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Sexual selection can be shaped by spatial variation in environmental features among populations. Differences in sexual selection among populations generated through the effects of the environment could be shaped via four paths: differences in mean absolute fitness, differences in the means or variances of phenotypes, or differences in the absolute fitness-trait function relationship. Because sexual selection occurs only during the adult life stage, most studies have focused on identifying environmental features that influence these metrics of fitness and trait distributions among adults. However, these adult features could also be affected by environmental factors experienced in early life stages that then shape the trajectory for sexual selection during the adult life stage. Here we investigated how among-population variation in environmental conditions during the juvenile (larval) stage of two species of

Parasite-driven population divergence in hosts can be exacerbated by environmental factors affecting host parasitism, as well as by increasing sexual selection against parasitized hosts. Environmental factors can influence parasitism directly by affecting parasite survival, and indirectly by affecting host condition, which can in turn shape host sexual selection. To disentangle these potential alternative paths, we used a damselfly (host) - water mite (parasite) system to examine how environmental factors directly and indirectly drive heterogeneity in parasitism across populations and influence the strength of sexual selection acting against parasitized males. We found substantial heterogeneity in parasitism across populations, driven mainly by lake pH, and damselfly density. Although this heterogeneity in parasitism did not translate directly into variation in sexual selection, the density of predatory fish increased sexual selection strength, likely through the effects on damselfly condition. These results imply that parasitism alone may not cause differences in sexual selection across populations, but when linked with underlying environmental conditions, parasitism can increase the strength of selection. More broadly, these results suggest that elucidating how parasitism may drive sexual selection requires consideration of the intwined effects of ecological processes.

Whole-genome duplications (WGDs) have long been considered the causal mechanism underlying dramatic increases to morphological complexity due to the neo-functionalization of paralogs generated during these events. Nonetheless, an alternative hypothesis suggests that behind the retention of most paralogs is not neo-functionalization, but instead the degree of the inter-connectivity of the intended gene product, as well as the mode of the WGD itself. Here, we explore both the causes and consequences of WGD by examining the distribution, expression, and molecular evolution of microRNAs (miRNAs) in both gnathostome vertebrates as well as chelicerate arthropods. We find that although the number of miRNA paralogs tracks the number of WGDs experienced within the lineage, few of these paralogs experienced changes to the seed sequence, and thus are functionally equivalent relative to their mRNA targets. Nonetheless, in gnathostomes, although the retention of paralogs following the 1R autotetraploidization event is similar across the two subgenomes, the paralogs generated by the gnathostome 2R allotetraploidization event are retained in higher numbers on one subgenome relative to the second, with the miRNAs found on the preferred subgenome showing both higher expression of mature miRNA transcripts and slower molecular evolution of the precursor miRNA sequences. Importantly, WGDs do not result in the creation of miRNA novelty, nor do WGDs correlate to increases in complexity. Instead, it is the number of miRNA seed sequences in the genome itself that not only better correlate to instances in complexification, but also mechanistically explain why complexity increases when new miRNA families are established.

Resource dynamics influence the contemporary ecology of consumer-resource mutualisms. Suites of resource traits, such as floral nectar components, also evolve in response to different selective pressures, changing the ecological dynamics of the interacting species at the evolutionary equilibrium. Here we explore the evolution of resource-provisioning traits in a biotically pollinated plant that produces nectar as a resource for beneficial consumers. We develop a mathematical model describing natural selection on two quantitative nectar traits: maximum nectar production rate and maximum nectar reservoir volume. We use this model to examine how nectar production dynamics evolve under different ecological conditions that impose varying cost-benefit regimes on resource provisioning. The model results predict that natural selection favors higher nectar production when ecological factors limit the plant or pollinator’s abundance (e.g., a lower productivity environment or a higher pollinator conversion efficiency). We also find that nectar traits evolve as a suite in which higher costs of producing one trait select for a compensatory increase in investment in the other trait. This empirically explicit approach to studying the evolution of consumer-resource mutualisms illustrates how natural selection acting via direct and indirect pathways of species interactions generates patterns of resource provisioning seen in natural systems.

