<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>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>

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.

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.

Natural populations exhibit a non-random mating behavior and it is assumed that mate preferences causing the non-random mating behavior play a role in sympatric speciation. In my thesis, I have proposed a new model of a mate preference that is based on ecological performance and I have named it fitness-based mating. Individuals that express this mate preference choose primarily fit partners. Fitness-based mating is modelled for haploid, diallelic populations. Individuals are distributed across two niches, and genomes are simplified to two loci. The first locus is subject to natural selection, and the second-locus genotype gives the strength of the mate preference. The population is separated into females and males, among which only females exhibit the mate preference. Ecological selection acts on both sexes alike. With the model I have investigated how female choosiness based on direct advantages offered by their partners can cause and maintain a polymorphic population. Fitness-based mating is an evolutionary successful mating strategy. It spreads in a population due to its amplifying effect on the reproductive success and on the attractiveness of its carriers. A polymorphism arises naturally in the model. The emergence of a stable polymorphism of traits underlying ecological selection is of special interest, because a polymorphism can be a precursor of speciation.

Sexual selection is responsible for a great diversity of elaborate male traits. A general female preference for males that have exaggerated traits drives this process, but the reasons females exhibit this preference are often unclear. Recent advances in understanding signal evolution have emerged from studies of receiver psychology that focus on how receivers perceive and process communication signals. I apply the perspective of receiver psychology to understand female preference for elaborate signals in túngara frogs (Physalaemus pustulosus). Male túngara frogs produce advertisement calls of variable complexity. Females exhibit a strong preference for complex to simple calls, but previous studies have not found consistent patterns of preference between calls of variable complexity. In my doctoral research, I investigate the function of variable complexity in túngara frogs. Specifically, I address the following questions: 1) Are calls of variable complexity especially relevant to females in certain contexts? Do males respond to female behavior by increasing their production of complex calls? 2) Does male to female proximity influence female response to call complexity? 3) Are females constrained by their perceptual biology in discriminating differences in call complexity? 4) Can females remember attractive males over silences between bouts of advertising? Is working memory for attractive males dependent upon signal complexity? And 5) Does signal memorability increase with signal complexity in a linear relationship? These studies provide several new perspectives to an understanding of female preference for elaborate signals. Phonotaxis experiments demonstrate that females use elicitation behaviors to influence male production of complex calls, that proximity influences female response to signal elaboration, that females are constrained by their perceptual biology in discriminating between complex calls, that memory can influence the evolution of signal complexity, and that memorability and signal complexity share a non-linear relationship. / text

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.

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.

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