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.

In Western Europe the incidence of tick-borne encephalitis (TBE) has increased over the last 30 years, coupled with changes in distribution. Modifications in the TBE enzootic cycle, through a combination of changes in temperature, vertebrate abundance and habitat suitability may have increased the risk of TBE in recent years. In Switzerland, analysis using satellite-derived climate data demonstrated that the environment of areas with TBE since the 1980s and areas that recently became endemic for TBE have become more similar between 2001 and 2009. This was coupled with an increase in April, May and June temperature, which could have affected the tick population and/or human exposure to ticks. Deer and boar abundance also changed in some cantons. In Germany, spatio-temporal modelling demonstrated the importance of temperature, vertebrate abundance and unemployment in the incidence and distribution of TBE between 2001 and 2009. Changes in TBE reporting, April, May and June temperature, vertebrate abundance and pesticide use may have contributed to increases in TBE in 1992 and 2001. Human exposure patterns, however, appear to be as important as the enzootic cycle in shaping the incidence of TBE, not only in determining the overall trend but also in interacting with the weekly, seasonal and yearly patterns of tick hazard to give the observed incidence. In Switzerland, in weeks with warm, sunny weather, human exposure to ticks is promoted and short-term increases in tick bites are seen. Human outdoor activity also shifts the seasonal pattern of tick bites, when compared with tick questing. There was no apparent increase in time spent in outdoor activities between the 1990s and 2000s in Italy, Germany and Austria, but survey data demonstrated that walking and hiking were already popular activities across Europe by the 1990s. The popularity of mushroom and berry foraging as a source of income in Latvia, Lithuania and Poland, coupled with the expense of vaccination, provide an inverse link between economic wellbeing and TBE risk. Correspondingly, in 2009, the economic recession was associated with an increase with TBE in these three countries.

Sexual selection acts on traits that increase the reproductive success of an individual in competition with other individuals of the same sex over reproductive opportunities, through intra-sexual competition and inter-sexual mate choice. Because males benefit more from remating than females, they are subject to more intense sexual selection. Modern genetic tools have shown that females often mate promiscuously, thus prolonging sexual selection after insemination through intra-sexual (sperm competition) and inter-sexual (cryptic female choice) episodes. Despite increasing interest in post-copulatory sexual selection, the implications of sperm competition, cryptic female choice and underpinning mechanisms remain little understood. This thesis adopts an integrated approach to quantify the relative importance of post-copulatory episodes in the operation of sexual selection, elucidate their proximate mechanisms in the red junglefowl Gallus gallus. By combining behavioural observations of replicate groups with paternity data, I show that female promiscuity decreased the total opportunity for sexual selection in a group, but accounts for an unexpectedly large proportion of the variance in male reproductive success. By comparing the operation of sexual selection on multiple male traits, I show that post-copulatory sexual selection reinforced pre-copulatory sexual selection for male social dominance and that female preferred to mate with compatible males. I used experiments to study the mechanisms of post-copulatory sexual selection by studying the effect of seminal fluid in sperm competition and cryptic female choice in relation to male status and relatedness. Following previous work indicating that seminal fluid products influence sperm quality in this species, I tested in vivo whether the seminal fluid of an ejaculate acts differentially towards sperm from the same ejaculate and rival sperm, and found no evidence for this idea. Finally, I show that cryptic female choice can drastically bias the outcome of sperm competition, and that female fowl might bias paternity toward unrelated males.

