Inbreeding can have profound negative effects on individuals by reducing fertility and viability. In populations, inbreeding depression can reduce growth rates and increases extinction risk. The aims of this thesis are to investigate inbreeding depression in male guppies (Poecilia reticulata) and to study the evolution of mechanisms for inbreeding avoidance in females, using guppies from a feral population in Queensland, Australia. Male guppies are highly polymorphic in their sexual ornamentation, indeed they show one of the most extreme polymorphisms observed in nature. Female guppies exhibit complex mate choice based on preferences for ornamentation, as well as social context. I aim is to examine how these factors of inbreeding avoidance alter sexual selection. In male guppies I found strong inbreeding depression in male sperm numbers, which is amplified under semi-natural compared to laboratory conditions (Chapter 2). Moreover, inbreeding depression results in low fertility under sperm competition: an experiment using artificial insemination techniques reveals that highly inbred males are heavily disadvantaged in gaining paternity (Chapter 3). On population level, inbreeding depression is manifest in reduced growth rates, predominantly in the early stages of inbreeding (Chapter 4). Population growth at inbreeding coefficients f=0.375-0.59 did not seem to lead to inbreeding depression, whereas lower levels of inbreeding reduced population growth. Although the growth rates in inbred populations appear normal, severe inbreeding depression is uncovered after outbred immigrants are added. Specifically, male immigrants are most efficient in short-term genetic rescue, probably due to insemination of large numbers of females whereas females are limited in the number of eggs they can produce (Chapter 4). Male ornamental traits show significant inbreeding depression in semi-natural conditions only (Chapters 2 & 3). Inbreeding avoidance mechanisms seem to have evolved in females: they prefer courtship displays of non-inbred males (Chapter 2), unfamiliar males (Chapter 5) and males with rare patterns (Chapter 6). This preference might increase the mating success of immigrants, and may have evolved to facilitate the avoidance of inbreeding. Together with context-independent preferences for ornament combinations (Chapter 6), it also offers an explanation for the maintenance of polymorphism in this species.

Inbreeding can have profound negative effects on individuals by reducing fertility and viability. In populations, inbreeding depression can reduce growth rates and increases extinction risk. The aims of this thesis are to investigate inbreeding depression in male guppies (Poecilia reticulata) and to study the evolution of mechanisms for inbreeding avoidance in females, using guppies from a feral population in Queensland, Australia. Male guppies are highly polymorphic in their sexual ornamentation, indeed they show one of the most extreme polymorphisms observed in nature. Female guppies exhibit complex mate choice based on preferences for ornamentation, as well as social context. I aim is to examine how these factors of inbreeding avoidance alter sexual selection. In male guppies I found strong inbreeding depression in male sperm numbers, which is amplified under semi-natural compared to laboratory conditions (Chapter 2). Moreover, inbreeding depression results in low fertility under sperm competition: an experiment using artificial insemination techniques reveals that highly inbred males are heavily disadvantaged in gaining paternity (Chapter 3). On population level, inbreeding depression is manifest in reduced growth rates, predominantly in the early stages of inbreeding (Chapter 4). Population growth at inbreeding coefficients f=0.375-0.59 did not seem to lead to inbreeding depression, whereas lower levels of inbreeding reduced population growth. Although the growth rates in inbred populations appear normal, severe inbreeding depression is uncovered after outbred immigrants are added. Specifically, male immigrants are most efficient in short-term genetic rescue, probably due to insemination of large numbers of females whereas females are limited in the number of eggs they can produce (Chapter 4). Male ornamental traits show significant inbreeding depression in semi-natural conditions only (Chapters 2 & 3). Inbreeding avoidance mechanisms seem to have evolved in females: they prefer courtship displays of non-inbred males (Chapter 2), unfamiliar males (Chapter 5) and males with rare patterns (Chapter 6). This preference might increase the mating success of immigrants, and may have evolved to facilitate the avoidance of inbreeding. Together with context-independent preferences for ornament combinations (Chapter 6), it also offers an explanation for the maintenance of polymorphism in this species.

<p>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, <i>Gallus</i> <i>gallus ssp.</i>, and its domestic subspecies the domestic fowl, <i>Gallus gallus</i> <i>domesticus</i>. 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.</p>

<p>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. </p><p>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. </p><p>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.</p>

<p>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 (<i>Ficedula hypoleuca</i>) and the collared flycatcher (<i>F. albicollis</i>). 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).</p><p>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.</p>

