The introduction of a new species can change the characteristics of other species within a community. These changes may affect discontiguous trophic levels via adjacent trophic levels. The invasion of an exotic host species may provide the opportunity to observe the dynamics of changing interspecific interactions among parasitoids belonging to different trophic levels. The exotic large white butterfly Pieris brassicae invaded Hokkaido Island, Japan, and quickly spread throughout the island. Prior to the invasion, the small white butterfly P. rapae was the host of the primary parasitoid Cotesia glomerata, on which both the larval hyperparasitoid Baryscapus galactopus and the pupal hyperparasitoid Trichomalopsis apanteroctena depended. At the time of the invasion, C. glomerata generally laid eggs exclusively in P. rapae. During the five years following the invasion, however, the clutch size of C. glomerata in P. rapae gradually decreased, whereas the clutch size in P. brassicae increased. The field results corresponded well with laboratory experiments showing an increase in the rate of parasitism in P. brassicae. The host expansion of C. glomerata provided the two hyperparasitoids with an opportunity to choose between alternative hosts, that is, C. glomerata within P. brassicae and C. glomerata within P. rapae. Indeed, the pupal hyperparasitoid T. apanteroctena shifted its preference gradually to C. glomerata in P. brassicae, whereas the larval hyperparasitoid B. galactopus maintained a preference for C. glomerata in P. rapae. These changes in host preference may result from differential suitability of the two host types. The larval hyperparasitoid preferred C. glomerata within P. rapae to C. glomerata within P. brassicae, presumably because P. brassicae larvae attacked aggressively, thereby hindering the parasitization, whereas the pupal hyperparasitoid could take advantage of the competition-free resource by shifting its host preference. Consequently, the invasion of P. brassicae has changed the host use of the primary parasitoid C. glomerata and the pupal hyperparasitoid T. apanteroctena within a very short time.

Genotype × environment interactions can facilitate coexistence of locally adapted specialists. Interactions evolve if adaptation to one environment trades off with performance in others. We investigated whether evolution on one host genotype traded off with performance on others in long-term experimental populations of different genotypes of the protozoan Paramecium caudatum, infected with the bacterial parasite Holospora undulata. A total of nine parasite selection lines evolving on three host genotypes and the ancestral parasite were tested in a cross-infection experiment. We found that evolved parasites produced more infections than did the ancestral parasites, both on host genotypes they had evolved on (positive direct response to selection) and on genotypes they had not evolved on (positive correlated response to selection). On two host genotypes, a negative relationship between direct and correlated responses indicated pleiotropic costs of adaptation. On the third, a positive relationship suggested cost-free adaptation. Nonetheless, on all three hosts, resident parasites tended to be superior to the average nonresident parasite. Thus genotype specificity (i.e., patterns of local adaptation) may evolve without costs of adaptation, as long as direct responses to selection exceed correlated responses.

Because disease resistance is a hallmark signature of pathogen-mediated selection pressure on hosts, studies of resistance structure (the spatial distribution of disease resistance genes among conspecific host populations) can provide valuable insights into the influence of pathogens on host evolution and spatial variation in the magnitude of their effects. To date few studies of wild plant–pathogen interactions have characterized resistance structure by sampling across the host's biogeographic range, and only a handful have paired such investigations with studies of disease levels under natural conditions. I used a greenhouse cross-inoculation experiment to characterize genetic resistance of 16 populations of California dwarf flax (Hesperolinon californicum) to attack by multiple samples of the rust fungus Melampsora lini. I documented a latitudinal cline in resistance structure, manifest across the host's biogeographic range, which mirrored almost identically a cline in infection prevalence documented through field surveys of disease in study populations. These results provide empirical evidence for clinal patterns of antagonistic selection pressure, demonstrate that such patterns can be manifest across broad biogeographic scales, and suggest that rates of disease prevalence in wild plant populations may be tightly linked to the distribution of host resistance genes. Tests for local adaptation of the fungus revealed evidence of the phenomenon (significantly greater infection in sympatric plant–fungal pairings) as well as the potential for substantial bias to be introduced into statistical analyses by spatial patterns of host resistance structure.

