Sturgeon species (order Acipenseriformes) are prone for interspecific hybridization. Anthropogenic activities in river basins influence sturgeon reproduction by destruction of their natural spawning grounds. Consequently, spawning areas, as well as the time of spawning of sturgeon species overlap and different sturgeon species reproduce concurrently. This increases the probability of meeting of heterospecific gametes and pre-zygotic postcopulatory reproductive barriers, comprising of sperm competition and cryptic female choice, may play an important role in preventing undesirable interspecific hybridization. The goal of this study was to evaluate the role of interspecific sperm competition and cryptic female choice during interspecific hybridization of sterlet (Acipenser ruthenus) and Siberian sturgeon (Acipenser baerii). Reproductive characteristics (fertilization rate and hatching rate) were described in each of experimental and control groups showing similar values for competitive and non-competitive trials. Parentage assignment was performed in hatched larvae using combination of mitochondrial DNA and microsatellite DNA markers. Obtained results revealed higher fertilization success of sterlet spermatozoa, when these competed for fertilization with spermatozoa of Siberian sturgeon. Total reproductive success of starlet spermatozoa was 78.9 % and Siberian sturgeon 21.1 %. Contrary, when spermatozoa did not compete for fertilization, males of analysed species showed equal fertilization success. In the trials, where eggs of both studied species were mixed and fertilized by sperm from each species separately, eggs of any species did not show a tendency to bias fertilization by spermatozoa of conspecific males. Probably, there are no pre-zygotic postcopulatory reproductive barriers that prevent interspecific hybridization of sterlet and Siberian sturgeon at the gametic level.

A critical mechanism underpinning current biological diversity is the extent to which one species mates with, or avoids mating with, another. However, little is known about the factors that mediate hybridisation, especially during the initial and rarely observed stages of secondary contact when interspecific interactions have not responded to selection. In particular, whilst hybridisation is ultimately a behavioural phenomenon, the role of behaviour in mediating hybridisation and how it is influenced by environmental and circumstantial factors is rarely investigated. Recently introduced species provide us with unequalled opportunities to study these factors. In this thesis I examine the role of behavioural mechanisms, in particular male-male competition and mate choice, in mediating mating patterns between two genetically and phenotypically distinct lineages of European wall lizard (Podarcis muralis) that have come into recent secondary contact through human introductions. In Chapter Two, I investigated how sexual selection during allopatry is responsible for creating stark differences in phenotypic traits such as body size and weapon performance evident in the two lineages today, ultimately explaining the strong biases in dominance during territorial disputes between males. However, I also show that even given this asymmetry in male competitive ability, the extent to which it extrapolates into greater access to females in naturalistic, outdoor enclosures depends strongly on the spatial clustering of basking sites, a critically important resource for many ectotherms. In contrast to initial predictions suggested by asymmetries in male competition outlined in the previous chapter, in Chapter Three I show that both paternity and courtship behaviour was strongly assortative in the outdoor enclosures. Further investigation through staged experiments on olfactory mate choice, mating trials and analyses on specific behavioural data obtained in an enclosure experiment, I show that lineage based dominance actually contributes to assortative mating patterns in conjunction with weak conspecific male choice. In contrast, female choice seems to play no role in mediating the mating patterns observed between the two lineages. In Chapter Four I had the rare opportunity to examine the morphological and behavioural factors that predict why animals should hybridise in the first place, using the data obtained in the enclosure experiment above. I found that hybridisation was particularly common between small individuals of the larger lineage and large individuals of the smaller lineage; a result that corroborates the mechanisms determining the assortative patterns uncovered in Chapter Three. Additionally, hybridisation rates were particularly high in less dominant individuals, which I suggest is due to subordinate males having reduced opportunities for courting conspecific females due to male-male competition, requiring them to become less ‘choosy’ and therefore more likely to mate with heterospecifics. Finally, secondary contact cannot occur without at least one lineage coming into a new environment, and yet relatively little attention is paid to how this environmental change can affect the signals involved in intraspecific communication and mate choice. In Chapter Five I show that a change in the amount of time male lizards spend thermoregulating (a likely consequence of arriving in a new environment) significantly changes the chemical composition of their scent marks. However, whilst female lizards were able to detect these effects, they did not seem to base their mating decisions on them. Nevertheless, this result raises interesting questions about the potential function and consequences of this plasticity, and highlights the importance of considering plasticity in chemical communication in heterogeneous environments. Overall, this thesis shows the critically important role of behaviour in mediating intra- and interspecific mating patterns during recent secondary contact. In particular, it highlights how the direction and extent of hybridisation and competition are influenced by the degree to which differing morphological and behavioural phenotypes interact over a heterogeneous environment, particularly during the initial stage of secondary contact when mate choice has not had the chance to respond to the selective pressures of hybridisation.

Sexual selection plays an important role in the evolution of animals. Today we already know that it takes place not only before copulation (precopulatory sexual selection), but also after copulation. This type of sexual selection is called postcopulatory sexual selection, and occurs mainly in promiscuous species, where females mate with multiple males. In this case, sperm competition occurs in the female reproductive tract. To increase the likelihood of their reproductive success, males began to develop surprisingly diverse sperm adaptations at the morphological, physiological, or behavioural levels. These adaptations often affect sperm velocity (motility), which is a key factor for successful egg fertilization. However, the result of reproductive success can also be influenced by females, who may prefer sperm with a certain phenotype in the process of cryptic female choice, and thus, for example, obtain better genes for offspring. In some species, females even can have the ability to sort and store sperm in specialized organs in which the sperms are nourished for some time, and then used to fertilize the egg. Even in this case, the storage of sperm is often affected by sperm morphology. Due to these mechanisms of postcopulatory sexual selection, sperm are under strong selection pressure, which can...

