In the light of increasing pesticides resistance in agricultural pests and in insect vectors of human diseases, leading to the rise in occurrence of mosquito-borne diseases, new, efficient and environmentally friendly methods of pest control are needed. Sterile Insect Technique (SIT), relying on mass releases of radiation sterilised males to reduce reproductive potential of target pest populations, although not new, offers an alternative to the use of pesticides and is an environmentally non-polluting method of insect control. Many insect species, however, are not very amenable to classical SIT, due to detrimental side-effects of radiation treatment. We propose a new method, a genetically engineered modification of classical SIT, replacing radiation with genetically induced sterility. Based on conditional expression of male-germline targeted nucleases which introduce double strand breaks into the male germline DNA to render males sterile, this method emulates SIT mechanism, at the same time eliminating radiation and associated detrimental side-effects. Different variants of such a system were investigated in this project, eventually leading to the creation of functional conditional male-sterility systems in two model organisms – the Yellow fever mosquito, Aedes aegypti and the Mediterranean fruit fly, Ceratitis capitata. Both systems utilise chimeric nuclease composed of protamine and FokI cleavage domain fusion. The sperm-specificity and the conditionality of the sterile phenotype have been achieved through the use of tetracycline repressible expression system driven by the β2-tubulin promoter in Ceratitis capitata and by the Topi promoter in Aedes aegypti.

A combination of habitat loss, degradation and fragmentation are causing declines in butterfly species across Europe, with habitat specialists more vulnerable than generalists. The influence of climate change is serving to complicate matters, particularly at the edge of geographic ranges. Improving the quality of habitats is a key tool for the conservation of threatened species at both a local and landscape scale and this requires a thorough understanding of the ecology and habitat requirements of species. From this perspective this thesis examines and compares the ecology of two closely butterfly species, Polyommatus bellargus (the Adonis Blue) and Polyommatus coridon (the Chalkhill Blue) in southern England; calcareous grassland specialists, found in a highly fragmented ecosystem at the northern most edge of their geographic range. Chapter 2 explores whether there has been an expansion in the larval niche of P. bellargus due to a warming climate in recent decades, and finds no indication of change once concurrent local habitat change has been accounted for. Chapter 3 examines the role that differences in microhabitat requirements play in differentiating the larval niche of P. bellargus and P. coridon, showing P. coridon has broader microhabitat requirements. Chapter 4 experimentally examines the specificity of mutualistic interactions between the larvae of each butterfly species with the ants Myrmica sabuleti and Lasius niger. Chapter 5 uses microsatellites to explore the genetic structure of both P. bellargus and P. coridon, showing (as predicted) that P. coridon has a less fragmented population structure than P. bellargus, but both species have high levels of inbreeding.

Decision making is a framework we impose on a vast universe of possible behaviors to better understand the advantages and disadvantages of acting in different ways. Decisions under uncertainty are of particular interest because stochasticity is a feature of environments both today and throughout evolutionary history. As a result, we might expect decision makers (DMs) to have evolved mechanisms to handle variability. In this dissertation, I examine common decision processes in several model species: European starlings (Sturnus vulgaris), Banded tetras (Astyanax fasciatus) and humans (Homo sapiens). The broad range of approaches discussed include currencies DMs are expected to maximize (Risk Sensitivity Theory, Expected Utility), the currencies DMs do maximize (e.g. long versus short term rate maximizing models), the representation of outcomes in memory (Scalar Expectancy Theory) as well as explicit choice mechanisms (Sequential Choice Model). The first section of this thesis discusses responses to risk, offering humans and starlings choices between options which deliver certain or variable outcomes. Starlings demonstrate sensitivity to changes in the probability of variable outcomes and strong support for local rate maximization. Humans appear similarly sensitive to their own accuracy when task difficulty is varied. When the DM’s affective state was manipulated, neither humans nor starlings exhibit changes in risk preferences, but the effectiveness of these manipulations used could not be confirmed. Another topic of inquiry is the effect of the DM’s state at the time of valuation learning. State dependent valuation learning is demonstrated for the first time in a fish species, but Within Trial Contrast is not observed in starlings. Lastly, two experiments find strong support for the Sequential Choice Model, a promising new model of the mechanism of choice. Taken together, these experiments offer a glimpse into shared decision processes, but leave open questions about the mechanisms through which value is acquired.