Many well-studied coevolutionary interactions between predators and prey or hosts and parasites are mediated by quantitative traits. In some interactions, such as those between cuckoos and their hosts, interactions are mediated by the degree of phenotype matching among species, and a significant body of theory has been developed to predict the coevolutionary dynamics and outcomes of such interactions. In a large number of other cases, however, interactions are mediated by the extent to which the phenotype of one species exceeds that of the other. For these cases—which are arguably more numerous—few theoretical predictions exist for coevolutionary dynamics and outcomes. Here we develop and analyze mathematical models of interspecific interactions mediated by the extent to which the quantitative trait of one species exceeds that of the other. Our results identify important differences from previously studied models based on trait matching. First, our results show that cyclical dynamics are possible only if the strength of coevolutionary selection exceeds a threshold and stabilizing selection acts on the interacting traits. Second, our results demonstrate that significant levels of genetic polymorphism can be maintained only when cyclical dynamics occur. This result leads to the unexpected prediction that maintenance of genetic polymorphism is enhanced by strong selection. Finally, our results demonstrate that there is no a priori reason to expect the traits of interacting species should match in any literal sense, even in the absence of gene flow among populations.

Wright's adaptive topography describes gene frequency evolution as a maximization of mean fitness in a constant environment. I extended this to a fluctuating environment by unifying theories of stochastic demography and fluctuating selection, assuming small or moderate fluctuations in demographic rates with a stationary distribution, and weak selection among the types. The demography of a large population, composed of haploid genotypes at a single locus or normally distributed phenotypes, can then be approximated as a diffusion process and transformed to produce the dynamics of population size, N, and gene frequency, p, or mean phenotype, 𝑧̄. The expected evolution of p or 𝑧̄ is a product of genetic variability and the gradient of the long-run growth rate of the population, 𝑟̃, with respect to p or 𝑧̄. This shows that the expected evolution maximizes 𝑟̃, the mean Malthusian fitness in the average environment minus half the environmental variance in population growth rate. Thus, 𝑟̃ as a function of p or 𝑧̄ represents an adaptive topography that, despite environmental fluctuations, does not change with time. The haploid model is dominated by environmental stochasticity, so the expected maximization is not realized. Different constraints on quantitative genetic variability, and stabilizing selection in the average environment, allow evolution of the mean phenotype to undergo a stochastic maximization of 𝑟̃. Although the expected evolution maximizes the long-run growth rate of the population, for a genotype or phenotype the long-run growth rate is not a valid measure of fitness in a fluctuating environment. The haploid and quantitative character models both reveal that the expected relative fitness of a type is its Malthusian fitness in the average environment minus the environmental covariance between its growth rate and that of the population.

How adaptive evolution occurs with individually deleterious but jointly beneficial mutations has been one of the major problems in population genetics theory. Adaptation in this case is commonly described as a population's escape from a local peak to a higher peak on Sewall Wright's fitness landscape. Recent molecular genetic and computational studies have suggested that genetic robustness can facilitate such peak shifts. If phenotypic expressions of new mutations are suppressed under genetic robustness, mutations that are otherwise deleterious can accumulate in the population as neutral variants. When the robustness is perturbed by an environmental change or a major mutation, these variants become exposed to natural selection. It is argued that this process promotes adaptation because allelic combinations enriched under genetic robustness can then be positively selected. Here, I propose simple two- and three-locus models of adaptation with partial genetic robustness as suggested by recent studies. The waiting time until the fixation of an adaptive haplotype was observed in stochastic simulations and compared to the expectation without robustness. It is shown that peak shifts can be delayed or accelerated depending on the conditions of genetic robustness. The evolutionary significance of these processes is discussed.

