Male pregnancy is a complex and energetically costly form of male parental care found exclusively in the fish family Syngathidae, which includes pipefishes, seahorses, and seadragons. The novelty of the male brood pouch raises questions about how this trait evolved and what role it plays in sexual selection. One possibility is that brood pouch evolution was partially shaped by postcopulatory sexual selection. We used the sexually dimorphic Gulf pipefish, Syngnathus scovelli, to test for postcopulatory sexual selection within broods and tradeoffs between successive male pregnancies as a function of female attractiveness. Offspring survivorship within a pregnancy was affected by the size of a male's mate, the number of eggs transferred, and the male's mating history. These results indicate the males invest more resources into broods from large, attractive mates to the detriment of future broods. Next, we investigated the effects of food limitation on male parental care strategies. Our data suggests that male Gulf pipefish sacrifice investment in future reproduction, via somatic growth, in favor of current reproduction. A positive relationship between number of failed eggs and male growth rate in our low-food treatments suggests that males may derive an energetic benefit from unsuccessful eggs in the brood pouch. Finally, we used a paired design, to investigate the effect of perceived female attractiveness on offspring survivorship. We found that, in general, males prefer the largest female available. Within a replicate, we found that the male that preferred their mate more, regardless of stimulus female size, also had higher offspring survivorship in the resulting brood. This result is exciting because it shows not only that cryptic male choice affects offspring survivorship in Gulf pipefish but also that mate preference can have a prolonged effect on an individual’s reproductive success. Ultimately, our data suggests that the evolution of the brood pouch has produced a trait that not only nurtures the offspring but also exert cryptic male choice.

Directional selection is prevalent in nature yet phenotypes tend to remain relatively constant, suggesting a limit to trait evolution. The genetic basis of evolutionary limits in unmanipulated populations, however, is generally not known. Given widespread pleiotropy, opposing selection on a focal trait may arise from the effects of the underlying alleles on other fitness components, generating net stabilizing selection on trait genetic variance and thus limiting evolution. Here, I look for the signature of stabilizing selection for a suite of cuticular hydrocarbons (CHCs) in Drosophila serrata. Despite strong directional sexual selection on CHCs, genetic variance differed between high and low fitness individuals and was greater among the low fitness males for seven of eight CHCs. Univariate tests of a difference in genetic variance were non-significant but have low power. My results implicate stabilizing selection, arising through pleiotropy, in generating a genetic limit to the evolution of CHCs in this species.

Sexual selection drives the evolution of male morphology, life history, physiology, and behaviour across taxa. Here I examine the mechanisms of sexual selection that arise at various stages in mating interactions to identify congruence or conflict between the traits selected by choice and competition. I first examine plasticity of developing male Australian redback spiders (Latrodectus hasselti) and show that male metabolic rates vary adaptively to facilitate the scramble to reach virgins. Next, I show that females cease sex pheromone production after mating and re-advertise receptivity later in their reproductive season effectively creating two windows in which males may compete. I show that females discriminate against males that do not meet a threshold courtship duration suggesting that courtship is the trait selected through choice. However, male-male competition leads to reductions in courtship effort provided to females. During the first window paternity is split equally if rival males mate in quick succession with a virgin female. However, if the second mating is delayed, there is a strong bias in the paternity of the second male. A delay in the second mating is beneficial to females as it reduces longevity costs of polyandry. However, delays in the initial mating decrease female longevity, likely because of elevated metabolic rates of virgins. My research shows that the trait favoured by female choice is in opposition to selection via male-male competition. Females’ sex pheromone production yields windows during which mating will optimize female, but not male, fitness. Studies that isolate the mechanisms of sexual selection are valuable in that they can identify the traits under selection. However, my research shows that considering these processes in isolation can lead to incorrect inferences about the net effect of sexual selection.