Species diversity varies dramatically across the surface of the Earth. A key step in the accumulation of species diversity is the ability of species to coexist in biological communities. Thus, identifying the mechanisms underlying community assembly is a major challenge for ecologists seeking to explain patterns in species diversity and composition. Recently some consensus has been reached on the set of processes that influence community assembly: speciation, demographic stochasticity, niche-based fitness trade-offs among species and dispersal. However, it is unclear how the importance of a particular process changes with spatial scale, which interactions exist among processes at large spatial scales and the extent to which niche-based resource partitioning among species explains differences in diversity among communities. Neotropical birds offer an ideal opportunity to address these uncertainties because of their high diversity and the existence of detailed information on their evolutionary history and ecology. In this thesis, I first use trait and phylogenetic metrics of community structure to show that both habitat filtering and interspecific competition shape community composition at the scale of individual bird territories (~1-2 ha). Second, I use simulations of community assembly to show that trait-based metrics of community structure outperform phylogenetic metrics for detecting niche-based community assembly, and that both sets of metrics often have low power when multiple processes influence community composition. Third, taking a trait-based, species-level approach, I show that both habitat filtering and interspecific competition influence species occurrence at regional scales (~75000 km²), and interact with dispersal ability so that their effect on species occurrence is increased for species with greater dispersal ability. Finally, using a combination of trait- and isotope-based methods to quantify resource partitioning, I show that species' niche widths do not change and niche overlap is reduced at high compared to low species richness. Taken together, these results suggest that both habitat filtering and interspecific competition (i.e. niche-based processes) influence community assembly from local to regional scales. However, at least at regional scales, the degree to which these processes are important for determining the occurrence of any given species depends on that species's dispersal ability. They also suggest, based on niche-based interspecific competition influencing community composition, that differences in species richness among communities are in part explained by differences among sites in the breadth of available niche space, not by increased ecological specialisation or niche overlap.

Inclusive fitness theory concerns the study of social traits. Often, individuals differ in their phenotype (e.g. size, weight, nutritional state) independently of their genetic make up, that is, individuals differ in their quality. Individuals can then be classified into different “classes” according to their quality, which enable us to understand social evolution in class-structured populations. This is important because individuals in natural populations often differ in quality, either because of intrinsic factors (e.g. size), or extrinsic factors (e.g. resource availability). My thesis concerns the evolution of social traits in class-structured populations. In chapter 1, I make a brief introduction to my thesis, providing the abstract of each chapter. In chapter 2, I outline a general theory of individual quality, where I show how individual quality impacts social evolution in two fundamental ways. In chapter 3, I show that resource heterogeneity greatly influences the evolution of conditional social behaviour. In chapter 4, I show that temporal group-size heterogeneity promotes the evolution of both conditional helping and harming. In chapter 5, I analyse the effect of individual quality on kin selection. I find that individual quality has an important impact in kin selection, which can lead to extreme forms of social behaviour. In chapter 6, I show that stable environments promote the evolution of negative density-dependent dispersal, while unstable environments promote the evolution of positive density-dependent dispersal. In chapter 7, I show that budding and low local quality promote the evolution of dispersal and cooperation.

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.