Theoretical quantitative genetics provides a framework for reconstructing past selection and predicting future patterns of phenotypic differentiation. However, the usefulness of the equations of quantitative genetics for evolutionary inference relies on the evolutionary stability of the additive genetic variance–covariance matrix (G matrix). A fruitful new approach for exploring the evolutionary dynamics of G involves the use of individual-based computer simulations. Previous studies have focused on the evolution of the eigenstructure of G. An alternative approach employed in this paper uses the multivariate response-to-selection equation to evaluate the stability of G. In this approach, I measure similarity by the correlation between response-to-selection vectors due to random selection gradients. I analyze the dynamics of G under several conditions of correlational mutation and selection. As found in a previous study, the eigenstructure of G is stabilized by correlational mutation and selection. However, over broad conditions, instability of G did not result in a decreased consistency of the response to selection. I also analyze the stability of G when the correlation coefficients of correlational mutation and selection and the effective population size change through time. To my knowledge, no prior study has used computer simulations to investigate the stability of G when correlational mutation and selection fluctuate. Under these conditions, the eigenstructure of G is unstable under some simulation conditions. Different results are obtained if G matrix stability is assessed by eigenanalysis or by the response to random selection gradients. In this case, the response to selection is most consistent when certain aspects of the eigenstructure of G are least stable and vice versa.

We derive formulas that can be applied to estimate the effective population size N_e for organisms with two sexes reproducing once a year and having constant adult mean vital rates independent of age. Temporal fluctuations in population size are generated by demographic and environmental stochasticity. For populations with even sex ratio at birth, no deterministic population growth and identical mean vital rates for both sexes, the key parameter determining N_e is simply the mean value of the demographic variance for males and females considered separately. In this case Crow and Kimura's generalization of Wright's formula for N_e with two sexes, in terms of the effective population sizes for each sex, is applicable even for fluctuating populations with different stochasticity in vital rates for males and females. If the mean vital rates are different for the sexes then a simple linear combination of the demographic variances determines N_e, further extending Wright's formula. For long-lived species an expression is derived for N_e involving the generation times for both sexes. In the general case with nonzero population growth and uneven sex ratio of newborns, we use the model to investigate numerically the effects of different population parameters on N_e. We also estimate the ratio of effective to actual population size in six populations of house sparrows on islands off the coast of northern Norway. This ratio showed large interisland variation because of demographic differences among the populations. Finally, we calculate how N_e in a growing house sparrow population will change over time.

Using phylogeny-based methods to identify evolutionary transitions has become an integral part of evolutionary biology. Here, we demonstrate the potential for these methods to give statistically well-supported but misleading inferences about character evolution. We also show how inferences of character evolution can be informed using GIS-based methods to reconstruct ancestral environmental regimes. We reconstruct a phylogeny for marsupial frogs (Hemiphractidae) using nuclear and mitochondrial DNA sequences and estimate patterns of life-history evolution across the resulting tree. We find that Gastrotheca species with complex life cycles (i.e., egg, tadpole, and adult stages) are phylogenetically nested among species and genera with direct development (i.e., egg and adult stages only). Assuming a single rate for gains and losses in likelihood reconstructions, there is strong statistical support for the hypothesis that the tadpole stage was lost early in the phylogeny but reappeared within Gastrotheca. Assuming different rates of gain and loss, the model with significantly higher statistical support, the tadpole stage seems to have been lost multiple times but never regained. Given that both hypotheses cannot be correct, at least one reconstruction model must be giving well-supported but misleading results. Several lines of evidence (including GIS-based reconstructions of the ancestral climatic regime) suggest that the former hypothesis is correct, and that the tadpole stage has evolved from direct development within Gastrotheca, the only known case of such a reversal in frogs.

