Natural and sexual selection and sexual conflict are forces shaping the evolution of reproductive behaviour, while constrained by factors like environment, physiology and life-history trade-offs. Parental care is costly both in terms of time and energy. In fish, filial cannibalism is a strategy for caring males to compensate for some of the energy loss. Human impact like eutrophication also alters the basics for animals living in that environment. It is fundamental to any species to adjust its behaviour to a changing environment. Studying sand goby males, I found trade-offs both regarding parental care allocations, like ventilation vs. predator defence, and investment in present vs. future reproductive success. Paternal sand gobies exposed to water with low oxygen levels increased their fanning effort but did not compensate by eating more eggs, even though an increased current parental effort should affect future reproductive success negatively. Investigating if patterns of filial cannibalism change with time of season, I found no correlation. Theory predicts that it should pay more to eat eggs early, when future mating potential is higher than late in the season. However, as early hatching fry are likely to gain higher fitness through larger size the next breeding season, this may provide an opposing selection pressure. In species with male care the only way a female can affect the level of post-spawning care is by choosing a good mate. A female preference to spawn in nests that already contain eggs of other females has been interpreted as a means to avoid filial cannibalism through a dilution effect or to decrease the costs of search time. Yet, in my study females did not avoid filial cannibalism by preferring large clutches to small ones. Oxygenation of the eggs might be a key factor, since both large and small females preferred spawning in nests with small clutches. Thus, as in most animals, trade-offs clearly govern the reproductive behaviour of sand gobies.

The evolutionary goal of individuals is reproduction and sexual selection favours traits improving reproductive success. When males invest less than females in offspring, males have potentially a higher reproductive rate than females. This typically results in sex-specific reproductive strategies of male-male competition and female choice of mating partner. Under polyandry, sexual selection can continue after copulation as sperm competition and cryptic female choice. This thesis focuses on male and female pre- and post-copulatory reproductive strategies in the promiscuous red junglefowl, Gallus gallus ssp., and its domestic subspecies the domestic fowl, Gallus gallus domesticus. Males impose high re-mating rates on females, which triggers female resistance in copulations. In addition, when sexual harassment increases, females re-mate at times of day when male mating propensity is lower, to avoid intense sexual harassment. Males allocate sperm supplies differentially according to (i) variation in female polyandry and own competitive ability, (ii) earlier sperm investment in a female, and (iii) female reproductive quality, signalled by female comb size. Males also perform ‘aspermic’ copulations (i.e. copulations with no semen transfer), which inhibit polyandry and in turn reduce sperm competition. In mating opportunities with relatives, males do not avoid inbreeding. However, females avoid inbreeding before copulation through kin recognition and after copulation by selecting against related males’ sperm. These results show that selection on males to re-mate at higher rates than females and copulate indiscriminately according to partner relatedness, trigger counteracting female responses, creating the potential for sexual conflict over fertilisation. Teasing apart pre- and post-copulatory strategies and the contribution of each sex therefore becomes crucial in order to understand the evolution of reproductive strategies and the mechanisms affecting paternity.

Competition over mating opportunities is a conspicuous characteristic of animal behaviour. In many butterfly species the males establish territories in places advantageous for encountering females. This thesis addresses questions about how territoriality has evolved and is maintained in butterflies. The studies have been conducted using the speckled wood butterfly, Pararge aegeria, as a model species. Males of P. aegeria are found in sunspots on the forest floor (paper I-V), on the lookout for females visiting the sunspots. However, males are only found in sunspots above a certain size (paper III). This behavior is maintained by a mating success advantage, where using large sunspots instead of small sunspots as perching areas generates a higher reproductive output (paper I). The mating success asymmetry is not explained by female choice or by a female preference for large sunspots per se (paper I, V), but rather the large sunspot facilitates visual performance of perching males and improves flight pursuit and interception of females (paper III). Winners of territorial contests gain sole ownership of large sunspot territories, while losers search for a new suitable sunspot territory (paper I, II & IV) or use smaller, suboptimal sunspots as perching sites (paper II). Territorial contests between P. aegeria males are not settled due to an obvious morphological/physiological asymmetry (paper I). Rather, variation in resource value and motivational asymmetries are important for settling contests (paper IV). A majority of male-female interactions (paper V) and matings (paper I) are initiated by a perching male detecting and intercepting a flying female. Furthermore, females can affect their chances of being detected by a perching male by behaving more conspicuously (paper V). This thesis highlights the role of female behaviour, variation in resource value and motivation asymmetries to understand the evolution of territoriality in butterflies.