Whole genome duplications (WGDs) have long been considered the causal mechanism underlying the dramatic increase in vertebrate morphological complexity relative to invertebrates. This is due to the retention and neo-functionalization of paralogues generated during these events, evolving new regulatory circuits, and ultimately morphological novelty. Nonetheless, an alternative hypothesis suggests that behind the retention of most paralogues is not neo-functionalization, but instead the degree of the inter-connectivity of the intended gene product, as well as the mode of the WGD itself. Here, we explore both the causes and consequences of WGD by examining the distribution, expression, and molecular evolution of microRNAs (miRNAs) in both gnathostome vertebrates as well as chelicerate arthropods. We find that although the number of miRNA paralogues tracks the number of WGDs experienced within the lineage, few of these paralogues experienced changes to the seed sequence, and thus are functionally equivalent relative to their mRNA targets. Nonetheless, the paralogues generated by the gnathostome 2R allotetraploidization event are retained in higher numbers on one sub-genome relative the second, with the miRNAs found on the preferred set of paralogons showing both higher expression of mature miRNA transcripts and slower molecular evolution of the precursor miRNA sequences. Importantly, WGDs do not result in the creation of miRNA novelty, nor do WGDs correlate to increases in complexity. Instead, it is the number of miRNA seed sequences in the genome itself that not only better correlate to instances in complexification, but also mechanistically explain why complexity increases when new miRNA families are established.

The neutral theory of biodiversity explored the structure of a community of ecologically equivalent species. Such species are expected to display community drift dynamics analogous to neutral alleles undergoing genetic drift. While entire communities of species are not ecologically equivalent, recent field experiments have documented the existence of guilds of such neutral species embedded in real food webs. What demographic outcomes of the interactions within and between species in these guilds are expected to produce ecological drift versus coexistence remains unclear. To address this issue, and guide empirical testing, we consider models of a guild of ecologically equivalent competitors feeding on a single resource to explore when community drift should manifest. We show that community drift dynamics only emerge when the density-dependent effects of each species on itself are identical to its density-dependent effects on every other guild member. In contrast, if each guild member directly limits itself more than it limits the abundance of other guild members, all species in the guild are coexisting, even though they all are ecologically equivalent with respect to their interactions with species outside the guild (i.e. resources, predators, mutualists). Hence, considering only interspecific ecological differences generating density dependence, and not fully accounting for the preponderance of mechanisms causing intraspecific density dependence, will provide an incomplete picture for segregating between neutrality and coexistence. We also identify critical experiments necessary to disentangle guilds of ecologically equivalent species from those experiencing ecological drift, as well as provide an overview of ways of incorporating a mechanistic basis into studies of species coexistence and neutrality. Identifying these characteristics, and the mechanistic basis underlying community structure, is not merely an exercise in clarifying the semantics of coexistence and neutral theories, but rather reflects key differences that must exist among community members in order to determine how and why communities are structured.

Much of ecological theory presumes that natural selection should foster species coexistence by phenotypically differentiating competitors so that the stability of the community is increased, but whether this will actually occur is a question of the ecological dynamics of natural selection. I develop an evolutionary model of consumer-resource interactions based on MacArthur's and Tilman's classic works, including both resource and apparent competition, to explore what fosters or retards the differentiation of resources and their consumers. Analyses of this model predict that consumers will differentiate only on specific ranges of environmental gradients (e.g., greater productivity, weaker stressors, lower structural complexity), and where it occurs, the magnitude of differentiation also depends on gradient position. In contrast to "limiting similarity" expectations, greater intraspecific phenotypic variance results in less differentiation among the consumers because of how phenotypic variation alters the fitness landscapes driving natural selection. In addition, the final structure of the community that results from the coevolution of these interacting species may be highly contingent on the initial properties of the species as the community is being assembled. These results highlight the fact that evolutionary conclusions about community structure cannot be based on ecological arguments of community stability or coexistence but rather must be explicitly based on the ecological dynamics of natural selection.