Resources, sex ratio, and seed production by hermaphrodites covary among natural populations of many gynodioecious plant species, such that they are functionally “more dioecious” as resources become more limiting. Strong correlations among these three factors confound our understanding of their relative roles in maintaining polymorphic sexual systems. We manipulated resource availability and sex ratio and measured their effects on relative fertility and phenotypic selection through the maternal fitness of females and hermaphrodites of Fragaria virginiana. Two results were particularly surprising. First, hermaphrodites showed little variability in fecundity across resource treatments and showed strong positive and context-dependent selection for fruit set. This suggests that variation in hermaphrodite seed production along resource gradients in nature may result from adaptation rather than plasticity. Second, although females increased their fecundity with higher resources, their fertility was unaffected by sex ratio, which is predicted to mediate pollen limitation of females in natural populations where they are common. Selection on petal size of females was also weak, indicating a minimal effect of pollinator attraction on variation in the fertility of female plants. Hence, we found no mechanistic explanation for the complete absence of high-resource high female populations in nature. Despite strong selection for increased fruit set of hermaphrodites, both the strength of selection and its contribution to the maintenance of gynodioecy are severely reduced under conditions where females have high relative fecundity (i.e., low resources and high-female sex ratios). High relative fertility plus high female frequency means that the evolution of phenotypic traits in hermaphrodites (i.e., response to selection via seed function) should be manifested through females because most hermaphrodites will have female mothers. Fruit set was never under strong selection in females; hence, selection to increase fruit set hermaphrodites will be less effective in maintaining their fruiting ability in natural populations with low resources and high female frequency. In sum, both sex ratio and resource availability influence trait evolution indirectly—through their effects on relative fertility of the sexes and patterns of selection. Sex ratio did not impose strong pollen limitation on females but did directly moderate the outcome of natural selection by biasing the maternal sex of the next generation. This direct effect of sex ratio on the manifestation of natural selection is expected to have far greater impact on the evolution of traits, such as seed-producing ability in hermaphrodites and the maintenance of sexual polymorphisms in nature, compared to indirect effects of sex ratio on relative fertility of the sexes.

How introduced plants, which may be locally adapted to specific climatic conditions in their native range, cope with the new abiotic conditions that they encounter as exotics is not well understood. In particular, it is unclear what role plasticity versus adaptive evolution plays in enabling exotics to persist under new environmental circumstances in the introduced range. We determined the extent to which native and introduced populations of St. John's Wort (Hypericum perforatum) are genetically differentiated with respect to leaf-level morphological and physiological traits that allow plants to tolerate different climatic conditions. In common gardens in Washington and Spain, and in a greenhouse, we examined clinal variation in percent leaf nitrogen and carbon, leaf δ¹³C values (as an integrative measure of water use efficiency), specific leaf area (SLA), root and shoot biomass, root/shoot ratio, total leaf area, and leaf area ratio (LAR). As well, we determined whether native European H. perforatum experienced directional selection on leaf-level traits in the introduced range and we compared, across gardens, levels of plasticity in these traits. In field gardens in both Washington and Spain, native populations formed latitudinal clines in percent leaf N. In the greenhouse, native populations formed latitudinal clines in root and shoot biomass and total leaf area, and in the Washington garden only, native populations also exhibited latitudinal clines in percent leaf C and leaf δ¹³C. Traits that failed to show consistent latitudinal clines instead exhibited significant phenotypic plasticity. Introduced St. John's Wort populations also formed significant or marginally significant latitudinal clines in percent leaf N in Washington and Spain, percent leaf C in Washington, and in root biomass and total leaf area in the greenhouse. In the Washington common garden, there was strong directional selection among European populations for higher percent leaf N and leaf δ¹³C, but no selection on any other measured trait. The presence of convergent, genetically based latitudinal clines between native and introduced H. perforatum, together with previously published molecular data, suggest that native and exotic genotypes have independently adapted to a broad-scale variation in climate that varies with latitude.