The sex-role reversal and male pregnancy found in syngnathids are highly unusual traits in vertebrates. Reproductive hormones likely influence development and regulation of these traits. However, very few studies have examined the underlying hormonal mechanisms that mediate female competitive behavior and male pregnancy. New methodologies and better husbandry practices have made such studies more feasible in recent years. Research on a relatively small number of species has suggested that androgens are likely regulators of spermatogenesis and the development of the male brood pouch prior to pregnancy. Androgens are also potential candidates for mediating sex-role reversed behavior in female syngnathids. The goal of this dissertation was to examine the role of androgens in the male reproductive cycle and female intrasexual competitive behavior in the sex-role reversed Gulf pipefish, Syngnathus scovelli. From review of the literature, I developed a model for the hormonal regulation of the male reproductive cycle in seahorses. I predicted that androgens would be low during the early stages of pregnancy and increase during the end of pregnancy as males go through another cycle of spermatogenesis in preparation for the next mating event. My study of 11-ketotestosterone and testis mass across the reproductive cycle in male S. scovelli supported this model. I also conducted several studies on the role of androgens in female competitive behavior. I determined that treatment with 11-ketotestosterone the evening prior to an intrasexual interaction resulted in an increase in competitive behavior in large over small test females. Conversely, treatment with 11-ketotestosterone one hour prior to an intrasexual interaction resulted in a decrease in competitive behavior in large over small females when stimulus female behavior was controlled. A comparative study of competitive and courtship behavior in S. scovelli and the closely related S. floridae suggested that sexual selection has affected competitive and courtship behavior in both males and females of these species. The diversity of reproductive patterns exhibited by syngnathids suggests that they will provide a unique opportunity to assess how hormonal regulation of reproductive behavior and function has evolved within this lineage.

In this thesis, different genetic tools are used to investigate pre- and postzygotic barriers to gene exchange and their role in speciation in the pied flycatcher (Ficedula hypoleuca) and the collared flycatcher (F. albicollis). This species complex consists of four genetically distinct clades that apparently diverged in allopatry (I). Sequencing of introns from autosomal and Z-linked genes from the two species reveals signs of selection on the Z-chromosome. Sexual selection acting on Z-linked genes might explain this pattern (II). By using large-scale genotyping of single nucleotide polymorphisms (SNPs), introgression is observed at autosomal- but not Z-linked loci, mostly from the pied- to the collared flycatcher. Male plumage characters and genes involved in hybrid fitness are largely mapped to the Z-chromosome (III). By studying mate choice of female hybrids I show that there is a link between female preferences and the Z chromosome (IV). The rate of introgression in island versus clinal hybrid zones is consistent with regional differences in hybrid fertility. Asymmetric gene flow from allopatry on the islands may oppose reinforcement, leading to introgression and a partial breakdown of postzygotic isolation. Adaptive introgression may explain the high rate of introgression observed at one of the genetic markers (V). For late breeding female collared flycatchers it appears to be adaptive to use pied flycatchers as social fathers but conspecific males as genetic fathers. Additionally, females in mixed species pairs may reduce hybridization costs by producing an excess of male hybrid offspring that are more fertile than females (VI). In conclusion, the Z-chromosome appears to play a major role in flycatcher speciation. Sexual selection and reinforcement are important mechanisms in the divergence of these birds. However, gene flow from allopatry, introgression of adaptive genes and adaptive hetrospecific pairing by late breeding collared flycatcher females may work in the opposite direction.

