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The evolution of behavioral isolation is often the first step toward speciation. While past studies show that behavioral isolation will sometimes evolve as a by-product of divergent ecological selection, we lack a more nuanced understanding of factors that may promote or hamper its evolution. The environment in which mating occurs may be important in mediating whether behavioral isolation evolves for two reasons. Ecological speciation could occur as a direct outcome of different sexual interactions being favored in different mating environments. Alternatively, mating environments may vary in the constraint they impose on traits underlying mating interactions, such that populations evolving in a "constraining" mating environment would be less likely to evolve behavioral isolation than populations evolving in a less constraining mating environment. In the latter, mating environment is not the direct cause of behavioral isolation but rather permits its evolution only if other drivers are present. We test these ideas with a set of 28 experimental fly populations, each of which evolved under one of two mating environments and one of two larval environments. Counter to the prediction of ecological speciation by mating environment, behavioral isolation was not maximal between populations evolved in different mating environments. Nonetheless, mating environment was an important factor as behavioral isolation evolved among populations from one mating environment but not among populations from the other. Though one mating environment was conducive to the evolution of behavioral isolation, it was not sufficient: assortative mating only evolved between populations adapting to different-larval environments within that mating environment, indicating a role for ecological speciation. Intriguingly, the mating environment that promoted behavioral isolation is characterized by less sexual conflict compared to the other mating environment. Our results suggest that mating environments play a key role in mediating ecological speciation via other axes of divergent selection. [ABSTRACT FROM AUTHOR]

Many genes are differentially expressed between males and females and patterns of sex-biased gene expression (SBGE) vary among species. Some of this variation is thought to have evolved in response to differences in mate competition among species that cause varying patterns of sex-specific selection. We used experimental evolution to test this by quantifying SBGE and sex-specific splicing in 15 Drosophila melanogaster populations that evolved for 104 generations in mating treatments that removed mate competition via enforced monogamy, or allowed mate competition in either small, simple, or larger, structurally more complex mating environments. Consistent with sex-specific selection affecting SBGE, initially sex-biased genes diverged in expression more among treatments than unbiased genes, and there was greater expression divergence for male- than female-biased genes. It has been suggested the transcriptome should be "feminized" under monogamy because of the removal of sexual selection on males; we did not observe this, likely because selection differs in additional ways between monogamy vs. polygamy. Significant divergence in average expression dimorphism between treatments was observed and, in some treatment comparisons, the direction of the divergence differed across different sex-bias categories. There was not a generalized reduction in expression dimorphism under enforced monogamy. [ABSTRACT FROM AUTHOR]

Prezygotic isolation is often stronger between sympatric as opposed to allopatric taxa, but the underlying cause can be difficult to infer from comparative studies alone. Experimental evolution, where evolutionary responses to treatments manipulating the presence/absence of heterospecific individuals are tracked, can provide a powerful complementary approach. We used experimental evolution to investigate a naturally occurring pattern of reproductive character displacement in the mushroom-feeding fly, Drosophila subquinaria. In nature, female D. subquinaria from populations sympatric with the closely related Drosophila recens discriminate more strongly against heterospecific males than do females from allopatric populations. Starting with 16 replicate allopatric populations of D. subquinaria , we manipulated the presence/absence of D. recens during mating (experimental sympatry vs. control) and, when present, we allowed hybrids to live or kill them each generation. Across 12 generations, heterospecific offspring production from no-choice mating trials between D. subquinaria females and D. recens males declined in both experimental sympatry treatments relative to the control, suggesting increased sexual isolation. Male cuticular hydrocarbon profiles also evolved, but only in the hybrids killed treatment. Our results strongly imply that the existing reproductive character displacement in wild D. subquinaria populations was an evolutionary response to selection arising from secondary contact with D. recens. [ABSTRACT FROM AUTHOR]