For sexual selection to be important in plants, it must occur at pollen load sizes typical of field populations. However, studies of the impact of pollen load size on pollen competition have given mixed results, perhaps because so few of these studies directly examined the outcome of mating when pollen load size was varied. We asked whether seed paternity after mixed pollination of wild radish was affected by pollen load sizes ranging from 22 to 220 pollen grains per stigma. We examined the seed siring abilities of 12 pollen donors across 11 maternal plants. Seed paternity was statistically indistinguishable across the pollen load sizes even though, overall, the pollen donors sired different numbers of seeds. This lack of effect of pollen load size on seed paternity may have occurred because fruit abortion and early abortion or failure of fertilization of seeds increased as load size decreased. Thus, failures of fruits and seeds sired by poorer pollen donors may keep seed paternity constant across pollen load sizes.

Mating signals act as behavioral barriers to gene flow in many animal taxa, yet little is known about how signal evolution within populations contributes to the formation of these barriers. Although variation in mating signals among populations is known to affect mating behavior, there is no direct evidence that the evolution of mating signals changes signal effectiveness within a natural population. Making use of historical recordings of bird song, I found that both male and female white-crowned sparrows (Zonotrichia leucophrys) respond more strongly to current than to historical songs, indicating that historical songs are less effective as signals in the current contexts of both mate choice and male–male competition. Finding that historical signals are less effective suggests that signal evolution within populations may ultimately contribute to the formation of behavioral barriers to gene flow between populations.

In many parentally fed species, siblings compete for food not only by begging and scrambling, but also by violently attacking each other. This aggressive competition has mostly been studied in birds, where it is often combined with dominance subordination, aggressive intimidation, and siblicide. Previous experimental and theoretical studies proposed several life-history, morphological, and behavioral variables that may facilitate the evolution of broodmate aggression, and explain its taxonomic distribution. Here we apply phylogenetic comparative analyses for the first time to test the influence of five hypothesized facilitators of the evolution of broodmate aggression, analyzing 69 species in seven avian families using two quantitative measures of aggression: incidence and intensity. We show that incidence and intensity of aggression increase with long nestling periods and indirect feeding, and small brood size is associated with intense aggression. Large food parcels were not correlated with either the incidence or intensity of aggression. Our study suggests that indirect feeding, long nestling periods, and small broods, possibly in combination with other factors, have tended to favor the evolution of aggressive broodmate competition.

We used a long-term population band-resight survey database, a parallel reproduction database, and multistate mark–recapture analysis to assess the costs of reproduction, a keystone concept of life-history evolution, in Nazca boobies (Sula granti) from Punta Cevallos, Isla Española, Galápagos, Ecuador. We used eight years of resight and breeding data to compare models that included sex- and state-specific survival probabilities and probabilities of transition between reproductive states using multistate mark–recapture models. Models that included state-specific effects were compared with models lacking such effects to evaluate costs of reproduction. The top model, optimizing the trade-off of model simplicity and fit to the data using the Akaike Information Criterion (AIC), showed evidence of a temporally varying survival cost of reproduction: nonbreeders showed higher annual survival than breeders did in some years. Because increasing investment among breeders showed no negative association with survival and subsequent breeding success, this evidence indicates a cost to both males and females of initiating, but not of continuing, a reproductive attempt. In some cases, breeders reaching the highest reproductive state (fledging an offspring) showed higher survival or subsequent breeding success than did failed breeders, consistent with differences in overall quality that promote both survival and reproduction. Although a male-biased adult sex ratio was observed in this population of Nazca boobies, models of state- and sex-specific survival and transition probabilities were not supported, indicating that males and females do not incur different costs of reproduction, and that the observed sex ratio bias is not due to sex-specific adult mortality.