The risk of meiotic segregation errors increases dramatically during a woman’s thirties, a phenomenon known as the maternal age effect. In addition, several lines of evidence indicate that meiotic cohesion deteriorates as oocytes age. One mechanism that may contribute to age-induced loss of cohesion is oxidative damage. In support of this model, we recently reported (Perkins et al. in Proc Natl Acad Sci U S A 113(44):E6823–E6830, 2016) that the knockdown of the reactive oxygen species (ROS)–scavenging enzyme, superoxide dismutase (SOD), during meiotic prophase causes premature loss of arm cohesion and segregation errors in Drosophila oocytes. If age-dependent oxidative damage causes meiotic segregation errors, then the expression of extra SOD1 (cytosolic/nuclear) or SOD2 (mitochondrial) in oocytes may attenuate this effect. To test this hypothesis, we generated flies that contain a UAS-controlled EMPTY, SOD1, or SOD2 cassette and induced expression using a Gal4 driver that turns on during meiotic prophase. We then compared the fidelity of chromosome segregation in aged and non-aged Drosophila oocytes for all three genotypes. As expected, p{EMPTY} oocytes subjected to aging exhibited a significant increase in nondisjunction (NDJ) compared with non-aged oocytes. In contrast, the magnitude of age-dependent NDJ was significantly reduced when expression of extra SOD1 or SOD2 was induced during prophase. Our findings support the hypothesis that a major factor underlying the maternal age effect in humans is age-induced oxidative damage that results in premature loss of meiotic cohesion. Moreover, our work raises the exciting possibility that antioxidant supplementation may provide a preventative strategy to reduce the risk of meiotic segregation errors in older women. Electronic supplementary material The online version of this article (10.1007/s00412-019-00702-y) contains supplementary material, which is available to authorized users.

1. Recent studies demonstrate that ecological and evolutionary processes can occur over similar temporal and spatial scales and might thus frequently interact.Although concepts such as the evolving metacommunity, diffuse (co)evolution and community genetics integrate multi-species dynamics, most experimental studies usually consider how evolution affects only one focal species. Hence, our understanding of evolution in multi-species communities is still underdeveloped. 2. We highlight key community and evolutionary mechanisms and their interactions to facilitate a broader understanding of evolution in multi-species communities. We propose a framework that explicitly considers interactions between each of the four analogous processes of evolutionary biology (selection, gene flow, genetic drift and mutation) and community ecology (species sorting, dispersal, ecological drift and speciation). 3. Focusing on interactions between processes of evolutionary biology and community ecology enables explorations of the full range of eco-evolutionary dynamics in multi-species communities and guides the design of novel experiments. Furthermore, the proposed framework develops a shared language between evolutionary biologists and community ecologists and indicates new research avenues. 4. Overall, we propose that explicitly incorporating interactions between these evolutionary and community processes to study eco-evolutionary dynamics in multi-species communities will better inform broader questions about the maintenance of diversity and the resilience of diverse communities to disturbances, both natural and manmade.

That species must differ ecologically is often viewed as a fundamental condition for their stable coexistence in biological communities. Yet, recent work has shown that ecologically equivalent species can coexist when reproductive interactions and sexual selection regulate population growth. Here, we review theoretical models and highlight empirical studies supporting a role for reproductive interactions in maintaining species diversity. We place reproductive interactions research within a burgeoning conceptual framework of coexistence theory, identify four key mechanisms in intra- and interspecific interactions within and between sexes, speculate on novel mechanisms, and suggest future research. Given the preponderance of sexual reproduction in nature, our review suggests that this is a neglected path towards explaining species diversity when traditional ecological explanations have failed.

Females in many animal species must discriminate between conspecific and heterospecific males when choosing mates. Such mating preferences that discriminate against heterospecifics may inadvertently also affect the mating success of conspecific males, particularly those with more extreme phenotypes. From this expectation, we hypothesized that female mate choice should cause Enallagma females (Odonata: Coenagrionidae) to discriminate against conspecific males with more extreme phenotypes of the claspers males use to grasp females while mating - the main feature of species mate recognition in these species. To test this, we compared cerci sizes and shapes between males that were captured while mating with females to males that were captured at the same time but not mating in three Enallagma species. In contrast to our hypothesis, we found only one of forty comparisons of shape variation that was consistent with females discriminating against males with more extreme cerci shapes. Instead, differences in cerci shape between mating and single males suggested that females displayed directional preferences on 1-4 aspects of cerci shape in two of the species in our samples. These results suggest that whereas some directional biases in mating based on cerci shape occur, the intraspecific phenotypic variation in male cerci size and shape is likely not large enough for females to express any significant incidental discrimination among conspecifics with more extreme shapes.

