Biologists have long used demographic approaches to answer questions in ecology and evolution. The utility of these approaches has meant a constant development and refinement of methods. A key milestone has been the development of phenotype structured population models that link ecology and evolution. Moreover, biostatistical research steadily improves methods to coax demographic information from scarce data. In this thesis, I build upon some of the recent advances in the field. My first three studies focus on the consequences of sex differences in life history for population dynamics. Firstly, I test whether males matter for the dynamics of African lion (Panthera leo) populations via a previously unquantified mechanism: the inheritance of phenotype from father to offspring. Secondly, I develop a method to estimate age-specific mortality rates for both sexes in species where one of the sexes disperses around the age of maturity. Thirdly, I apply this method to study variation in mortality between the sexes and between two populations of African lions. After these three chapters, which make contributions to the field of sex-structured population dynamics, I focus on the integration of phenotype structured modelling and quantitative genetics. I illustrate how heritability of a quantitative character that develops with age depends on (i) viability selection, (ii) fertility selection, (iii) the development of the phenotype with age, and (iv) phenotype inheritance from parents to offspring. Our results question the adequacy of quantitative genetics methods to obtain unbiased estimates of heritability for wild populations. This thesis advances our understanding of population development over ecological time scales. This knowledge has applications in conservation and population management, but also contributes to untangling evolutionary processes in wild animals.

The main object of this thesis was to find out whether grammar school students prefer zoological objects in biology lessons presented a) on photo in their natural habitat or b) the real specimen presented in liquid in glass cylinders and burettes. I have chosen these research questions: 1. In which form do pupils prefer zoological objects, the real specimen presented in liquid in glass cylinders or burettes or in their natural habitat on a photograph? 2. How do these preferences differ during the course of their high school study? 3. How do these preferences differ according to gender? 4. Are these preferences influenced by a specific animal? 5. Are these preferences influenced by other factors, e.g. by the appearance and personal experiences with live nature or frequency of using these objects in biology lessons. The research was implemented by using a questionnaire and a preferential test and was given to first and third year students from five Prague high schools. The conclusion of my research study shows that students prefer neither animals on photographs nor the real specimen presented as a liquid preparation. My research did not show that the preferences of the tested students were influenced by any of the tested factors, e.g. type of living, popularity of biology or students' grade in...

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.

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.

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.

Intraguild competition and predation have been recognised as important ecological factors influencing the population dynamics of carnivores. The effects of these interactions are often asymmetrical due to a size-related dominancy hierarchy. However, it has been suggested that competitively subordinate carnivores can minimise the costs of predation and competition through spatial and temporal avoidance. Here I investigate the ecological and behavioural mechanisms by which cheetahs (Acinonyx jubatus) coexist with competitively stronger lions (Panthera leo) and spotted hyaenas (Crocuta crocuta). Fieldwork was carried out in the Okavango Delta, northern Botswana, between October 2008 and August 2011. A total of 20 Global Positioning System (GPS) radio-collars were fitted on all known cheetahs (n=6), lion prides (n=5) and spotted hyaena clans (n=6) in the study area (approx. 3 000 km²). Pre-programmed radio-collars recorded locations and activity continuously for each individual and these data were complemented with direct behavioural observations. Cheetah data were analysed with respect to the temporal and spatial likelihood of encountering lions and spotted hyaenas. Results suggest that the response to the risks posed by other predators is species-specific, habitat-specific and dependent on the immediacy of the risk. Resource partitioning was not the main mechanism for coexistence as cheetahs overlapped extensively with lions and spotted hyaenas in time, space and habitat use. Instead, cheetahs adjusted their spatial distribution in response to immediate risks or adapted their habitat use depending on their vulnerability (e.g. behaviours such as feeding or with differing levels of moonlight at night). In general, cheetah temporal and spatial distribution is a hierarchal process, firstly driven by resource acquisition and thereafter fine-tuned by predator avoidance. In addition, habitat heterogeneity seemed to be key in facilitating coexistence. Understanding the behavioural mechanisms that interacting apex predators adopt to regulate these negative interactions could be crucial to carnivore conservation, especially as human-related habitat loss is forcing species into ever smaller areas.