The ability to cope with environmental change is fundamental to a species' evolution. Organisms can respond to seasonal environmental variation through phenotypic plasticity. The substantial plasticity in body mass of temperate species has often been considered a simple consequence of change in environmental quality, but could also have evolved as an adaptation to seasonality. We investigated the genetic basis of, and selection acting on, seasonal plasticity in body mass for wild bighorn sheep ewes (Ovis canadensis) at Ram Mountain, Alberta, under two contrasting environmental conditions. Heritability of plasticity, estimated as mass-specific summer and winter mass changes, was low but significant. The additive genetic variance component of relative summer mass change was greater under good environmental conditions (characterized by a population increase and high juvenile survival) than under poor conditions (population decrease and low juvenile survival). Additive genetic variance of relative winter mass change appeared independent of environmental conditions. We found evidence of selection on summer (relative) and winter (relative and absolute) mass change. For a given mass, more plastic individuals (with greater seasonal mass changes) achieve greater fitness through reproduction in the following year. However, genetic correlations between mass parameters were positive. Our study supports the hypothesis that seasonal plasticity in body mass in vertebrates is an adaptation that evolved under natural selection to cope with environmental variation but genetic correlations with other traits might limit its evolutionary potential.

Trade-offs between reproduction and life span are ubiquitous, but little is known about their underlying mechanisms. Here we combine treatment with the juvenile hormone analog (JHa) methoprene and experimental evolution in Drosophila melanogaster to study the potential role of juvenile hormone (JH) in mediating such trade-offs at both the physiological and evolutionary level. Exposure to JHa in the larval medium (and up to 24 h posteclosion) increased early life fecundity but reduced life span of normal (unselected) flies, supporting the physiological role of JH in mediating the trade-off. This effect was much smaller for life span, and not detectable for fecundity, in fly lines previously bred for 19 generations on a medium containing JHa. Furthermore, these selection lines lived longer than unselected controls even in the absence of JHa treatment, without a detectable reduction in early life fecundity. Thus, selection for resistance to JHa apparently induced some evolutionary changes in JH metabolism or signaling, which led to longer life span as a correlated response. This supports the hypothesis that JH may mediate evolution of longer life span, but—contrary to our expectation—this apparently does not need to trade-off with fecundity.

Many species have mitochondrial DNA lineages that are phylogenetically intermixed with other species, but studies have rarely tested the cause of such paraphyly. In this study, we tested two hypotheses that could explain mitochondrial paraphyly of Holarctic gadwalls (Anas strepera) with respect to Asian falcated ducks (A. falcata). First, hybridization could have resulted in falcated duck mitochondrial DNA (mtDNA) introgressing into the gadwall gene pool. Second, gadwalls and falcated ducks could have diverged so recently that mtDNA lineages have not sorted to reciprocal monophyly. We used coalescent analyses of three independent loci to distinguish between these two hypotheses. Two lines of evidence support introgression. First, analyses of the three loci combined show that some introgression is necessary to explain current genetic diversity in gadwalls. Second, we generated alternative predictions regarding time since divergence estimated from mtDNA: falcated ducks and gadwalls would have diverged between 65,000 and 700,000 years before present (ybp) under the introgression hypothesis and between 11,000 and 76,000 ybp under the incomplete lineage sorting hypothesis. The two independent nuclear introns indicated that these species diverged between 210,000 and 5,200,000 ybp, which did not overlap the predicted time for incomplete lineage sorting. These analyses also suggested that ancient introgression (∼14,000 ybp) has resulted in the widespread distribution and high frequency of falcated-like mtDNA (5.5% of haplotypes) in North America. This is the first study to use a rigorous quantitative framework to reject incomplete lineage sorting as the cause of mitochondrial paraphyly.

Developmental failure caused by excess sperm (polyspermy) is thought to be an important mechanism driving the evolution of gamete-recognition proteins, reproductive isolation, and speciation in marine organisms. However, these theories assume that there is heritable variation in the susceptibility to polyspermy and that this variation is related to the overall affinity between sperm and eggs. These assumptions have not been critically examined. We investigated the relationship between ease of fertilization and susceptibility to polyspermy within and among three congeneric sea urchins. The results from laboratory studies indicate that, both within and among species, individuals and species that produce eggs capable of fertilization at relatively low sperm concentrations are more susceptible to polyspermy, whereas individuals and species producing eggs that require higher concentrations of sperm to be fertilized are more resistant to polyspermy. This relationship sets the stage for selection on gamete traits that depend on sperm availability and for sexual conflict that can influence the evolution of gamete-recognition proteins and eventually lead to reproductive isolation.

