate of deforestation continues to increase throughout the tropics. Despite a wealth of literature on the effects of tropical forest disturbance on the diversity and composition of a variety of taxa, there is still no clear consensus on the value of disturbed forests for biodiversity. 2. If forest management practises are to be sustainable in the long-term they should maintain both biodiversity and ecosystem functioning (the interactions and processes of the ecosystem), as well as a timber harvest. However, few studies have investigated the extent to which ecosystem functioning is reduced in logged forests. The effects of different logging intensities on a variety of taxa, and the ecosystem processes with which they are associated, were assessed in the Danum Valley Conservation Area in Sabah (Malaysian Borneo). 3. Even under high logging intensities, the forests of Sabah appear to have been managed in a way that maintains timber yields in the short-term. However, other aspects of forest structure had been affected, which could have important consequences ecologically, and for the long-term sustainability of timber harvests. 4. Combining field studies with manipulative experiments allows assessment of the impacts of species changes associated with habitat modification on measures of ecosystem functioning. Dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) functional group richness and composition were manipulated in a series of field experiments. Certain functional groups and species were found to have a greater impact on ecosystem functioning than others; nevertheless a full complement of species was needed to maintain full ecosystem functioning. 5. Dung beetles appeared to be relatively robust to changes in forest structure associated with selective logging, but species richness was reduced with high-intensity logging. There was a corresponding decrease in ecosystem functioning (dung and seed removal) with a decrease in species richness, and a decrease in the biomass of large nocturnal tunnellers, suggesting that although some species are dominant, rare species are also needed to preserve full ecosystem functioning. 6. A complex interaction between birds and ants resulted in reduced herbivory of seedlings of the important timber tree, Parashorea malaanonan, in some instances. However, this interaction was not affected by either selective or high intensity logging. Seedfall of P. malaanonan, was reduced in logged forest compared to primary forest. Despite insect seed predation being higher in primary forest, there was still successful recruitment during a non-mast year. Parasitism of insect-predated seeds was found to be inversely density dependent, and was higher in logged forest where seed predation was lower. 7. The results of this thesis suggest that the forests of Sabah appear to be being logged under a management system that is compatible with sustainable timber management, but not necessarily sustainable forest management. Low intensity selective logging seems to preserve much of the original forest structure, biodiversity and ecosystem functioning compared to logging at higher intensities. However, ecosystem processes were variable in their response to logging, suggesting that management decisions should be based on the consideration of multiple taxa and processes.

The evolutionary processes that shape individual variation in continuous behavioural traits remain poorly understood. While the emerging discipline of animal personality is providing increasing evidence that consistent individual differences in behaviour have significant fitness consequences, cognitive traits are yet to be explored in the same manner. My general objective in this thesis was twofold. First, I aimed to examine the ecological significance and fitness consequences of the cognitive trait innovative problem solving-performance, using a population of great tits (Parus major). Second, I aimed to explore the mechanisms underlying the functional significance of ‘exploration behaviour’ a captive measure of the reactive- proactive personality axis, focusing specifically on foraging and risk-taking behaviour. This two-trait approach was expected to shed light on whether personality and cognition simultaneously influence fundamental behaviours. By carrying out behavioural assays on birds temporarily held in captivity, I showed that success at solving a food-motivated problem was repeatable within individuals, consistent between two different tasks and independent of exploration behaviour. Problem-solving performance was positively related to clutch size and fledgling number, established when birds were released back into the wild. Furthermore, when rearing offspring, solvers had shorter working day lengths than non-solvers and foraged over smaller ranges without compromising either provisioning quantity or quality. However, solver females were also more likely to desert their broods than non-solvers and consequently there was little evidence to suggest that directional selection acted on problem-solving performance. In comparison to non-solvers, solver males were also found to be poorer at competing for limited food resources during the winter. Together these findings suggest that costs and benefits are associated with problem-solving performance, which together may act to maintain variation in this trait. My thesis also provides some of the first evidence that exploration behaviour is related to both foraging and risk-taking behaviour in the wild. In comparison to relatively slow explorers, fast exploring males were better competitors at feeders during the winter. Relatively fast and slow explorers also differed in a number of foraging behaviours during offspring provisioning, although not always in the direction predicted from captive work on other populations. Finally, while exploration behaviour was positively correlated with risk- taking behaviour amongst breeding females during incubation, it did not predict nest desertion in response to the risk associated with being trapped by field workers. Collectively these findings suggest that personality measured in captivity has specific but not general power to predict behaviour in the wild. An important facet of evolution is that natural selection is rarely likely to act on a single trait in isolation owing to correlations between traits. This thesis demonstrates how important behaviours in the wild can be simultaneously linked to multiple sources of consistent behavioural variation. It also represents the first large-scale investigation of how variation in a cognitive trait relates to natural behaviour, reproductive life-history variation and fitness in wild animals. Using this individual-based approach in a natural setting may prove to be a useful tool for understanding how selection acts on cognitive traits.

