The intersex genetic correlation for fitness r fm w À Á , a standardized measure of the degree to which male and female fitness covary genetically, has consequences for important evolutionary processes, but few estimates are available and none have explored how it changes with environment. Using a half-sibling breeding design, we estimated the genetic (co)variance matrix (G) for male and female fitness, and the resulting r fm w , in Drosophila serrata. Our estimates were performed in two environments: the laboratory yeast food to which the population was well adapted and a novel corn food. The major axis of genetic variation for fitness in the two environments, accounting for 51.3 per cent of the total genetic variation, was significant and revealed a strong signal of sexual antagonism, loading negatively in both environments on males but positively on females. Consequently, estimates of r fm w were negative in both environments (K0.34 and K0.73, respectively), indicating that the majority of genetic variance segregating in this population has contrasting effects on male and female fitness. The possible strengthening of the negative r fm w in this novel environment may be a consequence of no history of selection for amelioration of sexual conflict. Additional studies from a diverse range of novel environments will be needed to determine the generality of this finding.

Sexually antagonistic genetic variation can pose limits to the independent evolution and adaptation of the sexes. The extent of sexually antagonistic variation is reflected in the intersex genetic correlation for fitness (r w FM ). Previous estimates of this correlation have been mostly limited to populations in environments to which they are already well adapted, making it difficult to gauge the importance of sexually antagonistic genetic variance during the early stages of adaptation, such as that occurring following abrupt environmental change or upon the colonization of new habitat. Here we assayed male and female lifetime fitness in a population of Drosophila serrata in four novel laboratory environments. We found that r w FM varied significantly across environments, with point estimates ranging from positive to negative values of considerable magnitude. We also found that the variability among estimates was because, at least in part, of significant differences among environments in the genetic variances of both male and female fitness, with no evidence of any significant changes in the intersex covariance itself, although standard errors of these estimates were large. Our results illustrate the unpredictable nature of r w FM in novel environments and suggest that, although sexually antagonistic genetic variance can be pronounced in some novel environments, it may have little effect in constraining the early stages of adaptation in others.

Theory predicts that costly sexual displays should evolve condition dependence if the marginal fitness gain from trait exaggeration is greater for high- than for low-condition individuals and that the strength of condition dependence should increase with the strength of directional selection. While there is substantial support for the first prediction, evidence for the latter is much weaker. We undertook a quantitative test of this prediction for a multivariate sexual display consisting of a suite of contact pheromones termed "cuticular hydrocarbons" (CHCs) in Drosophila serrata. We performed a dietary manipulation of condition (i.e., the pool of metabolic resources available for allocation to fitness-enhancing traits) within a half-sibling breeding design, thereby also providing insight into the genetic basis of condition dependence. As predicted, the linear combination of CHCs under the strongest sexual selection from female mate preferences was unusually condition dependent relative to other CHC combinations within the population ([Formula: see text]). A significant positive correlation also existed between the strengths of condition dependence and sexual selection among different CHC blends ([Formula: see text], [Formula: see text]). Finally, sires varied in their response to the dietary manipulation, demonstrating significant genetic variance in condition dependence. Our results are consistent with the evolution of heightened condition dependence of sexual displays in response to persistent sexual selection.

A female's mate preference is a potentially complex function relating variation in multiple male phenotypes with her probability of accepting individual males as a mate. Estimating the quantitative genetic basis preference functions within a population is empirically challenging yet key to understanding preference evolution. We employed a recently described approach that uses random-coefficient mixed models in the analysis of function-valued traits. Using a half-sibling breeding design in a laboratoryadapted Drosophila serrata population, we estimated the genetic (co)variance function of female preference for male sexual displays composed of nine contact pheromones. The breeding design was performed across two environments: the food to which the population was well adapted and a novel food that reduced average female productivity by 35%. Significant genetic variance in female preference was detected and the majority (64.2%) was attributable to a single genetic dimension (eigenfunction), suggesting that preferences for different pheromones are not genetically independent. The second eigenfunction, accounting for 24% of the total genetic variance, approached significance in a conservative test, suggesting the existence of a second, independent genetic dimension. There was no evidence that the genetic basis of female preference differed between the two environments, suggesting the absence of genotype-by-environment interactions and hence a lack of condition-dependent preference expression.