Male harm arises when traits that increase reproductive success in competition with other males also harm females as a side effect. The extent of harm depends on male and female phenotypes, both of which can diverge between populations. Within a population, harm is inferred when increased exposure to males reduces female fitness, but studies of the divergence of male harm rarely manipulate male exposure. Here, we quantify male harm and compare its magnitude between two lab populations of Drosophila serrata that were derived from a common ancestor 7 years earlier and subsequently held under conditions that minimized environmental differences. We manipulated female exposure to males in a factorial design involving all four combinations of males and females from these populations, providing insight into divergence in both sexes. Our results reveal substantial harm to females and provide stronger evidence of divergence in males than in females. Using these and other published data, we discuss conceptual issues surrounding the quantification and comparison of harm that arise because it involves a comparison of multiple quantities (e.g., female fitness under varying male exposure), and we demonstrate the increased insight that is gained by manipulating male exposure to quantify these quantities. [ABSTRACT FROM AUTHOR]

The epicuticular compounds (ECs) of insects serve both to waterproof the cuticle and, in many taxa, as pheromones that are important for various social interactions, including mate choice within populations. However, ECs have not been individually identified in many species and most studies of their role in mate choice have been performed in a laboratory setting. Here we newly identify and quantify the ECs of the antler fly, Protopiophila litigata Bonduriansky, and use a cross-sectional selection analysis to quantify their association with male mating success in the wild across two years (2013 and 2017). The ECs of antler flies include straight-chain and methylated alkanes, alkenes, and a family of branched wax esters. We find all ECs to be shared between males and females but also demonstrate sexual dimorphism in the abundance of several. Male EC relative abundances were significantly associated with mating success in both years, although the multivariate direction of selection differed significantly between the years. Surprisingly, only two of the 18 compounds (or groups of compounds) we identified were similarly associated with mating success across the sampling years. In 2017, we further partitioned sexual selection into intra- and intersexual components, revealing selection on ECs to be significant via female choice but not male–male competition. Our study is one of few to investigate the potential role of ECs in mating success in the wild and adds to a growing body of evidence demonstrating significant temporal variability in selection in natural populations. [ABSTRACT FROM AUTHOR]

A longstanding focus in evolutionary physiology concerns the causes and consequences of variation in maintenance metabolism. Insight into this can be gained by estimating the sex‐specific genetic architecture of maintenance metabolism alongside other, potentially correlated traits on which selection may also act, such as body mass and locomotor activity. This may reveal potential genetic constraints affecting the evolution of maintenance metabolism. Here, we used a half‐sibling breeding design to quantify the sex‐specific patterns of genetic (co)variance in standard metabolic rate (SMR), body mass and daily locomotor activity in Drosophila melanogaster. There was detectable additive genetic variance for all traits in both sexes. As expected, SMR and body mass were strongly and positively correlated, with genetic allometry exponents (bA ± SE) that were close to 2/3 in females (0.66 ± 0.16) and males (0.58 ± 0.32). There was a significant and positive genetic correlation between SMR and locomotor activity in males, suggesting that alleles that increase locomotion have pleiotropic effects on SMR. Sexual differences in the genetic architecture were largely driven by a difference in genetic variance in locomotor activity between the sexes. Overall, genetic variation was mostly shared between males and females, setting the stage for a potential intralocus sexual conflict in the face of sexually antagonistic selection. [ABSTRACT FROM AUTHOR]

Territoriality (i.e. defense of a resource) is the outcome of behavioral interactions that can result in selective advantages in many vertebrates and invertebrates. Since territoriality is expressed in a social context, an individuals' territoriality may change according to the phenotype of the opponents that they are confronted with (termed "indirect effects"). Defending a territory may also confer energetic costs to individuals, which could be reflected in their standard metabolic rate (SMR), a key component of an ectotherms' energy budget. Here, we measured territoriality using dyadic contests, body mass, and SMR using flow-through respirometry, twice in each of 192 adult male Drosophila melanogaster. Territoriality, body mass, and (whole-animal) SMR were all significantly repeatable. However, essentially all the among-individual variation in SMR was shared with body mass, as indicated by a very strong among-individual correlation (r ind) between body mass and SMR. The among-individual correlation between territoriality and SMR also tended to be positive, suggesting the presence of underlying metabolic costs to territoriality. Although indirect effects on territoriality were present but weak (accounting for 8.4% of phenotypic variance), indirect effects on territoriality were negatively and significantly correlated with body mass. This indicates that larger individuals tended to suppress their opponents territoriality. Variation among individuals in their ability to suppress territoriality in others was not associated with their own territoriality or SMR. [ABSTRACT FROM AUTHOR]