Studies of the evolution of the major histocompatibility complex (MHC) have been central to the understanding sexual selection and pathogen-mediated selection. The European badger Meles meles is well suited for exploring such questions because of its life history characteristics, reproductive biology and mating system. In this thesis, I examined both MHC class I and class II genes. Seven putatively functional sequences were found for class I genes and four for class II DRB genes. Evidence of past balancing selection of both genes was demonstrated by the d_Nd_S ratio, by positive selection at the antigen-binding site (ABS) and by trans-species polymorphism of alleles within other mustelids and carnivores. MHC class I genes also showed evidence of concerted evolution, but domains showed different evolutionary histories. MHC genes may influence microbiota and odour of an individual and influence mating preferences. I examined the bacterial community of the subcaudal gland secretion and demonstrated a high number of bacterial species (56 operational taxonomic units), which cubs exhibited a higher diversity than adults. The microbiota may lead to an individual-specific odour as a cue signaling the MHC genotype of potential mating partners. I report the first evidence for a MHC- based mating preference in carnivores. Female badgers showed a MHC-assortative mate choice towards breeding with males that had functionally similar MHC genes, for MHC class II DRB genes. This applied to neighbouring-group matings. I also found considerable annual fluctuation in the occurrence of MHC-based mate choice. Based on genome-wide background in the same mating randomizations I found no evidence of inbreeding, which indicated that MHC similarity was apparently the actual target of mate choice. In line with MHC-assortative mate choice, MHC heterozygosity had no influence on the co-infection status. Individual MHC alleles did, however, associate with resistance and susceptibility to specific pathogens, suggesting that MHC diversity may be driven and maintained by pathogen-mediated selection through rare-allele advantages and/or fluctuating selection. My study of genetic characteristics, mate choice and pathogen pressures in a wild population revealed past and contemporary evolutionary process of the MHC genes. This increases knowledge of how the MHC may affect mating behaviour and sexual selection, ultimately influencing population processes.

As ecosystems become increasingly fragmented, there has been a proliferation of research into fields such as resource use, movement ecology and habitat connectivity. To understand how species may adapt to threats associated with habitat fragmentation it is necessary to study these processes in dispersing individuals. However, this is seldom done. Dispersal is one of the most important life-history traits involved in species persistence and evolution, but the consequences of dispersal are determined primarily by those that survive to reproduce. Although dispersal is most effectively studied as a three-stage process (departure, transience and settlement), empirical studies rarely do so and an investigation into the entire process has probably never been carried out on any one species. Here I investigate the survival, resource use, movement ecology and connectivity of African lions (Panthera leo) in all three dispersal phases in addition to adulthood. I make use of a longterm dataset incorporating radio-telemetry and observational data from lions in Hwange National Park, Zimbabwe. Dispersal is inherently risky and my results show that male lions that disperse while young suffer high mortality, young dispersal being brought about by high off-take of territorial males. Dispersing males may be aware of risks associated with territorial adults as they position themselves far from them and utilise habitats and resources differently. However, dispersers, compared to adult males and females, are far less averse of risky, anthropogenic landscapes, suggesting they are the demographic most prone to human-lion conflict. The ontogenetic movement behaviour of lions reflects a transition from directional movement during transience, suggestive of sequential search strategies, to random or periodic use of a fixed territory after settlement. In terms of habitat connectivity, I show that radically different conclusions emerge depending on which demographic is used to parameterise connectivity models. Understanding the shifting mechanisms that species adopt throughout ontogeny is critical to their conservation in an increasingly fragmented world.

Few studies of tropical fish communities compare fish distributions across the full range of near-shore tropical marine habitats. As a result, our understanding of tropical marine fish communities is often biased towards habitats preferred by researchers. The distribution and habitat preferences of all 136 fish species occurring at 106 stations in three bays off Tortola in the British Virgin Islands were assessed. At a species level, habitat type was often the key factor influencing fish distribution. Of the 44 species occurring at more than 10 stations, 3 were sand specialists, 5 were strongly associated with mangroves, 7 were dependent on seagrass and algal beds and 14 were found only on the forereef. Eleven species were widely distributed both on the forereef and in bays, and 9 of them showed clear evidence of ontogenetic partitioning. The juveniles all preferred bay habitat types and moved onto the forereef as they approached sexual maturity. At a community level, five distinct fish assemblages were found. The assemblage types were classified according to their distribution: 1) forereef, 2) reef flat, 3) non-mangrove associated seagrass, 4) mangrove associated seagrass and 5) eutrophic areas. Forereef stations were the most species-rich with 24 species per station while eutrophic stations had the most depauperate communities with only 4 species per station. Variation in fish species richness at each station was largely explained by a simple habitat complexity index. It accounted for 70% of the variation in fish species richness and 21% of the variation in fish abundance. Rugosity and variety of growth forms were the most important predictors of species richness, but the height of the habitat architecture was the most useful predictor of fish abundance. Artificial reefs were constructed to test the effects of each habitat complexity variable experimentally. Increasing rugosity, variety of growth forms and percentage hard substrate increased the observed number of species but increasing the variety of hole-sizes, and height had no effect. The only complexity variable that had a significant effect on fish abundance was percentage hard substrate. In addition to the static substrate structure, long-spined sea urchins Diadema antillarum affected fish distribution because small fish shelter from predators in their spines. The urchins increase species richness and abundance in low complexity seagrass beds, but on artificial reefs, where shelter was not a limiting factor, the effect was less pronounced. Organic pollution is another factor that negatively affects the fish community by reducing fish species richness and abundance. Poor water quality often alters the natural habitat, confounding observations, but fish species richness was reduced in polluted areas even when artificial reefs were used as habitat controls. The applications of these findings to the management and conservation of fish in the British Virgin Islands are discussed.

