Females sample more males at high nesting densities, but ultimately obtain less attractive mates

Background: Sexual selection is largely driven by the availability of mates. Theory predicts that male competition and female choice should be density-dependent, with males competing more intensely at relatively high density, and females becoming increasingly discriminating when there are more males from whom to choose. Evidence for flexible mating decisions is growing, but we do not understand how environmental variation is incorporated into mate sampling strategies. We mimicked threespine stickleback (Gasterosteus aculeatus) breeding conditions in pools with high and low densities of nesting males and allowed females to search for mates to determine whether 1) mate search strategies change with the density of breeding males and 2) pre-copulatory components of mate choice (signalling, competition, search patterns, and mating decisions) are modified in parallel. Results: While females sampled more males at high male density, suggesting greater opportunity for sexual selection, the expanded search did not result in females choosing males with more attractive sexual signals. This is likely because red throat colouration was twice as great when half as many males competed. Instead, females chose similarly at high and low male density, using a relative strategy to compare male traits amongst potential suitors. Reduced throat colour could reflect a trade-off with costly male competition. However, we did not observe more intense competition at higher relative density. Density-dependent signalling appears largely responsible for females associating with males who have more attractive signals at low density. If we lacked knowledge of plasticity in signalling, we might have concluded that females are more discriminating at low male density. Conclusions: To understand interactions between mate choice and population dynamics, we should consider how components of mate choice that precede the mating decision interact

Females sample more males at high nesting densities, but ultimately obtain less attractive mates

Background: Sexual selection is largely driven by the availability of mates. Theory predicts that male competition and female choice should be density-dependent, with males competing more intensely at relatively high density, and females becoming increasingly discriminating when there are more males from whom to choose. Evidence for flexible mating decisions is growing, but we do not understand how environmental variation is incorporated into mate sampling strategies. We mimicked threespine stickleback (Gasterosteus aculeatus) breeding conditions in pools with high and low densities of nesting males and allowed females to search for mates to determine whether 1) mate search strategies change with the density of breeding males and 2) pre-copulatory components of mate choice (signalling, competition, search patterns, and mating decisions) are modified in parallel. Results: While females sampled more males at high male density, suggesting greater opportunity for sexual selection, the expanded search did not result in females choosing males with more attractive sexual signals. This is likely because red throat colouration was twice as great when half as many males competed. Instead, females chose similarly at high and low male density, using a relative strategy to compare male traits amongst potential suitors. Reduced throat colour could reflect a trade-off with costly male competition. However, we did not observe more intense competition at higher relative density. Density-dependent signalling appears largely responsible for females associating with males who have more attractive signals at low density. If we lacked knowledge of plasticity in signalling, we might have concluded that females are more discriminating at low male density. Conclusions: To understand interactions between mate choice and population dynamics, we should consider how components of mate choice that precede the mating decision interact

Diversification under sexual selection: the relative roles of mate preference strength and the degree of divergence in mate preferences

The contribution of sexual selection to diversification remains poorly understood after decades of research. This may be in part because studies have focused predominantly on the strength of sexual selection, which offers an incomplete view of selection regimes. By contrast, students of natural selection focus on environmental differences that help compare selection regimes across populations. To ask how this disparity in focus may affect the conclusions of evolutionary research, we relate the amount of diversification in mating displays to quantitative descriptions of the strength and the amount of divergence in mate preferences across a diverse set of case studies of mate choice. We find that display diversification is better explained by preference divergence rather than preference strength; the effect of the latter is more subtle, and is best revealed as an interaction with the former. Our findings cast the action of sexual selection (and selection in general) in a novel light: the strength of selection influences the rate of evolution, and how divergent selection is determines how much diversification can occur. Adopting this view will enhance tests of the relative role of natural and sexual selection in processes such as speciation

Diversification under sexual selection: the relative roles of mate preference strength and the degree of divergence in mate preferences

The contribution of sexual selection to diversification remains poorly understood after decades of research. This may be in part because studies have focused predominantly on the strength of sexual selection, which offers an incomplete view of selection regimes. By contrast, students of natural selection focus on environmental differences that help compare selection regimes across populations. To ask how this disparity in focus may affect the conclusions of evolutionary research, we relate the amount of diversification in mating displays to quantitative descriptions of the strength and the amount of divergence in mate preferences across a diverse set of case studies of mate choice. We find that display diversification is better explained by preference divergence rather than preference strength; the effect of the latter is more subtle, and is best revealed as an interaction with the former. Our findings cast the action of sexual selection (and selection in general) in a novel light: the strength of selection influences the rate of evolution, and how divergent selection is determines how much diversification can occur. Adopting this view will enhance tests of the relative role of natural and sexual selection in processes such as speciation

Data from: Plastic responses to parents and predators lead to divergent social behaviour in sticklebacks

Population divergence in antipredator defense and behaviour occurs rapidly and repeatedly. Genetic differences, phenotypic plasticity, or parental effects may all contribute to divergence, but the relative importance of each of these mechanisms remains unknown. We exposed juveniles to parents and predators to measure how induced changes contribute to shoaling behaviour differences between two threespine stickleback species (benthics and limnetics: Gasterosteus spp). We found that limnetics increased shoaling in response to predator attacks while benthics did not alter their behaviour. Care by limnetic fathers led to increased shoaling in both limnetic and benthic offspring. Shoaling helps limnetics avoid trout and avian predation; our results suggest this adaptive behaviour is the result of a combination of paternal effects, predator-induced plasticity, and genetic differences between species. These results suggest that plasticity substantially contributes to the rapid divergence in shoaling behaviour across the post-Pleistocene radiation of sticklebacks

Data from: Predator experience overrides learned aversion to heterospecifics in stickleback species pairs

Predation risk can alter female mating decisions because the costs of mate searching and selecting attractive mates increase when predators are present. In response to predators, females have been found to plastically adjust mate preference within species, but little is known about how predators alter sexual isolation and hybridization among species. We tested the effects of predator exposure on sexual isolation between benthic and limnetic threespine sticklebacks (Gasterosteus spp.). Female discrimination against heterospecific mates was measured before and after females experienced a simulated attack by a trout predator or a control exposure to a harmless object. In the absence of predators, females showed increased aversion to heterospecifics over time. We found that predator exposure made females less discriminating and precluded this learned aversion to heterospecifics. Benthic and limnetic males differ in coloration, and predator exposure also affected sexual isolation by weakening female preferences for colourful males. Predator effects on sexual selection were also tested but predators had few effects on female choosiness among conspecific mates. Our results suggest that predation risk may disrupt the cognitive processes associated with mate choice and lead to fluctuations in the strength of sexual isolation between species

Data from: Male competition fitness landscapes predict both forward and reverse speciation

Speciation is facilitated when selection generates a rugged fitness landscape such that populations occupy different peaks separated by valleys. Competition for food resources is a strong ecological force that can generate such divergent selection. However, it is unclear whether intrasexual competition over resources that provide mating opportunities can generate rugged fitness landscapes that foster speciation. Here we use highly variable male F2 hybrids of benthic and limnetic threespine sticklebacks, Gasterosteus aculeatus Linnaeus, 1758, to quantify the male competition fitness landscape. We find that disruptive sexual selection generates two fitness peaks corresponding closely to the male phenotypes of the two parental species, favouring divergence. Most surprisingly, an additional region of high fitness favours novel hybrid phenotypes that correspond to those observed in a recent case of reverse speciation after anthropogenic disturbance. Our results reveal that sexual selection through male competition plays an integral role in both forward and reverse speciation