Understanding the genetic basis of fitness differences has been a major goal for evolutionary biologists over the last two decades. Although there are many studies investigating how natural selection can promote local adaptation, few have succeeded to find the link between genotype and fitness of the phenotype. Polymorphic genes of the major histocompatibility complex (Mhc) are excellent candidates for such associations as they are a central component of the vertebrate immune system, playing an important role in parasite resistance, and hence can have direct effects on survival of their bearers. Although associations between Mhc and disease resistance are frequently documented, the epidemiological basis of the host-parasite interaction is often lacking and few studies have investigated the role that Mhc genes play in individual variation in fitness; thus comparatively little is known about the fitness consequences of Mhc in wild populations. Furthermore, the majority of work to date has involved testing associations between Mhc genotypes and disease. However, the mechanism by which any direct selection on the Mhc acts, depends on how genotypes map to the functional properties of Mhc molecules. The aim of this thesis was to characterize Mhc alleles in terms of their predicted functional properties and to investigate whether and how selection operates on Mhc class I functional variation using the great tit (Parus major) population at Wytham Woods as a model host species. Through a comprehensive characterization effort and the use of 454 pyrosequencing platform, I performed a detailed analysis of genetic variation at Mhc class I exon 3 and grouped alleles with similar antigen-binding affinities into supertypes to classify functionally distinct Mhc types. There was extreme complexity at the Mhc class I of the great tit both in terms of allelic diversity and gene number. A total of 862 alleles were detected from 857 individuals; the highest number yet characterized in a wild bird species. The functional alleles were clustered into 17 supertypes; there was clear evidence that functional alleles were under strong balancing selection. To understand the role of Mhc in disease resistance, I examined the linkage between Mhc supertypes, Plasmodium infection and great tit survival, and showed that certain functional variants of Mhc confer resistance to two divergent Plasmodium parasite species that are common in the environment. I further investigated the fitness consequences of functional variation at Mhc, using mark-recapture methods and long-term breeding data; and tested the hypotheses that selection: (i) maximizes Mhc diversity; (ii) optimizes Mhc diversity, or (iii) favours specific functional variants. I found that the presence of three different supertypes was associated with three different components of individual fitness: adult survival, annual recruitment probabilities and lifetime reproductive success. In contrast, there was no evidence for a selective advantage of Mhc functional diversity, either in terms of maximal or optimal supertype diversity. Finally, I explored the role that Mhc plays in female mate choice decisions and examined the reproductive fitness consequences of Mhc-dependent mating patterns. There was little evidence to suggest that functional dissimilarity at Mhc has any influence on female mate choice decisions or that dissimilarity at Mhc affects the reproductive output of the social pair. Overall, this thesis provides strong support for the suggestion that selection favours specific functional variants of Mhc, possibly as a result of supertype-specific resistance or susceptibility to parasites that exert strong selective pressures on their hosts; whereas there is no support for selection favouring maximal or optimal Mhc diversity. More importantly it demonstrates that functional variants of Mhc class I loci are an important determinant of individual fitness in natural populations.

Great tits (Aves: Passeriformes; Parus major) and blue tits (Parus caeruleus) nested in boxes in Wytham Woods near Oxford. The breeding densities of both species were limited by the availability of nest sites. The larger great tits were dominant in obtaining nest boxes. This was most important where breeding sites were scarce. The two species did not maintain interspecific territories or interfere with interspecific nest site spacing beyond the immediate vicinity of the nest. Blue and great tit numbers fluctuated in parallel where nest sites were not limiting resources. Annual changes in breeding numbers of great tits were negatively related to blue tit breeding density but great tit density did not seem to affect changes in the blue tit population. Overlap in the feeding sites of blue and great tits was greatest during the summer and interference competition was lowest at this time. The nestling diets of the two species were very similar. Despite an apparent abundance of food for nestlings, adults were pressed to feed large broods. Food for nestlings was probably a limiting and depletable resource. The date of clutch initiation of great tits, but not blue tits, was retarded at high densities of blue or great tits. The clutch sizes of both species were probably negatively affected by high breeding densities of congeners but the results were not clear cut. Heavy great tit fledglings are most likely to survive to breed. Great tit fledglings were heavier at low densities of blue tits. An experiment in which blue tit young were removed from a section of the Woods, showed that great tit nestlings were heavier and developed faster, and that female condition was better, than in a control section or section where blue tit broods were supplemented. In terms of resource competition, blue tits were the better scramble competitors and great tits the better interference competitors.

Thesis (doctoral)--Swedish University of Agricultural Sciences, 2007. / Thesis documentation sheet inserted. Includes appendix of four papers and manuscripts co-authored with Bo Söderström. Includes bibliographical references. Also issued electronically via World Wide Web in PDF format; online version lacks appendix.

In this master thesis i was tested the effectiveness of the myrmecomorphy.The animals which ones were tested are Phrurolithus festivus the myrmecomorphic spider and the predator, great tit(Parus major). I want to know the adventages and disadventages of myrmecomorph mimicry compared whit other common invertebrates in natural habitat of great tits.