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Postcopulatory sexual selection—sperm competition and cryptic female choice—has become a major area of research over the past 40 years. Within this field there are many outstanding questions at every level of analysis, from proximate to ultimate. The fitness consequences for both sexes in the period after copulation and before fertilization are considerable, but are obscured within the female reproductive tract. Our understanding of postcopulatory mechanisms is especially sparse in taxa other than birds and insects. Nearly nothing is known in reptiles except that multiple paternity is common and widespread, and often results from long-term sperm storage across breeding seasons. We present some of the very first data on the determinants of fertilization success in the context of sperm competition in reptiles, a group that accounts for 30% of terrestrial vertebrates. In the first chapter, "Asymmetric gametic isolation between two populations of red-sided garter snakes", we discuss the use of between-population crosses to reveal gametic isolation. The effect of population density and operational sex ratios on mating systems and the speciation process has fueled theoretical debate. We attempted to address these issues using two populations of red-sided garter snakes (Thamnophis sirtalis parietalis) from Manitoba, Canada. Our study populations differ markedly in their density mating aggregations, with a 10-fold difference between them. Using microsatellite markers for paternity analysis of litters produced from within and between population crosses. We found that the population with highest aggregation density, and presumably with the highest level of sexual conflict (i.e., when the evolutionary interests of the sexes differ) over mating, was also the population that exhibited homotypic sperm precedence. The less dense population showed a distinct postcopulatory male-size advantage. We also demonstrated that sperm stored within the female over hibernation can father 20-30% of offspring in a litter. In the second chapter, "Sperm competition and mate-order effects in red-sided garter snakes", we test whether females use mate-order effects to ensure that a larger (fitter) male will sire her offspring. Does that second male should have precedence in sperm competition? We tested for second-male precedence using singly-mated females that mated with a second male. Average proportion of paternity was shared equally among the first (P₁, i.e., proportion of offspring from a litter fathered by the first male to mate) and second males (P₂) to mate, and stored sperm (P[subscript ss]). This may be a case where last male precedence breaks down with more than two males. All females were spring virgins (they had not mated that spring, but may have stored sperm from fall matings); thus sperm stored presumably from fall matings is important in this system. As the interval between matings increased P₁ increased at the expense of P[subscript ss]. As the second male to mate's copulation duration increased, P₁ also increased at the expense of P₂. This last result may indicate female influence over sperm transfer during coerced matings. Copulatory plugs (CPs) are found in many taxa, but the functional significance is debated. Male garter snakes produce a gelatinous copulatory plug during mating that occludes the opening of the female reproductive tract for approximately two days. In chapter three, "Not just a chastity belt: the role of mating plugs in red-sided garter snakes revisited", we experimentally tested the role of the CPs. In snakes, sperm are produced in the testes and delivered through the ductus deferens, and the copulatory plug is thought to be produced by the sexual segment of the kidney and conveyed through the ureter. We manipulated the delivery of the two fluids separately by ligating the ducts. We confirmed that the CP is not formed in ureter-ligated males and that sperm leaks out immediately after copulation. The CP is analogous to a spermatophore. The protein matrix contains most of the sperm which are liberated as the plug dissolves within the female's vaginal pouch. One of the fundamental principles in sperm competition is that increased sperm numbers increase the odds of winning in competitions for fertilization success and males will adjust their ejaculate relative to competition and the quality of his mate. In chapter four, "Sperm depleted males and the unfortunate females who mate with them", we detect significant among-male variation in the number of sperm ejaculated, and that male mate-order reduces sperm numbers. Male sperm numbers drop significantly from one mating to the next, and this has implications for sperm competiveness, as Thamnophis sirtalis exhibits a disassociated reproductive tactic, in that sperm stores are produced outside the breeding season, and thus cannot be replenished after mating. Interestingly, however, the on average the mobility of the sperm increased for a male's second mating. Therefore, increased sperm quality may compensate for reduced numbers in a competitive context. Further, females increase their remating rate when mating with males that are unable to deliver sperm. In chapter five, "Sexual conflict during mating in red-sided garter snakes as evidenced by genital manipulation", we revisited the CP in the context of sexual conflict. Sex-differences in optimal copulation duration can be a source of conflict, as increased copulation duration may be advantageous for males as it delays female remating. Males of many species actively guard females to prevent them from remating, and in some cases males produce copulatory plugs to prevent remating. If precopulatory choice is limited at the time of her first mating, conflict may be especially onerous to a female. The size of the plug is influenced by the copulation duration. We experimentally tested the contribution of male and female control over copulation duration. We ablated the largest basal spine on the male's hemipene and found a reduction in copulation duration and an increase in the variation of plug mass. Further, we anesthetized the female's cloaca and found copulation duration increased, which suggests that males benefit from increased copulation duration while females actively try to reduce copulation duration. Therefore, sexual conflict is manifest in divergent copulation duration optima for males and females. / Graduation date: 2013