Standard metabolic rate (SMR), defined as the minimal energy expenditure required for self‐maintenance, is a key physiological trait. Few studies have estimated its relationship with fitness, most notably in insects. This is presumably due to the difficulty of measuring SMR in a large number of very small individuals. Using high‐throughput flow‐through respirometry and a Drosophila melanogaster laboratory population adapted to a life cycle that facilitates fitness measures, we quantified SMR, body mass, and fitness in 515 female and 522 male adults. We used a novel multivariate approach to estimate linear and nonlinear selection differentials and gradients from the variance‐covariance matrix of fitness, SMR, and body mass, allowing traits specific covariates to be accommodated within a single model. In males, linear selection differentials for mass and SMR were positive and individually significant. Selection gradients were also positive but, despite substantial sample sizes, were nonsignificant due to increased uncertainty given strong SMR‐mass collinearity. In females, only nonlinear selection was detected and it appeared to act primarily on body size, although the individual gradients were again nonsignificant. Selection did not differ significantly between sexes although differences in the fitness surfaces suggest sex‐specific selection as an important topic for further study. [ABSTRACT FROM AUTHOR]

Mating/fertilization success and fecundity are influenced by sexual interactions among individuals, the nature and frequency of which can vary among different environments. The extent of local adaptation for such adult fitness components is poorly understood. We allowed 63 populations of Drosophila melanogaster to independently evolve in one of three mating environments that alter sexual interactions: one involved enforced monogamy, while the other two permitted polygamy in either structurally simple standard fly vials or in larger "cages" with added complexity. Adult male and female reproductive fitness were measured after 16 and 28 generations, respectively, via full reciprocal transplants. In males, reciprocal local adaptation was observed between the monogamy and simple polygamy treatments, consistent with the evolution of reproductively competitive males under polygamy that perform poorly under monogamy because they harm their only mate. However, males evolved in the complex polygamy treatment performed similarly or better than all other males in all mating environments, consistent with previous results showing higher genetic quality in this treatment. Differences in female fitness were more muted, suggesting selection on females was less divergent across the mating treatments and echoing a common pattern of greater phenotypic and expression divergence in males than females. [ABSTRACT FROM AUTHOR]

Abundant evidence supports a role for sexual selection in the evolution of reproductive isolation, and it is thus unsurprising that much attention has been given, both conceptually and empirically, to understanding its role in speciation. In doing so, debate has arisen on how sexual selection fits within the much used ecological versus mutation‐order classification of speciation mechanisms, with sexual selection often presented as a distinct third alternative. We argue that models of speciation by sexual selection include a fundamental role of divergent selection between environments or mutation order in initiating the process. Rather than representing a unique mechanism, sexual selection layers a coevolutionary process between males and females on top of the classic mechanisms such that the evolution of each sex can now be driven by divergent selection, mutation order, and selection arising from interactions with the other sex. In addition to blurring the distinction between ecological and mutation‐order speciation, this coevolutionary process is likely to speed divergence. Sexual selection is not unique in this way; coevolutionary processes can similarly arise from ecological interactions between populations or species, with similar results. Ultimately, understanding the contribution of sexual selection to speciation will require identifying the processes that drive the divergence of mating biases. [ABSTRACT FROM AUTHOR]