This thesis proposes a collection of analytical and computational methods for inferring an underlying social structure of a given population, observed only via timestamped occurrences of its members across a range of locations. It shows that such data streams have a modular and temporally-focused structure, neither fully ordered nor completely random, with individuals appearing in "gathering events". By exploiting such structure, the thesis proposes an appropriate mapping of those spatio-temporal data streams to a social network, based on the co-occurrences of agents across gathering events, while capturing the uncertainty over social ties via the use of probability distributions. Given the extracted graphs mentioned above, an approach is proposed for studying their community organisation. The method considers communities as explanatory variables for the observed interactions, producing overlapping partitions and node membership scores to groups. The aforementioned models are motivated by a large ongoing experiment at Wytham woods, Oxford, where a population of Parus major wild birds is tagged with RFID devices and a grid of feeding locations generates thousands of spatio-temporal records each year. The methods proposed are applied on such data set to demonstrate how they can be used to explore wild bird sociality, reveal its internal organisation across a variety of different scales and provide insights into important biological processes relating to mating pair formation.

This thesis describes an investigation into silk spinning, with the objective of producing high performance silk fibres in the laboratory using a novel spinning device based upon observations on natural spinning glands and processes. After an in-depth literature review the work is reported in two sections: natural and artificial spinning. The literature provides fragmented data on different aspects of natural silk production, and artificial spinning has not yet reproduced fibres with the properties of native silk fibres, despite unfounded claims of biomimetic spinning. The first half of the thesis looks at natural silk spinning. The work started with a general study of the morphology of spider and silkworm spinning ducts: First, how the silk fibre develops as the dope flows through the gland; and second the relationship between silk fibre properties and both gland morphology and spinning speed. More detailed studies using histochemical and spectroscopic investigations showed that the silk ducts of the spider Nephila edulis and the silkworm Bombyx mori both contain β-chitin, despite an evolutionarily distant common ancestor. Finally, observations showed that the duct of N. edulis consists of alternating nanoporous discs, and FEA modelling indicated that the duct is optimised for mechanical integrity and permeability. The second half of the thesis describes the development of a spinning device that uses natural silk dope mainly taken from B. mori as feedstock. It begins with a description of the gradual development of the engineering aspects of the spinning device, to meet challenges raised during the spinning investigation. The development of a centrifugal capillary rheometer, for practical quantitative insights into rheological processes is then presented. Finally the spinning investigation is reported: first, the screening of spinning in glass capillaries based upon natural gland dimensions and flow rates, which have been shown to induce fibrillation in silk dope in a rheometer, and also included initiation of instability through heat applied along the capillary; second, the final spinning evaluation, using lessons learned from all the screening trials throughout the project, but also including a key development of a hydrophobic coating on the capillary tip to inhibit droplet formation and massively increase the process stability and ease of fibre production. The main conclusions from this work are that good silk fibre cannot be spun by flow shear stress alone; and, that heat instability induces indiscriminate gelation of the silk, whose disordered molecular structure gives poor silk fibre properties. The body of work behind these conclusions provides fundamental background information and new insights that will contribute to the next stages of development of artificial silk spinning, from obtaining a better understanding of the biology of natural spinning glands to the engineering difficulties of implementing the bioinspired principles.