Eco-evolutionary dynamics integrates the reciprocal interactions betweenecology and evolution. These two branches of biology traditionally assumethe other as static for simplicity. However, increasing evidence shows thatthis simplification may not always hold because ecology and evolution canoperate in similar timescales. This thesis theoretically explores how thereciprocal interactions may influence ecological and evolutionary outcomesin four different eco-evolutionary contexts.Many species of non-social animals live in groups. Aggregating ingroups often has both benefits and costs that depend on group size. Thanksto the benefits of aggregation, population growth likely depends positivelyon population density when it is small. This phenomenon, the Allee effect,has been hypothesized to explain the evolution of aggregation behavior. Ifind that the Allee effect alone does not lead to the evolution whenpopulation dynamics is explicitly accounted for. Some other mechanisms,such as frequent needs for colonizing new patches or anti-aggregation,should be invoked to explain why aggregation behavior could evolve.Phenotypic plasticity is the ability of a genotype to express distinctphenotypes when exposed to different environments. Although it is oftenshown to be adaptive and not costly, highly plastic organisms are rare. Paststudies demonstrated some potential reasons. I test another possibility; costsmay arise from sexual selection because highly plastic individuals may beless preferred as a mate. I show that, even in the absence of the direct cost ofplasticity, the level of plasticity remained low at intermediate strengths ofassortative mating. This pattern is robust across wide ranges of parametervalues.Ecological speciation occurs when ecologically divergent selectionbetween environments causes reproductive isolation between divergingsubpopulations. Several verbal models of ecological speciation emphasizethe roles of phenotypic plasticity in promoting speciation. The complexprocesses involved in speciation, however, are difficult to be evaluated byverbal accounts. I quantitatively test the proposed idea in a mechanisticmodel of ecological speciation in the presence and absence of plasticity. Ifind conditions under which plasticity can promote or hinder ecologicalspeciation. Plasticity facilitates speciation by producing a gap in thedistributions of expressed phenotypes, which serves as a barrier to gene flowin an assortatively mating population.Ecosystem ecology and evolutionary biology are the least integratedfields in ecology and evolution. Natural selection operating at the individuallevels on traits governing ecosystem functions may affect ecosystemproperties, which may feedback to individuals. I reviewed this idea anddemonstrate the feedback loop by using a simple consumer-resource model.

Sex ratio theory suggests that the strength of intersexual selection will increase as a population more male-biased; reflecting increased selectivity in mate choice. Populations of pond turtle have varying adult sex ratios, in painted turtles (Chrysemys picta), reported sex ratios range from female biased (1:3) to male biased (3:1). I investigated the effect of sex ratio on male reproductive success (quantified as “fertilization success”) in painted turtles. I examined the mating system of painted turtles in a female-biased population using microsatellite paternity analysis, relating variation in male fertilization success to male phenotype and offspring survival, employed ex situ behavioural observation to clarify the mechanism behind the variation in male fertilization success and used agent-based modeling to simulate the effects of changing sex ratio, population density and proportions of male phenotype on male fertilization success. Small males contributed sperm to a greater number of clutches than did larger males, but were not more likely to reproduce in a season than larger males. There was no offspring fitness advantage related to male body size and no relationship between male claw length and fertilization success. Large male painted turtles courted at a higher frequency than small males. I found no relationship between male courtship behavior and claw length. Females showed no preference for males of any phenotype. Agent-based simulations were based on the distribution of best fit from the observed data; an amalgam of two Poisson distributions, each with its own probability of success and proportional representation in the final distribution. Increased female sex ratio bias, increased population density and increased proportions of “more successful” males all increased the mean and variance of male fertilization success, based on increased encounter rate among turtles. Small and large male painted turtles enjoy different fertilization success. It is uncertain whether this difference is based on active female choice, cryptic female choice, sperm competition or a combination of factors. Sex ratio simulations predict the opposite result as that predicted by sex ratio theory. These contrary results should be compared to simulations manipulating choosiness and field data from painted turtle populations to clarify mechanisms influencing male reproductive success.