Sexual interactions among adults can generate selection on both males and females with genome-wide consequences. Sexual selection through males is one component of this selection that has been argued to play an important role in purging deleterious alleles. A common technique to assess the influence of sexual selection is by a comparison of experimental evolution under enforced monogamy versus polygamy. Mixed results from past studies may be due to the use of highly simplified laboratory conditions that alter the nature of sexual interactions. Here, we examine the rate of purging of 22 gene disruption mutations in experimental polygamous populations of Drosophila melanogaster in each of two mating environments: a simple, high-density environment (i.e., typical fly vials), and a lower density, more spatially complex environment. Based on past work, we expect sexual interactions in the latter environment to result in stronger selection in both sexes. Consistent with this, we find that mutations tend to be purged more quickly in populations evolving in complex environments. We discuss possible mechanisms by which environmental complexity might modulate the rate at which deleterious alleles are purged and putatively ascribe a role for sexual interactions in explaining the treatment differences in our experiment. [ABSTRACT FROM AUTHOR]

The reinforcement of premating barriers due to reduced hybrid fitness in sympatry may cause secondary sexual isolation within a species as a by-product. Consistent with this, in the fly Drosophila subquinaria, females that are sympatric with D. recens mate at very low rates not only with D. recens, but also with conspecific D. subquinaria males from allopatry. Here, we ask if these effects of reinforcement cascade more broadly to affect sexual isolation with other closely related species. We assay reproductive isolation of these species with D. transversa and find that choosy D. subquinaria females from the region sympatric with D. recens discriminate strongly against male D. transversa, whereas D. subquinaria from the allopatric region do not. This increased sexual isolation cannot be explained by natural selection to avoid mating with this species, as they are allopatric in geographic range and we do not identify any intrinsic postzygotic isolation between D. subquinaria and D. transversa. Variation in epicuticular hydrocarbons, which are used as mating signals in D. subquinaria, follow patterns of premating isolation: D. transversa and allopatric D. subquinaria are most similar to each other and differ from sympatric D. subquinaria, and those of D. recens are distinct from the other two species. We suggest that the secondary effects of reinforcement may cascade to strengthen reproductive isolation with other species that were not a target of selection. These effects may enhance the divergence that occurs in allopatry to help explain why some species are already sexually isolated upon secondary contact. [ABSTRACT FROM AUTHOR]

Several lines of evidence implicate sexual isolation in both initiating and completing the speciation process. Although its existence is straightforward to demonstrate, understanding the evolution of sexual isolation requires identifying the underlying phenotypes responsible so that we can determine how these have diverged. Here, we study geographic variation in female mate preferences for male sexual displays in the fly Drosophila subquinaria. Female D. subquinaria that are sympatric with its sister species D. recens discriminate strongly against both D. recens and allopatric conspecific males, whereas females from allopatric populations do not. Furthermore, female mate preferences target at least in part a suite of cuticular hydrocarbons (CHCs) in males and geographic variation in CHCs mirrors the pattern of mate discrimination. In this study, we quantify female mate preferences for male CHCs from populations that span the geographic range of D. subquinaria. We find that the direction of linear sexual selection varies significantly between populations that are sympatric versus allopatric with D. recens in a pattern of reproductive character displacement. Differences in preference partially align with existing differences in CHCs and patterns of sexual isolation, although discrepancies remain that suggest the involvement of additional traits and/or more complex, nonlinear preference functions. [ABSTRACT FROM AUTHOR]

Sexually dimorphic traits have a history of directional selection for exaggeration in at least one sex. Theory suggests that traits targeted by persistent selection should evolve heightened condition dependence whereby their expression reflects the availability and processing efficiency of metabolic resources. This joint dependence of sexual dimorphism and condition dependence on directional selection should result in a positive correlation between the extent of sexual dimorphism and the strength of condition dependence. However, because direct phenotypic evidence is predominately from species with highly exaggerated male traits, it remains unclear whether condition-dependent sexual dimorphism is characteristic of species with more typical levels of dimorphism. We manipulated condition via larval diet and quantified sex-specific responses in adult body size and shape in a moderately dimorphic dipteran species, the antler fly Protopiophila litigata. While dimorphism did not increase with diet quality within any trait, among traits the extent of dimorphism was positively associated with the strength of condition dependence in males but not females, as previously reported for highly dimorphic species. This finding suggests that a shared developmental basis to condition dependence and sexual dimorphism can arise via sexual selection on males even in lineages lacking highly exaggerated male traits. [ABSTRACT FROM AUTHOR]