New Caledonian crows (NC crows; Corvus moneduloides) are the most prolific avian tool-users and crafters, using up to three unique tool types derived from numerous plant materials. Since the discovery that wild populations of NC crows use and manufacture different tools in different locations with no measured environmental correlates to these distributions, the process by which NC crows acquire their tool-oriented behavior has been investigated. Two major findings were discovered in 2005: NC crows have a genetic predisposition to manipulate stick like objects, and they increase their rate of manipulation when exposed to social influences. Since then, much of the research into the sociality of wild NC crows has focused on direct social influences, especially the parent-juvenile relationship, yet no social network of wild NC crows has been described. In my thesis, I characterized a new proximity-logging device, Encounternet, and outline a four-step plan to assess error in animal borne devices; uncovered drivers, such as relatedness, space-use, and environmental factors, of wild NC crow sociality, and experimentally manipulated the social network, revealing immediate changes to the number of day-time and roosting partners, the breakdown of first-order relatedness driving sociality, and an increase in the amount of time NC crows associate; and revealed an indirect pathway via tools left behind by conspecifics allowing for the transmission of tool-properties between unrelated NC crows. Altogether, I furthered our understanding of wild NC crow sociality through the use of an animal-borne device, experimental manipulation in the wild measuring the response of the NC crow social network, and demonstrated the utility of animal-borne devices in mapping the network of a population of wild birds.

Inbreeding occurs when relatives mate and have offspring. Inbreeding depression is hypothesized to have influenced the evolution of mating systems and behavioural mechanisms of inbreeding avoidance in the animal kingdom. Inbreeding in the wild is difficult to measure, as in order to build a pedigree allowing us to identify matings between relatives, the identity of as many as possible members of a population needs to be known. For a long time, the main source of knowledge about inbreeding depression was based on laboratory and agricultural studies, which did not reflect the array of environmental pressures wild populations have to cope with. In consequence, the deleterious consequences of inbreeding have often been underestimated. This is problematic because accurate estimates of the effect size of inbreeding depression are needed to study the strength of selection on inbreeding avoidance mechanisms, and are also of importance to conservation genetics. The aim of this thesis was to use pedigree data to infer the occurrence and effects of inbreeding using over forty years of breeding events of the great tit Parus major from Wytham Woods, Oxfordshire. The effects of inbreeding on fitness were investigated across a life-history continuum, and across environments. I found that close inbreeding (f=0.25) resulted in pronounced inbreeding depression, which acted independently on hatching success, fledging success, and recruitment success, and reduced the number of fledged grand-offspring by 55%. My results therefore suggest that estimates of fitness costs of inbreeding must focus on the entire life cycle. I also show that the variation in the strength of inbreeding depression varies across environments, particularly so the more the environmental variable considered is linked to fitness. These results emphasise the need of using relevant environmental contrasts when investigating inbreeding by environment interactions. I further asked whether individuals involved in matings with relatives differed relative to individuals mating with unrelated partners. I did not find any evidence for clear predictors of inbreeding, and I show that inbreeding depression in our population is entirely independent of any tendency for low quality parental genotypes, or phenotypes, to inbreed. Neither did I find any evidence for active inbreeding avoidance: great tits did not mate less often with kin than expected based on several scenarios of random mating, nor did I find increased rates of extra-pair paternity among birds breeding with relatives. In fact, I observed quite the contrary, as birds mating with kin exhibited a higher than average rate of close inbreeding relative to all scenarios of random mating investigated, showed lower rates of extra-pair paternity and divorce than birds mated to unrelated partners. I hypothesise that cases of occasional inbreeding in this population may result from mis-imprinting or a related process whereby some birds develop particularly strong bonds that are at odds with all predictions of avoiding inbreeding. Finally, I asked to what extent natal dispersal, a behaviour that is often hypothesized to play an important role in avoiding inbreeding, indeed reduces the likelihood of inbreeding. I found that male and female individuals breeding with a relative dispersed over several-fold shorter distances than those outbreeding. This led to a 3.4 fold increase (2.3-5, 95% CI) in the likelihood of close inbreeding relative to the population average when individuals dispersed less than 200m. This thesis demonstrates that inbreeding has deleterious effects on a wild population of birds, occurring throughout an individual’s life, and is of varying strength across environments. My findings strongly support the theory that natal dispersal should be considered as a mechanism of prime importance for inbreeding avoidance.