Novel infectious diseases in humans are of great concern to public health authorities and researchers in epidemiology. Zoonotic pathogens in particular have the potential to cause epidemics without any or little warning. In this thesis, I investigate evolutionary and environmental conditions, and the interactions between both, which facilitate the zoonotic emergence of novel pathogens. I start with a list of the mechanisms and processes which might influence a zoonotic emergence, and identify some unsolved problems. I address these with multiple, theoretical models. First, I use a village-city model with different adaptation scenarios to examine the influence of spatial heterogeneity on the emergence process. I derive general analytical results for the statistical properties of emergence events, including the probability distribution of outbreak sizes. My results suggest that, for typical connection strengths between communities, spatial heterogeneity has only a weak effect on outbreak size distributions, and on the risk of emergence per introduction. Next, I extend the research on environmental conditions by looking at pathogen specialisation in multi-host systems. I derive threshold connectivities for which generalist pathogens, which infect multiple species and might therefore be more dangerous to cross into the human species, can sustain transmission and are not dominated by specialists, which can only cause sustained transmission chains in a single host species, but are able to cause emergences with little warning. My third research chapter is interested in the effect of the loss of biodiversity. I analytically derive expected prevalences for fast growing and slow growing species. If fast growing species tend to perform better in degraded environments, my analytical results suggest that the overall prevalence level of infectious diseases will rise as environments degrade, which facilitates the chance of zoonotic jumps. In my last research chapter, I examine the actual impact of a novel, emerging infectious disease. I use data from the recent `Swine flu' epidemic in England to estimate epidemiological parameters of the infectious agent. My results suggest that the majority of infected cases showed no or only mild symptoms. This reveals that more data than just the estimated number of cases are necessary to fully evaluate the danger of a possible zoonotic, emerging infectious disease. I conclude by discussing my results and the implications which these might have.

A number of emerging vector-borne diseases have the potential to be transmitted in the UK by native mosquitoes. Human infection by some of these diseases requires the presence of communities of multiple vector mosquito species. Mitigating the risk posed by these diseases requires an understanding of the spatial distributions of the UK mosquito fauna. Little empirical data is available from which to determine the distributions of mosquito species in the UK. Identifying areas at risk from mosquito-borne disease therefore requires statistical modelling to investigate and predict mosquito distributions. This thesis investigates the distributions of potential vector mosquitoes in the UK at landscape to national scales. A number of new methodological approaches for species distri- bution modelling are developed. These methods are then used to map and understand the distributions of mosquito communities with the potential to transmit diseases to humans. Chapter 2 reports the establishment of substantial populations of the West Nile virus (WNV) vector mosquito Culex modestus in wetlands in southern England. This represents a drastic shift in the species’ known range and an increase in the risk of WNV transmission where Cx. modestus is present. Chapter 3 develops and applies a new species interaction distribution model which identifies fish and ditch shrimp of the genus Palaemonetes as predators which may restrict the distribution of the potential WNV vector community in these wetlands. Chapter 4 develops a number of methods to make robust predictions of the probability of presence of a species from presence-only data, by eliciting and applying estimates of the species’ prevalence. Chapter 5 introduces a new Bayesian species distribution modelling approach which outperforms existing methods and has number of useful features for dealing with poor- quality data. Chapter 6 applies methods developed in the previous two chapters to produce the first high-resolution distribution maps of potential vector mosquitoes in the UK. These maps identify several wetland areas where vector communities exist which could maintain WNV transmission in birds and transmit it to humans. This thesis makes significant contributions to our understanding of the distributions of UK mosquito species. It also provides methods for species distribution modelling which could be widely applied in ecology and epidemiology.