Sexual selection arises from both intrasexual competition and mate choice. With respect to the evolution of male traits, there is a vast literature documenting the existence of female choice and male-male competition, and both have been shown to co-occur in many species. Despite numerous studies of these two components of male reproductive success in isolation, few have investigated whether and how they interact to determine total sexual selection. To address this, we investigate male territoriality in Drosophila serrata, a species in which female preference for male sexual pheromones (cuticular hydrocarbons or CHCs) have been extensively studied. We demonstrate that territoriality occurs, that it involves direct male-male aggressive interactions, and that it contributes to variation in male mating success. Results from a phenotypic manipulation also indicate that territorial success is condition-dependent, although a genetic manipulation of condition, involving three generations of full-sib inbreeding, failed to find a significant effect. Finally, selection assays also suggest that territorial success depends on male body size but not on CHCs, whereas the opposite is true for mating success. [ABSTRACT FROM AUTHOR]

Interactions between species can alter selection on sexual displays used in mate choice within species. Here we study the epicuticular pheromones of two Drosophila species that overlap partially in geographic range and are incompletely reproductively isolated. Drosophila subquinaria shows a pattern of reproductive character displacement against Drosophila recens, and partial behavioral isolation between conspecific sympatric versus allopatric populations, whereas D. recens shows no such variation in mate choice. First, using manipulative perfuming experiments, we show that females use pheromones as signals for mate discrimination both between species and among populations of D. subquinaria. Second, we show that patterns of variation in epicuticular compounds, both across populations and between species, are consistent with those previously shown for mating probabilities: pheromone compositions differ between populations of D. subquinaria that are allopatric versus sympatric with D. recens, but are similar across populations of D. recens regardless of overlap with D. subquinaria. We also identify differences in pheromone composition among allopatric regions of D. subquinaria. In sum, our results suggest that epicuticular compounds are key signals used by females during mate recognition, and that these traits have diverged among D. subquinaria populations in response to reinforcing selection generated by the presence of D. recens. [ABSTRACT FROM AUTHOR]

Directional selection is prevalent in nature, yet phenotypes tend to remain relatively constant, suggesting a limit to trait evolution. However, the genetic basis of this limit is unresolved. Given widespread pleiotropy, opposing selection on a trait may arise from the effects of the underlying alleles on other traits under selection, generating net stabilizing selection on trait genetic variance. These pleiotropic costs of trait exaggeration may arise through any number of other traits, making them hard to detect in phenotypic analyses. Stabilizing selection can be inferred, however, if genetic variance is greater among low- compared to high-fitness individuals. We extend a recently suggested approach to provide a direct test of a difference in genetic variance for a suite of cuticular hydrocarbons (CHCs) in Drosophila serrata. Despite strong directional sexual selection on these traits, genetic variance differed between high- and low-fitness individuals and was greater among the low-fitness males for seven of eight CHCs, significantly more than expected by chance. Univariate tests of a difference in genetic variance were nonsignificant but likely have low power. Our results suggest that further CHC exaggeration in D. serrata in response to sexual selection is limited by pleiotropic costs mediated through other traits. [ABSTRACT FROM AUTHOR]

The effects of sexual selection on population mean fitness are unclear and a subject of debate. Recent models propose that, because reproductive success may be condition dependent, much of the genome may be a target of sexual selection. Under this scenario, mutations that reduce health, and thus nonsexual fitness, may also be deleterious with respect to reproductive success, meaning that sexual selection may contribute to the purging of deleterious alleles. We tested this hypothesis directly by subjecting replicate Drosophila melanogaster populations to two treatments that altered the opportunity for sexual selection and then tracked changes in the frequency of six separate deleterious alleles with recessive and visible phenotypic effects. While natural selection acted to decrease the frequency of all six mutations, the addition of sexual selection did not aid in the purging of any of them, and for three of them appears to have hampered it. Courtship and mating have harmful effects in this species and mate choice assays showed that males directed more courtship and mating behavior toward wild-type over mutant females, providing a likely explanation for sexual selection's cost. Whether this cost extends to other mutations (e.g., those lacking visible phenotypic effects) is an important topic for future research. [ABSTRACT FROM AUTHOR]