<p>Chemical signals are the most widely used form of sexual communication throughout the living world. However, there is in general little knowledge about what these signals actually communicate. The role of chemical signals, i.e. pheromones, in sexual behaviour has traditionally been seen as restricted to mate attraction and species recognition. This thesis reviews the evidence for pheromones as indicators of mate quality, and then investigates three important factors for mate quality signals – individual variation, heritability and cost – by using the male pheromone of the lekking fruit fly <i>Drosophila grimshawi</i> as a model.</p><p>The experiments presented indicate that the pheromone of <i>D. grimshawi</i> has multiple functions, and that these functions vary with social context. Thus, with regard to females, the pheromone seems to act mainly as a species/mate recognition signal, since females show little preference for the amount of pheromone deposited by a male. Moreover, males invest less in pheromone production when subjected to females as compared to when subjected to rival males. However, the pheromone seems to be costly in production since males that invest much in pheromone deposition has a shorter lifespan. This suggests a function for the pheromone in male-male interactions. Males can distinguish their own pheromone depositions from those of a strange male, and also discriminate between pheromone depositions from one and two strange males. This might give them the ability to assess the size of a lek and the competitive capacities of rivals, information that should be useful when optimizing sexual behaviour.</p><p>In conclusion, the pheromone seems to act as an honest mate/competitor quality signal in some social contexts, and as a non-costly species/mate recognition signal in other. In addition, I show for the first time that a chemical signal has differential fitness costs, and that an insect is able to distinguish between individual odour signatures.</p>

<p>This thesis focuses on the evolutionary interactions between birds and a parasite, the mosquito-borne Sindbis virus (Togaviridae, <i>Alphavirus</i>). In conclusion, the results show that the Sindbis virus is widespread among birds, and that the fitness of infected hosts may be reduced by the virus. Furthermore, viruclearance ability was revealed by male plumage traits, and viraemia was related to hormonal- and social status.</p><p>The distribution of Sindbis virus infections among passerine birds was examined in five areas in Sweden. Almost all species tested were infected, and three species of thrushes weridentified as the main hosts. In a series of experimental infections, greenfinches (<i>Carduelis chloris</i>) kept in aviaries were used ahosts. First, the behavioural consequences of an infection were investigated. During the infection, birds tended to reduce thespontaneous locomotion activity, and when escaping from a simulated predator attack, infected birds had reduced take-off spee Furthermore, when comparing virus clearance rate between male greenfinches, I found that males with large yellow tail ornaments hafaster virus clearance rates as compared to those with smaller ornaments. Thus, male virus clearance ability was honestly revealed by the size of an ornament. Moreover, males with experimentally elevated testosterone levels experienced a delayed, but not increased viraemia as compared to controls. When the relationship between male social ranand viraemia was examined, I found no evidence that high-ranked males suffered reduced rank during the infection. Nevertheless, viraemipatterns of males were related to their social rank, so that low-ranked birds had a delayed viraemia as compared to high-ranked birds. </p>

This thesis focuses on the evolutionary interactions between birds and a parasite, the mosquito-borne Sindbis virus (Togaviridae, Alphavirus). In conclusion, the results show that the Sindbis virus is widespread among birds, and that the fitness of infected hosts may be reduced by the virus. Furthermore, viruclearance ability was revealed by male plumage traits, and viraemia was related to hormonal- and social status. The distribution of Sindbis virus infections among passerine birds was examined in five areas in Sweden. Almost all species tested were infected, and three species of thrushes weridentified as the main hosts. In a series of experimental infections, greenfinches (Carduelis chloris) kept in aviaries were used ahosts. First, the behavioural consequences of an infection were investigated. During the infection, birds tended to reduce thespontaneous locomotion activity, and when escaping from a simulated predator attack, infected birds had reduced take-off spee Furthermore, when comparing virus clearance rate between male greenfinches, I found that males with large yellow tail ornaments hafaster virus clearance rates as compared to those with smaller ornaments. Thus, male virus clearance ability was honestly revealed by the size of an ornament. Moreover, males with experimentally elevated testosterone levels experienced a delayed, but not increased viraemia as compared to controls. When the relationship between male social ranand viraemia was examined, I found no evidence that high-ranked males suffered reduced rank during the infection. Nevertheless, viraemipatterns of males were related to their social rank, so that low-ranked birds had a delayed viraemia as compared to high-ranked birds.

Chemical signals are the most widely used form of sexual communication throughout the living world. However, there is in general little knowledge about what these signals actually communicate. The role of chemical signals, i.e. pheromones, in sexual behaviour has traditionally been seen as restricted to mate attraction and species recognition. This thesis reviews the evidence for pheromones as indicators of mate quality, and then investigates three important factors for mate quality signals – individual variation, heritability and cost – by using the male pheromone of the lekking fruit fly Drosophila grimshawi as a model. The experiments presented indicate that the pheromone of D. grimshawi has multiple functions, and that these functions vary with social context. Thus, with regard to females, the pheromone seems to act mainly as a species/mate recognition signal, since females show little preference for the amount of pheromone deposited by a male. Moreover, males invest less in pheromone production when subjected to females as compared to when subjected to rival males. However, the pheromone seems to be costly in production since males that invest much in pheromone deposition has a shorter lifespan. This suggests a function for the pheromone in male-male interactions. Males can distinguish their own pheromone depositions from those of a strange male, and also discriminate between pheromone depositions from one and two strange males. This might give them the ability to assess the size of a lek and the competitive capacities of rivals, information that should be useful when optimizing sexual behaviour. In conclusion, the pheromone seems to act as an honest mate/competitor quality signal in some social contexts, and as a non-costly species/mate recognition signal in other. In addition, I show for the first time that a chemical signal has differential fitness costs, and that an insect is able to distinguish between individual odour signatures.

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.