Sexual selection drives the evolution of male morphology, life history, physiology, and behaviour across taxa. Here I examine the mechanisms of sexual selection that arise at various stages in mating interactions to identify congruence or conflict between the traits selected by choice and competition. I first examine plasticity of developing male Australian redback spiders (Latrodectus hasselti) and show that male metabolic rates vary adaptively to facilitate the scramble to reach virgins. Next, I show that females cease sex pheromone production after mating and re-advertise receptivity later in their reproductive season effectively creating two windows in which males may compete. I show that females discriminate against males that do not meet a threshold courtship duration suggesting that courtship is the trait selected through choice. However, male-male competition leads to reductions in courtship effort provided to females. During the first window paternity is split equally if rival males mate in quick succession with a virgin female. However, if the second mating is delayed, there is a strong bias in the paternity of the second male. A delay in the second mating is beneficial to females as it reduces longevity costs of polyandry. However, delays in the initial mating decrease female longevity, likely because of elevated metabolic rates of virgins. My research shows that the trait favoured by female choice is in opposition to selection via male-male competition. Females’ sex pheromone production yields windows during which mating will optimize female, but not male, fitness. Studies that isolate the mechanisms of sexual selection are valuable in that they can identify the traits under selection. However, my research shows that considering these processes in isolation can lead to incorrect inferences about the net effect of sexual selection.

Directional selection is prevalent in nature yet phenotypes tend to remain relatively constant, suggesting a limit to trait evolution. The genetic basis of evolutionary limits in unmanipulated populations, however, is generally not known. Given widespread pleiotropy, opposing selection on a focal trait may arise from the effects of the underlying alleles on other fitness components, generating net stabilizing selection on trait genetic variance and thus limiting evolution. Here, I look for the signature of stabilizing selection for a suite of cuticular hydrocarbons (CHCs) in Drosophila serrata. Despite strong directional sexual selection on CHCs, genetic variance differed between high and low fitness individuals and was greater among the low fitness males for seven of eight CHCs. Univariate tests of a difference in genetic variance were non-significant but have low power. My results implicate stabilizing selection, arising through pleiotropy, in generating a genetic limit to the evolution of CHCs in this species.

<p>A long-held assumption in animal behavior is that females and males differ fundamentally in their mating strategies. Females are thought to be more choosy because female reproduction typically is limited by parental investment. Males, on the other hand, are expected to compete among themselves for access to females or resources, since male reproduction is limited primarily by mating access. This dichotomy is challenged by the increasing realization that males can be choosy and females also compete aggressively. It remains unclear, however, if and how selection acts on aggressive behavior in the context of intrasexual competition among females (reviewed in Chapter 1). In this thesis, I use a population of free-living tree swallows (<i>Tachycineta bicolor<i>) to test predictions about the selective pressures shaping aggressive behavior in females. First, using an experimental manipulation of nest site availability, I demonstrate that more aggressive females have a competitive edge in acquiring nestboxes, a critical limiting resource required for breeding (Chapter 2). This result shows that more aggressive females are more likely to breed and, thus, that females experience direct selection to be aggressive in the context of competition for mating opportunities. Next, I demonstrate a fitness cost of female aggression (Chapter 3): high levels of aggression in females are not associated with the quantity of offspring, but instead, more aggressive females had offspring of lower quality (i.e. reduced mass). Using a cross-fostering approach, I explore the causal link between female aggression and offspring mass, and I find that a trade-off between female aggression and maternal care best accounts for this cost of aggression. Site differences may create variation in how selection shapes female aggression, but the overall finding that more aggressive females have lower quality offspring indicates that this cost may work counter to selection favoring aggressive behavior in the context of competition over nestboxes. Understanding the evolution of female aggressiveness in a biparental system is incomplete without examining how males may alter the selective environment shaping female behavior. In Chapter 4, I explore the potential role of a female's mate in offsetting the costs of aggression. Males appear to mitigate these costs for their female partners, but not by compensating for poor parenting by aggressive females. Instead, females invest more heavily in reproduction, laying more and larger eggs, when mated to a male that is more different from her own phenotype. If this differential investment outweighs the cost of aggressiveness in terms of offspring quality, then male phenotype may play a key role in understanding the selective pressures shaping the evolution of aggressive behavior in females. Altogether, this dissertation explores the costs and benefits of female aggressive behavior. The focus on aggressiveness as a sexually selected trait in females provides a much needed parallel to the wealth of information already known about the selective pressures shaping sexually selected traits in males.</p> / Dissertation