Despite the many study systems in which predation has played a major role in phenotypic diversification and speciation, the underlying selective regimes imposed by different predator assemblages have rarely been quantified. We did so for the damselfly genus Enallagma which strongly diverged in antipredator traits when the ancestral species occupying lakes containing fish (hereafter fish lakes) repeatedly invaded fishless lakes with dragonfly larvae as top predators (hereafter dragonfly lakes). In two selection experiments in field enclosures we quantified the selection on two key escape traits of two fish-lake Enallagma species associated with survival selection by fish in the ancestral fish lakes and by dragonfly predators in the invaded fishless, dragonfly lakes. In accordance with the different hunting modes, fish imposed selection for a decreased swimming propensity while dragonfly larvae imposed selection for increased swimming speed in one of the two species. In two complementary quantitative genetic rearing experiments, we found relatively low but significant broad-sense heritabilities for both escape traits. Integrating these estimates for the selection coefficients and the heritabilities suggests that the evolutionary increase in swimming speed associated with the habitat shift may have occurred rapidly. Our study suggests that the phenotypic evolution of ecologically important traits related to habitat shifts may occur at an ecological timescale.

This chapter explores the evolutionary dynamics that arise when different types of species mix together in a community either by invasion or by perturbation, as well as community mixing caused by climate change. In particular, it considers the features that promote or retard ecological differentiation of species. The chapter first describes a general model of evolutionary and ecological dynamics in a community before discussing adaptive differentiation at multiple trophic levels. It then examines differentiation of species with identical underlying parameters vs. different underlying parameters, along with intraguild predation and how ecological opportunity evolves within biological communities. It also investigates when neutral species will initially differentiate from one another to convert them into a set of coexisting species, and when differentiated species will initially converge to become ecologically more similar. The chapter shows that, when differentiation occurs, the type of traits underlying species interactions determine the ecological structure of the resulting community.

This chapter examines ecological opportunities that are available to species in various positions within a biological community, with particular emphasis on identifying the criteria necessary for an ecological opportunity to exist. Before discussing what performance capabilities a species must have to fill different types of ecological opportunities and what is required for invasibility of species into different functional positions in a community, the chapter considers the different frameworks that have been used to model species interactions. It then describes resource and apparent competition to show how resource availability from below and predation pressure from above can affect the types of species that can exploit specifc ecological opportunities. It also analyzes communities with three trophic levels, intraguild predation or omnivory, mutualism, the mechanisms that foster coexistence between one plant species and one pollinator species, and the case of one plant species with multiple pollinators.

This chapter examines how species in simple community modules evolve in order to adapt to one another. It first considers the ecological basis of natural selection before discussing the three general types of traits that underlie the trophic interactions between predators and their prey. It then explores how the type of traits involved in species interactions affect the dynamics of adaptation and describes the underlying dynamics of adaptation when all the species in a simple community module can coevolve, along with the influences of various system features on the outcome of this coevolution. The chapter focuses on the simultaneous dynamics of abundances and traits that result from species interactions by looking at the case involving one resource and one consumer, with only trait being ecologically important for each species, and how this simple consumer-resource system evolves when a third trophic level is added to the system.

This chapter examines the ecological and evolutionary dynamics of species across a metacommunity, and how these dynamics affect regional community structure. It begins with a discussion of the evolution of dispersal, focusing on when movement between local communities is and is not favored by natural selection, and how these various movement patterns shape local community structure. An example of the demographic consequences of dispersal is presented, and the evolution of dispersal in a temporally constant environment is analyzed. The chapter also considers the evolution of dispersal rates among communities along with local adaptation within each and explains how link species affect local abundances via their movement strategies. Finally, it explores the interplay between local adaptation and dispersal evolution, the impact of simultaneous spatial and temporal variation in environmental conditions on the evolution of dispersal among populations, and the evolution of phenotypic plasticity.

This chapter discusses the important insights that can be gained from the synthetic analysis of evolution in communities. First, fitness, despite being commonly associated only with evolution by natural selection, also plays a key role in both population and community ecology. Second, the same processes jointly drive the dynamics of species abundances and traits, and these dynamics can only be understood in the context of a community of interacting species. All combinations of abiotic factors and other species' abundances and traits that would permit a species to invade the community without any evolution are contained in the so-called ecological volume of invasibility, in contrast to an evolutionary volume of invasibility that defines the conditions for invasion or adaptation to occur. This chapter also considers speciation, disaggregation and recombination, the traits that influence species interactions, ecological differentiation of species, and the use of phylogeny to elucidate community assembly and structure.