The constraining effect of gene flow on adaptive divergence is often inferred but rarely quantified. We illustrate ways of doing so using stream populations of threespine stickleback (Gasterosteus aculeatus) that experience different levels of gene flow from a parapatric lake population. In the Misty Lake watershed (British Columbia, Canada), the inlet stream population is morphologically divergent from the lake population, and presumably experiences little gene flow from the lake. The outlet stream population, however, shows an intermediate phenotype and may experience more gene flow from the lake. We first used microsatellite data to demonstrate that gene flow from the lake is low into the inlet but high into the outlet, and that gene flow from the lake remains relatively constant with distance along the outlet. We next combined gene flow data with morphological and habitat data to quantify the effect of gene flow on morphological divergence. In one approach, we assumed that inlet stickleback manifest well-adapted phenotypic trait values not constrained by gene flow. We then calculated the deviation between the observed and expected phenotypes for a given habitat in the outlet. In a second approach, we parameterized a quantitative genetic model of adaptive divergence. Both approaches suggest a large impact of gene flow, constraining adaptation by 80–86% in the outlet (i.e., only 14–20% of the expected morphological divergence in the absence of gene flow was observed). Such approaches may be useful in other taxa to estimate how important gene flow is in constraining adaptive divergence in nature.

We assessed the extent to which traits related to ejaculate investment have evolved in lines of Drosophila melanogaster that had an evolutionary history of maintenance at biased sex ratios. Measures of ejaculate investment were made in males that had been maintained at male-biased (MB) and female-biased (FB) adult sex ratios, in which levels of sperm competition were high and low, respectively. Theory predicts that when the risk of sperm competition is high and mating opportunities are rare (as they are for males in the MB populations), males should increase investment in their few matings. We therefore predicted that males from the MB lines would (1) exhibit increased investment in their first mating opportunities and (2) deplete their ejaculates at a faster rate when mating multiply, in comparison to FB males. To investigate these predictions we measured the single mating productivity of males from three replicates each of MB and FB lines mated to five wild-type virgin females in succession. In contrast to the first prediction, there was no evidence for differences in productivity between MB and FB line males in their first matings. The second prediction was upheld: mates of MB and FB males suffered increasingly reduced productivity with successive matings, but the decline was significantly more pronounced for MB than for FB males. There was a significant reduction in the size of the accessory glands and testes of males from the MB and FB regimes after five successive matings. However, the accessory glands, but not testes, of MB males became depleted at a significantly faster rate than those of FB males. The results show that male reproductive traits evolved in response to the level of sperm competition and suggest that the ability to maintain fertility over successive matings is associated with the rate of ejaculate, and particularly accessory gland, depletion.

Gene flow between coexisting or nearby populations normally prevents genetic divergence and local adaptation. Despite this, there are an increasing number of reports of sympatric sister taxa, indicating potential divergence and speciation in the face of gene flow. A large number of such reported cases involve lake-dwelling fish, which are expected to run into few physical barriers to dispersal within their aquatic habitat. However, such cases may not necessarily reflect sympatric speciation if cryptic dispersal barriers are common in lakes and other aquatic systems. In this study, we examined genetic differentiation in perch (Perca fluviatilis L.) from nine locations in a single, small lake (24 km²), using microsatellites. We detected significant genetic differentiation in all but two pairwise comparisons. These patterns were not consistent with divergence by distance or the existence of kin groups. Instead, they suggest that cryptic barriers to dispersal exist within the lake, allowing small-scale genetic divergence. Such an observation suggests that allopatric (or parapatric) divergence may be possible, even in small, apparently homogenous environments such as lakes. This has important consequences for how we currently view evidence from nature for sympatric speciation.