Categorization of predators is fundamental for birds? life. However, the mechanism is not well explained yet. Two basic theories of categorization were tested in the laboratory conditions, where we have the possibility to focus on the behavior of each individuum. In the experiments we focused on the relevance of the key features and colour patterns. The responses of the Great Tit to variously modified dummies of Sparrowhawk were observed. The results confirm that the bird does not recognize the predator only on the basis of the key features and the colour patterns have a essential influence on the categorization. Categorization of the predator is a complex process and it cannot be explained only on the basis of one psychological theory. A combination of the key features and a prototype of a particular category learned throughout its life is probably used.

In this thesis the spatial cognitive experiment with great tits (Parus major) as a model species is described. In this experiment tits learnt to respond to one of four response keys according the presented stimuli. Two types of stimuli were used: one of them contained configural information about response keys, but the other didn´t. The experiment tested the difference of using these two types of spatial information in great tits in two experimental designs. In the first design stimuli were presented in computer screen, in the other they were presented in the floor of experimental chamber. The thesis then discusses the performance of tits in two types of stimuli and two versions of experimental design and compares these results with pigeons in similar experiment.

Social learning is a topic of many studies. We tested the effect of social learning on the acquisition of avoidance against aposematic prey. We have chosen wild-caught adult and naive hand-reared juvenile great tits (Parus major) as a model predator species, because their individual avoidance learnig of aposematic prey is well-studied. We used red and green paper dummies of bugs with a mealworm (Tenebrio molitor) stuck underneath as an artificial prey. Mealworms were soaked in water or in bitter-tasting solution of quinine. We used two types of conspecific demonstrators - naive birds that showed aversive reactions while tasting an unpalatable prey and experienced birds that were trained not to handle the unpalatable pray at all. We compared effects of both demonstrators on discrimination learning and we tested whether these effects differ in adult and juvenile birds. Observing an experienced demonstrator had an effect on the performance of observers at the beginning of learning process. The observers did not reject the unpalatable prey completely, but the number of trials in which they correctly chose the palatable prey was higher in comparison with birds that observed naive demonstrators and birds from the control group with no demonstrator. Latencies to handle the unpalatable prey in the first...

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.

In nature, environments are often variable and heterogeneous influencing ecological and evolutionary processes. This thesis focus on how animals interact with their environment and how that affects the reproductive decisions they make. Using empirical data collected from wild collared flycatcher populations, experiments and molecular approaches I try to unveil some of these relationships and the evolutionary, ecological and conservation implications of these findings are discussed. Firstly, collared flycatchers were shown to use breeding densities of their own and other species using similar resources when assessing costs and benefits associated to breeding in specific habitats. However, species will vary in how informative they are, and the worst competitor – with whom you overlap most in resources needs – also provides the best source of information. Collared flycatcher parents will also benefit differentially from investments in sons and daughters due to habitat characteristics and dispersal differences between the sexes. Here, I show that they will produce more of the sex that will give the highest expected fitness return given the environment they are in. These results also provide a reciprocal scenario to Clark's (1978) classical study of sex ratio adjustment in relation to local resource competition (LRC), as more of the natal philopatric sex is produced when LRC is low. Secondly, the effect of elaborated ornaments on paternity in the socially monogamous collared flycatcher was shown to be of more importance in areas where the intensity of intra- and intersexual conflicts are expected to be elevated. Hence, ornamentation by environmental interactions determines paternity, illustrating that sexual selection through extra-pair paternity is context dependent. Finally, even though the collared flycatcher populations that this thesis is based on have been studied on their breeding grounds for more then 25 years, we know little of where they are when they are not breeding. Here, stable isotope signatures in winter-grown feathers suggests that they may spend their winter with their breeding ground neighbours and do so repeatedly over years. Differences between breeding populations at this small scale should have many impactions for evolutionary and ecological processes as it will, for example, determine with whom individuals interact throughout their life.

By warning signalling prey advertises its unpalatability to the predators. Typical examples are true bugs (Heteroptera) combining optical and chemical signals. The aim of this study was to find out the effectiveness of chemical defence against wild-caught adults of great tits (Parus major) and blue tits (Cyanistes caeruleus) and towards hand-reared juveniles of great tits. The tested prey were adults and larvae of two invasive species of genus Oxycarenus (Heteroptera: Oxycarenidae) (aposematic O. lavaterae, non-aposematic O. hyalinipennis), adults of Horvathiolus superbus (Heteroptera: Lygaeidae) and crickets (Gryllus assimilis) as a control prey. We were focusing on the influence of seed bugs to the initial reaction, the learning process and to the displays of discomfort behaviour in tits. Reactions affected by the tit species, age and sex were compared in adults. Juveniles were divided into two independent experimental groups, one group was offered adults of genus Oxycarenus, the second was offered sunflower (Helianthus sp.) or Digitalis sp. seed fed adults of H. superbus. The first bug offered did not elicit an initial aversive reaction in tits, but birds reacted aversively to the bugs after having further handling experience. All juveniles, contrary to the adult tits, attacked at least one bug...