This chapter considers the main processes that operate at the regional and biogeographic scales to ultimately shape local community structure—namely, speciation and biogeographic mixing of taxa. It first defines what a “species” is before discussing the range of mechanisms that give rise to new species, and more specifically reproductive isolation. In particular, it examines the extent to which the speciation process directly induces differences in ecologically important traits between the progenitor and daughter species. It then explains how the phenotypic differences generated at the time of speciation determine what type of community member the new species begins as. It also shows how past climate change affected current local and regional community structure by periodic forcing of mass movements of species across Earth and causing increases in speciation and extinction rates. Finally, it evaluates the dynamics of invasive species and their role in habitat alteration today.

Evolutionary Community Ecology develops a unified framework for understanding the structure of ecological communities and the dynamics of natural selection that shape the evolution of the species inhabiting them. All species engage in interactions with many other species, and these interactions regulate their abundance, define their trajectories of natural selection, and shape their movement decisions. Mark McPeek synthesizes the ecological and evolutionary dynamics generated by species interactions that structure local biological communities and regional metacommunities. McPeek explores the ecological performance characteristics needed for invasibility and coexistence of species in complex networks of species interactions. This species interaction framework is then extended to examine the ecological dynamics of natural selection that drive coevolution of interacting species in these complex interaction networks. The models of natural selection resulting from species interactions are used to evaluate the ecological conditions that foster diversification at multiple trophic levels. Analyses show that diversification depends on the ecological context in which species interactions occur and the types of traits that define the mechanisms of those species interactions. Lastly, looking at the mechanisms of speciation that affect species richness and diversity at various spatial scales and the consequences of past climate change over the Quaternary period, McPeek considers how metacommunity structure is shaped at regional and biogeographic scales. Integrating evolutionary theory into the study of community ecology, Evolutionary Community Ecology provides a new framework for predicting how communities are organized and how they may change over time.

External male reproductive structures have received considerable attention as an early-acting cause of reproductive isolation (RI), because the morphology of these structures often evolves rapidly between populations. This presents the potential for mechanical incompatibilities with heterospecific female structures during mating and could thus prevent interbreeding between nascent species. Although such mechanical incompatibilities have received little empirical support as a common cause of RI, the potential for mismatch of reproductive structures to cause RI due to incompatible species-specific tactile cues has not been tested. We tested the importance of mechanical and tactile incompatibilities in RI betweenEnallagma annaandE. carunculatum,two damselfly species that diverged within the past ~250,000 years and currently hybridize in a sympatric region. We quantified 19 prezygotic and postzygotic RI barriers using both naturally occurring and lab-reared damselflies. We found incomplete mechanical isolation between the two pure species and between hybrid males and pure species females. Interestingly, where mechanical isolation was incomplete, females showed greater resistance and refusal to mate with hybrid or heterospecific males compared to conspecific males, which suggests that tactile incompatibilities involving male reproductive structures can influence female mating decisions and form a strong barrier to gene flow in early stages of speciation.

External male reproductive structures have received considerable attention as a cause of reproductive isolation (RI), because the morphology of these structures often evolves rapidly between populations. This rapid evolution presents the potential for mechanical incompatibilities with heterospecific female structures during mating and could thus prevent interbreeding between nascent species. Although such mechanical incompatibilities have received little empirical support as a common cause of RI, the potential for mismatch of reproductive structures to cause RI due to incompatible species-specific tactile cues has not been tested. We tested the importance of mechanical and tactile incompatibilities in RI between Enallagma anna and E. carunculatum, two damselfly species that diverged within the past ∼250,000 years and currently hybridize in a sympatric region. We quantified 19 prezygotic and postzygotic RI barriers using both naturally occurring and laboratory-reared damselflies. We found incomplete mechanical isolation between the two pure species and between hybrid males and pure species females. Interestingly, in mating pairs for which mechanical isolation was incomplete, females showed greater resistance and refusal to mate with hybrid or heterospecific males compared to conspecific males. This observation suggests that tactile incompatibilities involving male reproductive structures can influence female mating decisions and form a strong barrier to gene flow in early stages of speciation.