Mutual mate choice for homologous sexual display traits has been demonstrated in several recent studies yet little attention has been given to quantitative comparison of the strength and form of mate preferences between the sexes. Such comparisons may provide important insight into the evolution of mate choice for honest signals. In particular, because females generally provide the majority of resources for initial offspring development, female displays may trade-off with fecundity, causing preference evolution to differ between the sexes. Recent theory suggests that adaptive male preferences for honest displays in females are possible under certain conditions and may result in preferences that are convex (i.e., stabilizing) in form. We compared sexual selection on a suite of contact pheromones arising from mutual mate choice using nine separate geographic populations of Drosophila serrata. We show that the convex selection is stronger on females than on males overall in these populations, and that convex selection is the predominate form of nonlinear selection on females but not males. [ABSTRACT FROM AUTHOR]

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 laboratory-adapted 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. [ABSTRACT FROM AUTHOR]

Indirect genetics effects (IGEs)—when the genotype of one individual affects the phenotypic expression of a trait in another—may alter evolutionary trajectories beyond that predicted by standard quantitative genetic theory as a consequence of genotypic evolution of the social environment. For IGEs to occur, the trait of interest must respond to one or more indicator traits in interacting conspecifics. In quantitative genetic models of IGEs, these responses (reaction norms) are termed interaction effect coefficients and are represented by the parameter psi (Ψ). The extent to which Ψ exhibits genetic variation within a population, and may therefore itself evolve, is unknown. Using an experimental evolution approach, we provide evidence for a genetic basis to the phenotypic response caused by IGEs on sexual display traits in Drosophila serrata. We show that evolution of the response is affected by sexual but not natural selection when flies adapt to a novel environment. Our results indicate a further mechanism by which IGEs can alter evolutionary trajectories—the evolution of interaction effects themselves. [ABSTRACT FROM AUTHOR]

Many laboratory speciation experiments have raised allopatric populations in different environments to determine whether reproductive isolation evolves as a byproduct of adaptation (a form of ecological speciation). Few, however, have addressed the evolution of both pre- and postmating isolation or investigated the conditions affecting the process. We present results of an evolution experiment in which 12 lines of Drosophila melanogaster were derived from a common population and then independently evolved for more than 57 generations under alternative selection treatments. Six “desiccation” populations had access to food and water removed during a period of their adult lives generating strong desiccation selection, and six “starvation” populations had access to food but not water removed for the same period, generating a mild starvation stress. Substantial divergence of cuticular hydrocarbons occurred between the desiccation and starvation populations, key traits that have been implicated in sexual isolation in Drosophila. Despite this divergence, there was no detectable premating isolation between desiccation and starvation populations and postmating isolation was asymmetrical: the fitness of F1 hybrids was reduced in the desiccation but not the starvation environment. This asymmetry was likely caused by the absence of divergent selection: adaptation to desiccation appears to have come at no cost to performance in the starvation environment. Novel environments are thought to promote the evolution of reproductive isolation. Understanding the conditions that favor or hamper this remains a key challenge for speciation research. [ABSTRACT FROM AUTHOR]