Dispersal behavior plays an important role for the geographical distribution and population structure of any given species. Individual’s fitness, reproductive and competitive ability, and dispersal behavior can be determined by the age of the individual. Age-dependent as well as density-dependent dispersal patterns are common in many bird species. In this thesis, I first present age-dependent breeding ability and natal site fidelity in white storks (Ciconia ciconia); migratory birds breeding in large parts of Europe. I predicted that both the proportion of breeding birds and natal site fidelity increase with the age. After the seventies of the last century, following a steep population decline, a recovery of the white stork population has been observed in many regions in Europe. Increasing population density in the white stork population in Eastern Germany especially after 1983 allowed examining density- as well as age-dependent breeding dispersal patterns. Therefore second, I present whether: young birds show more often and longer breeding dispersal than old birds, and frequency of dispersal events increase with the population density increase, especially in the young storks. Third, I present age- and density-dependent dispersal direction preferences in the give population. I asked whether and how the major spring migration direction interacts with dispersal directions of white storks: in different age, and under different population densities. The proportion of breeding individuals increased in the first 22 years of life and then decreased suggesting, the senescent decay in aging storks. Young storks were more faithful to their natal sites than old storks probably due to their innate migratory direction and distance. Young storks dispersed more frequently than old storks in general, but not for longer distance. Proportion of dispersing individuals increased significantly with increasing population densities indicating, density- dependent dispersal behavior in white storks. Moreover, the finding of a significant interaction effects between the age of dispersing birds and year (1980–2006) suggesting, older birds dispersed more from their previous nest sites over time due to increased competition. Both young and old storks dispersed along their spring migration direction; however, directional preferences were different in young storks and old storks. Young storks tended to settle down before reaching their previous nest sites (leading to the south-eastward dispersal) while old birds tended to keep migrating along the migration direction after reaching their previous nest sites (leading to the north-westward dispersal). Cues triggering dispersal events may be age-dependent. Changes in the dispersal direction over time were observed. Dispersal direction became obscured during the second half of the observation period (1993–2006). Increase in competition may affect dispersal behavior in storks. I discuss the potential role of: age for the observed age-dependent dispersal behavior, and competition for the density dependent dispersal behavior. This Ph.D. thesis contributes significantly to the understanding of population structure and geographical distribution of white storks. Moreover, presented age- and density (competition)-dependent dispersal behavior helps understanding underpinning mechanisms of dispersal behavior in bird species. / Das Verständnis der Mechanismen, die dem Ausbreitungsverhalten und der Wahl des Neststandorts zugrunde liegen, gibt wichtige Einsichten in Strukturen und Dynamiken von Tierpopulationen. Der Gesundheitszustand, die Produktivität und Konkurrenzfähigkeit sowie das Ausbreitungsverhalten eines Individuums können über das Alter ermittelt werden. Alters- und dichteabhängige Veränderungen in Verbreitungsmustern kommen bei vielen Vogelarten vor. In der vorliegenden Studie untersuchten wir zunächst den Effekt des Alters auf die Reproduktivität, auf die Wahl des Neststandorts sowie auf die Geburtsorttreue des Weißstorchs (Ciconia ciconia). Wir fragten, ob sowohl der Anteil der brütenden Individuen als auch die Geburtsorttreue mit dem Alter zunimmt. Weißstörche sind Zugvögel, die während der Migration zumeist segelnd die Thermik nutzen und in weiten Teilen Europas brüten. Nach einem starken Bestandsrückgang konnte in vielen Regionen Europas ab den 1970er Jahren wieder ein positiver Trend in der Populationsentwicklung beobachtet werden. Die zunehmende Populationsdichte, besonders nach 1983 in der ostziehenden Subpopulation in den fünf Bundesländern der ehemaligen DDR, erlaubte die Analyse von dichte- und altersabhängigen Präferenzen in der Richtung der Brutstandorte sowie in der Verbreitungsfrequenz und -distanz. Wir untersuchten zudem die Alters- und Dichteabhängigkeit der Ausbreitungsrichtung einer Teilpopulation. Wir fragten, ob und wie die Hauptzugrichtung im Frühjahr mit der Verbreitungsrichtung interagiert: Beeinflussen Alter und Populationsdichte die Ausbreitungsrichtung? Der Anteil der brütenden Individuen, die älter als 22 Jahre sind, nahm innerhalb der beobachteten Teilpopulation ab, vermutlich aufgrund einer altersbedingten Abnahme des Gesundheitszustands. Junge Vögel zeigten eine starke Geburtsorttreue, was auf eine genetische Komponente in den Zugmustern junger Störche hinweist. Generell trat bei jungen Störchen häufiger Ausbreitungsverhalten auf als bei älteren Störchen. Eine signifikante Zunahme der Ausbreitungsdistanz von Individuen über die Zeit lässt auf eine dichteabhängige Komponente im Ausbreitungsverhalten der Weißstörche schließen. Weiterhin wurde eine signifikante Interaktion zwischen dem Alter sich ausbreitender Individuen und dem betrachteten Jahr gefunden. Demzufolge breiteten sich alte Vögel über die Zeit über größere Distanzen aus, vermutlich um der ansteigenden Konkurrenz, bedingt durch den wachsenden Bestandsdruck, zu entgehen.

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 thesis addresses decision mechanisms in foraging situations, using laboratory experiments with European starlings (Sturnus vulgaris). Building on previous work from the Behavioural Ecology Research Group, I chose the Sequential Choice Model (SCM; reviewed in Kacelnik et al., 2011 − Appendix 1) as a starting point, and tested its premises and predictions generalising it to different experimental protocols. Classical decision models do not relate choice preferences to behaviour towards isolated options, and assume that choices involve time-consuming evaluations of all alternatives. However, previous work found that starlings’ responses to isolated options predict preference in choices, and that response times to single-option encounters are not reliably longer than response times in choices. Since, in the wild, options are normally encountered sequentially, dealing with isolated options can be considered of greater biological, and possibly psychological, significance than simultaneous decisions. Following this rationale, the SCM postulates that when multiple simultaneous stimuli are met they are processed in parallel, each competing against the memory of background opportunities, rather than comparing present options to each other. At the time of launching this research, these ideas had only been applied to protocols involving just two deterministic alternatives and offering no chance to explore the influence of learning history (i.e., how animals learn to choose; see Chapter 4). To increase their relevance and offer more rigorous tests, I generalised them to situations with multiple (see Chapters 2, 4 and 5), and in some cases probabilistic alternatives (see Chapter 3), controlling the learning regime. I combined these extensions with tests of economic rationality (see Chapter 6), a concept that is presently facing sustained debates. Integrating the result of all experimental chapters (see Chapter 7), my results support the notion that behaviour in single-option encounters is fundamental to understand choice behaviour. The important issue of whether choices involve a decision time cost or the opposite, a shortening of response times, remains unsolved, as neither could be evidenced reliably.