Ecological speciation involves the evolution of reproductive isolation (RI) as a by-product of adaptation to different selective environments. Parallel patterns of non-random mating by environment type provide strong evidence that ecological speciation has occurred. The processes involved in the origins of RI are more difficult to detect however. One mechanism involves the correlated evolution of mating preferences and sexually selected traits. I developed a conceptual model for detecting RI under various scenarios of mate preference evolution. The model predicts that RI will not evolve if preferences are evolutionarily constrained relative to the preferred traits, but is detectable as long as preferences evolve in parallel. I then applied this framework to an empirical system with populations of guppies (Poecilia reticulata) adapted to low- and high-predation environments. I measured female mate preferences for male colour and size; traits which are divergent between the two environment types. Preference functions for colour also diverged in the predicted direction. The parallel pattern of preference divergence suggests that divergent natural selection from predators may be contributing to RI between guppy populations.

Female reproductive decisions have been suggested to be highly influenced by mate quality. I have studied whether offspring quality may be adjusted by females to match the attractiveness of males and how strong control females have over their reproductive investment focusing on egg size. This was done in the Banggai cardinalfish (Pterapogon kauderni), a sex-role reversed obligate paternal mouthbrooder where males invest heavily into reproduction. As this species is suitable for both laboratory experiments and field studies it is an ideal candidate for the study of reproductive investment. Mating was size-assortative and both males and females benefited from pairing with large partners. However, male size determined the reproductive output of a pair. Females courted large males more intensively and produced larger, but not fewer eggs when mated to large males as compared to small males. Further, this matching of egg size to mate attractiveness may be fast. Female courtship behaviours contained honest information regarding both clutch weight and egg maturity, traits that may be highly important for male mate choice. Surprisingly, males played an important part in territory defence suggesting relatively equal sex-roles in this species. Also, this species showed stable group structures which may be important for the evolution of female plasticity in reproductive investment due to high variance in quality of available mates. This thesis suggest that females have a remarkable control over their reproductive investments and that male quality may be highly influential on reproductive decisions regarding offspring quality. Furthermore, it suggest that sexual selection may have strong effects on the evolution of egg size and parental care on a whole.

Sexual selection has long been a working explanation for the elaboration of appreciable traits in plants and animals, but the idea that it is an equally potent agent of change at the level of individual molecules is relatively recent. Indications that genes associated with reproductive biology evolve especially rapidly planted this notion, but many details about the genomics of sex remain elusive. Numerous studies have characterized rapid sequence and expression divergence of sex-related molecules, but few if any have demonstrated convincingly that these patterns exist as a result of sexual selection. This dissertation describes several genome-scale studies related to reproduction and the sexes in teleost fishes, a group of animals underexploited in regard to this topic. Using commercial microarrays I measured the extent of sexually dimorphic gene expression in the zebrafish, Danio rerio. Sex-biased patterns of gene expression in this species are similar to those described in other animals. A number of genes expressed at high levels in ovaries and testes relative to the body were identified as a product of the study, and these data may be useful for future studies of reproductive genes in Danio fishes. In a second study, the recent advent of high throughput cDNA pyrosequencing was leveraged to characterize the relationships between tissue-, sex-, and species-specific expression patterns of genes and rates of sequence evolution in swordtail fishes (Xiphophorus). I discovered ample evidence for expression biases of all three types, and a generally positive but idiosyncratic relationship between the magnitude of expression bias and rates of protein-coding sequence evolution. Pyrosequencing of cDNA was also used to explore the possibility that postcopulatory sexual selection drives the rapid evolution of male pregnancy genes, a novel class of reproductive molecules unique to syngnathid fishes (seahorses and pipefishes). Genes differentially expressed in the male brooding tissues as a function of pregnancy status evolve more rapidly at the amino acid level than genes exhibiting static expression. Brooding tissue genes expressed during male pregnancy have evolved especially rapidly in polyandrous lineages, a finding that supports the hypothesized relationship between postcopulatory sexual selection and the adaptive evolution of reproductive molecules.