The net effect of sexual selection on nonsexual fitness is controversial. On one side, elaborate display traits and preferences for them can be costly, reducing the nonsexual fitness of individuals possessing them, as well as their offspring. In contrast, sexual selection may reinforce nonsexual fitness if an individual's attractiveness and quality are genetically correlated. According to recent models, such good-genes mate choice should increase both the extent and rate of adaptation. We evolved 12 replicate populations of Drosophila serrata in a powerful two-way factorial experimental design to test the separate and combined contributions of natural and sexual selection to adaptation to a novel larval food resource. Populations evolving in the presence of natural selection bad significantly higher mean nonsexual fitness when measured over three generations (13-15) during the course of experimental evolution (16-23% increase). The effect of natural selection was even more substantial when measured in a standardized, monogamous mating environment at the end of the experiment (generation 16; 52% increase). In contrast, and despite strong sexual selection on display traits, there was no evidence from any of the four replicate fitness measures that sexual selection promoted adaptation. In addition, a comparison of fitness measures conducted under different mating environments demonstrated a significant direct cost of sexual selection to females, likely arising from some form of male-induced harm. Indirect benefits of sexual selection in promoting adaptation to this novel resource environment therefore appear to be absent in this species, despite prior evidence suggesting the operation of good-genes mate choice in their ancestral environment. How novel environments affect the operation of good-genes mate choice is a fundamental question for future sexual selection research. [ABSTRACT FROM AUTHOR]

Mechanisms of speciation are not well understood, despite decades of study. Recent work has focused on how natural and sexual selection cause sexual isolation. Here, we investigate the roles of divergent natural and sexual selection in the evolution of sexual isolation between sympatric species of threespine sticklebacks. We test the importance of morphological and behavioral traits in conferring sexual isolation and examine to what extent these traits have diverged in parallel between multiple, independently evolved species pairs. We use the patterns of evolution in ecological and mating traits to infer the likely nature of selection on sexual isolation. Strong parallel evolution implicates ecologically based divergent natural and/or sexual selection, whereas arbitrary directionality implicates nonecological sexual selection or drift. In multiple pairs we find that sexual isolation arises in the same way: assortative mating on body size and asymmetric isolation due to male nuptial color. Body size and color have diverged in a strongly parallel manner, similar to ecological traits. The data implicate ecologically based divergent natural and sexual selection as engines of speciation in this group. [ABSTRACT FROM AUTHOR]

While the feasibility of bottleneck-induced speciation is in doubt, population bottlenecks may still affect the speciation process by interacting with divergent selection. To explore this possibility, I conducted a laboratory speciation experiment using Drosophila pseudoobscura involving 78 replicate populations assigned in a two-way factorial design to both bottleneck (present vs. absent) and environment (ancestral vs. novel) treatments. Populations independently evolved under these treatments and were then tested for assortative mating and male mating success against their common ancestor. Bottlenecks alone did not generate any premating isolation, despite an experimental design that was conducive to bottleneck-induced speciation. Premating isolation also did not evolve in the novel environment treatment, neither in the presence nor absence of bottlenecks. However, male mating success was significantly reduced in the novel environment treatment, both as a plastic response to this environment and as a result of environment-dependent inbreeding effects in the bottlenecked populations. Reduced mating success of derived males will hamper speciation by enhancing the mating success of immigrant, ancestral males. Novel environments are generally thought to promote ecological speciation by generating divergent natural selection. In the current experiment, however, the novel environment did not cause the evolution of any premating isolation and it reduced the likelihood of speciation through its effects on male mating success. [ABSTRACT FROM AUTHOR]

Presents a study that tested a prediction of the ecological model concerning the fitness of hybrids between two young, sympatric species of threespine sticklebacks. Description of the design of the transplant experiment; Growth rate of each species; Average body mass and other morphological traits of the samples.

Determines whether there is a sufficient experimental design for the detection of ecologically dependent isolation. Reasons for hybrid unfitness; Genetic interpretation model developed by Lynch in 1991; Two environments and environment-dependent gene effects.

Provides some evidence for the founder-flush model of speciation by demonstrating the evolution of significant premating isolation. Details on the founder-flush model of speciation; Experimental methodology; Results and discussion.

Examines the hypothesis of reinforcement, by investigating how the strength of premating isolation varies between sympatric and allopatric populations of threespine sticklebacks. Methodology used in study; Information on the results which proved to be consistent with the hypothesis; Alternatives to reinforcement; Discussion on the findings of the investigation.