The extent of variation in mating signals between environments can be an important determinant of reproductive isolation. We tested divergence in male courtship behavior between lake and stream three-spined stickleback (Gasterosteus aculeatus L.) in the Misty system, Vancouver Island. Presumably, genetically based divergence was assessed by comparing male behavior (aggressive, display, and nest activities) among Lake, Inlet, and Outlet ecotypes that were reared from fertilization to maturity in a common environment. Plastic components of behavior were assessed by comparing male behavior in presence of females from each of the 3 ecotypes. We found that Inlet males had less aggressive elements than Lake and Outlet males in their behavior, whereas Lake and Outlet males were similar in their behavior. We further found that males of all ecotypes performed less display and nest activities in the presence of Lake and Outlet than Inlet females and that the male mating behavior was influenced by the relative difference in male and female body sizes. We suggest that divergent selection has driven genetic divergence in aggressive behaviors, whereas other behavioral components respond more plastically to variation in female phenotype. We discuss the implications of these results for reproductive isolation.

Phenotypes tend to remain relatively constant in natural populations, suggesting a limit to trait evolution. Although stationary phenotypes suggest stabilizing selection, directional selection is more commonly reported. However, selection on phenotypes will have no evolutionary consequence if the traits do not genetically covary with fitness, a covariance known as the Robertson-Price Identity. The nature of this genetic covariance determines if phenotypes will evolve directionally or whether they reside at an evolutionary optimum. Here, we show how a set of traits can be shown to be under net stabilizing selection through an application of the multivariate Robertson-Price Identity. We characterize how a suite of male sexual displays genetically covaries with fitness in a population of Drosophila serrata. Despite strong directional sexual selection on these phenotypes directly and significant genetic variance in them, little genetic covariance was detected with overall fitness. Instead, genetic analysis of trait deviations showed substantial stabilizing selection on the genetic variance of these traits with respect to overall fitness, indicating that they reside at an evolutionary optimum. In the presence of widespread pleiotropy, stabilizing selection on focal traits will arise through the net effects of selection on other, often unmeasured, traits and will tend to be stronger on trait combinations than single traits. Such selection may be difficult to detect in phenotypic analyses if the environmental covariance between the traits and fitness obscures the underlying genetic associations. The genetic analysis of trait deviations provides a way of detecting the missing stabilizing selection inferred by recent metaanalyses. cuticular hydrocarbons | evolutionary stasis

In ecological speciation, reproductive isolation evolves as a consequence of adaptation to different selective environments. A frequent contributor to this process is the evolution of positive assortative mate choice between ecotypes. We tested this expectation for lake and inlet stream threespine stickleback (Gasterosteus aculeatus) from the Misty system (Vancouver Island, Canada), which show strong genetically based adaptive divergence and little genetic exchange in nature. This, and work on other stickleback systems, led us to expect positive assortative mating. Yet, our standard "no-choice" laboratory experiment on common-garden fish revealed no evidence for this-despite divergence in traits typically mediating assortative mating in stickleback. These results remind us that divergent natural selection may not inevitably lead to the evolution of positive assortative mate choice. The apparent lack of strong and symmetric reproductive barriers in this system presents a conundrum: why are such barriers not evident despite strong adaptive divergence and low gene flow in nature?

Sexually dimorphic traits are likely to have evolved through sexually antagonistic selection. However, recent empirical data suggest that intralocus sexual conflict often persists, even when traits have diverged between males and females. This implies that evolved dimorphism is often incomplete in resolving intralocus conflict, providing a mechanism for the maintenance of genetic variance in fitness-related traits. We used experimental evolution in Drosophila melanogaster to directly test for ongoing conflict over a suite of sexually dimorphic cuticular hydrocarbons (CHCs) that are likely targets of sex-specific selection. Using a set of experimental populations in which the transmission of genetic material had been restricted to males for 82 generations, we show that CHCs did not evolve, providing experimental evidence for the absence of current intralocus sexual conflict over these traits. The absence of ongoing conflict could indicate that CHCs have never been the target of sexually antagonistic selection, although this would require the existing dimorphism to have evolved via completely sexlinked mutations or as a result of former, but now absent, pleiotropic effects of the underlying loci on another trait under sexually antagonistic selection. An alternative interpretation, and which we believe to be more likely, is that the extensive CHC sexual dimorphism is the result of past intralocus sexual conflict that has been fully resolved, implying that these traits have evolved genetic independence between the sexes and that genetic variation in them is therefore maintained by alternative mechanisms. This latter interpretation is consistent with the known roles of CHCs in sexual communication in this species and with previous studies suggesting the genetic independence of CHCs between males and females. Nevertheless, direct estimates of sexually antagonistic selection will be important to fully resolve these alternatives.