Understanding the processes that govern mating behaviors is a fundamental goal of evolutionary biology and behavioral ecology. Population-level patterns of mate acquisition and offspring production, otherwise known as the genetic mating system, play a central role in the sexual selection on morphological and behavioral traits and may facilitate speciation. The central hypothesis of this research is that variation in environmental conditions, such as temperature, turbidity, and habitat, and demographic influences such as population density, sex ratios and temporal availability of mates, may limit mating and reproductive success in a predictive manner. Therefore the goal of this dissertation is to examine the contributions of geographic and temporal variation on the plasticity of the genetic mating system in two species of pipefish. The first study examined whether meaningful variation in the genetic mating system exists between two natural populations of the dusky pipefish, Syngnathus floridae. Results of this investigation provide evidence that the genetic mating system differs among different geographic locations. The second study considered the relative contributions of environmental conditions and population demographics on differences in the genetic mating system of dusky pipefish from five natural populations. The results of this investigation show strong trends for demographic and environmental factors to strongly influence the genetic mating system between populations. The third study considered how variation in the number of available mates predicts the outcome of sexual selection during the course of a breeding season in the broad-nosed pipefish, Sygnathus typhle. The results of this study indicate a strong influence of the operational sex ratio on the genetic mating system. In addition to these studies, a study was conducted to investigate whether phylogeographic relationships may be responsible for geographic variation in the genetic mating system of the dusky pipefish of pipefish. Mitochondrial DNA analysis does not substantiate subspecies designations for this species and microsatellite analysis show a clear pattern of isolation by distance. Taken together, these studies significantly enhance the understanding of how mating systems are organized over broad environmental gradients and temporal/spatial scales and to the evolution of sexual selection on the whole.

Marine and aquatic species relying on external fertilization must have special adaptation against sperm-dilution in the water. In addition mating competition may be another important source of limitation to an individual¡¦s reproductive success. If the two above mechanisms are in direct conflict, real adaptation would reflect whichever is more critical to one¡¦s fitness. In using the same logic, the unidirectional induction of spawning, i.e., females inducing males, but not males inducing females, has been reported as an evidence for the relative importance of natural selection, that is, sperm-dilution, in limiting the fitness of brittlestars in shallow waters. If the above deduction is correct, one would predict more adaptations to offset the adverse effect of sperm-dilution in various aspects of spawning-related characteristics. Two species of brittlestars. Ophiocoma dentata, living in subtidal zone, and O. scolopendrina, living in intertidal zone, were studied at Kenting, southern Taiwan. Experimental approaches were adopted to figure out if these two species have any adaptation in space and in time that can perceivably counteract the effect of sperm dilution. Spatially, the male O. dentata has the ability to search for females, and the males living with females have higher rates of responding to female spawn than unpaired males; the males do not actively search for eggs released in the water. In O. scolopendrina, neither male nor female had the ability to distinguish the sex of other individuals, they do not form pairs in nature. Males, however, have strong ability to search released eggs and then approach and spawn near the eggs. Temporally, the males of the O. scolopendrina, are sensitive to tidal rhythms in terms of inducibility by female spawns, they only spawn at the time of low tide. The possible effect of water level and photophase were both ruled out. In contrast, no such tidal rhythms of male inducibility was found in the subtidal O. dentata. Sexual selection needs not be invoked in any of the above behaviors, whereas natural selection against sperm dilution is an acceptable explanation. Last, the male O. scolopendrina displays interference behavior by blocking other males from approaching eggs are in the water. The blocked individuals are less likely to spawn.

No description available.

Thesis (Ph. D.)--University of Chicago, Faculty of the Division of the Biological Sciences and the Pritzker School of Medicine, Department of Ecology and Evolution, June 2001. / Includes bibliographical references. Also available on the Internet.