Natural selection has both genetic and ecological dynamics. The fitnesses of individuals change with their ecological context, and so the form and strength of selective agents change with abiotic factors and the phenotypes and abundances of interacting species. I use standard models of consumer-resource interactions to explore the ecological dynamics of natural selection and how various trait types influence these dynamics and the resulting structure of a community of coevolving species. Evolutionary optima favored by natural selection depend critically on the abundances of interacting species, and the traits of species can undergo dynamic cycling in limited areas of parameter space. The ecological dynamics of natural selection can also drive shifts from one adaptive peak to another, and these ecologically driven adaptive peak shifts are fundamental to the dynamics of niche differentiation. Moreover, this ecological differentiation is fostered in more productive and more benign environments where species interactions are stronger and where the selection gradients generated by species interactions are stronger. Finally, community structure resulting from coevolution depends fundamentally on the types of traits that underlie species interactions. The ecological dynamics of the process cannot be simplified, neglected, or ignored if we are to build a predictive theory of natural selection.

The biological traits of different species usually predispose them to dissimilar levels of extinction risk. Accordingly, we analysed genetic variation and population genetic structure of two endangered forest monkeys to determine how they were influenced by habitat change. The Tana River red colobus and mangabey are of similar body size, are sympatric and endemic to forests in Kenya. The mangabey is frugivorous, terrestrial and exhibits female philopatry but the colobus is folivorous, arboreal and both sexes exhibit natal dispersal. We predicted that both species would have low genetic variation due to high genetic drift, and possible inbreeding, because their populations are relatively small and subdivided in forest fragments. Nevertheless, we expected the mangabeys to have a higher genetic variation because they have a larger population, and their higher vagility would enhance gene flow. Thus, we expected a stronger population genetic structure in the colobus because lower vagility would constrain dispersal and increase genetic differentiation. Finally, we expected both species to exhibit isolation by distance due to greater gene flow among populations in close geographic proximity. We genotyped 146 colobus and 76 mangabeys with nine microsatellite markers and found that both species had relatively high heterozygosities but low allelic richness. Furthermore, allelic richness, heterozygosity and genetic differentiation were all higher in the mangabey and only the mangabey exhibited isolation by distance. Taken together, our results suggest that species with divergent behavioral ecologies may exhibit differential vulnerabilities to genetic erosion due to habitat change.

The importance of negative intraspecific density dependence to promoting species coexistence in a community is well accepted. However, such mechanisms are typically omitted from more explicit models of community dynamics. Here I analyze a variation of the Rosenzweig-MacArthur consumer-resource model that includes negative intraspecific density dependence for consumers to explore its effect on the coexistence of multiple consumers feeding on a single resource. This analysis demonstrates that a guild of multiple consumers can easily coexist on a single resource if each limits its own abundance to some degree, and stronger intraspecific density dependence permits a wider variety of consumers to coexist. The mechanism permitting multiple consumers to coexist works in a fashion similar to apparent competition or to each consumer having its own specialized predator. These results argue for a more explicit emphasis on how negative intraspecific density dependence is generated and how these mechanisms combine with species interactions to shape overall community structure.

In stream organisms, the landscape affecting intraspecific genetic and phenotypic divergence is comprised of two fundamental components: the stream network and terrestrial matrix. These components are known to differentially influence genetic structure in stream species, but to our knowledge, no study has compared their effects on genetic and phenotypic divergence. We examined how the stream network and terrestrial matrix affect genetic and phenotypic divergence in two stream salamanders, Gyrinophilus porphyriticus and Eurycea bislineata, in the Hubbard Brook Watershed, New Hampshire, USA. On the basis of previous findings and differences in adult terrestriality, we predicted that genetic divergence and phenotypic divergence in body morphology would be correlated in both species, but structured primarily by distance along the stream network in G. porphyriticus, and by overland distance in E. bislineata. Surprisingly, spatial patterns of genetic and phenotypic divergence were not strongly correlated. Genetic divergence, based on amplified DNA fragment length polymorphisms, increased with absolute geographic distance between sites. Phenotypic divergence was unrelated to absolute geographic distance, but related to relative stream vs. overland distances. In G. porphyriticus, phenotypic divergence was low when sites were close by stream distance alone and high when sites were close by overland distance alone. The opposite was true for E. bislineata. These results show that small differences in life history can produce large differences in patterns of intraspecific divergence, and the limitations of landscape genetic data for inferring phenotypic divergence. Our results also underscore the importance of explicitly comparing how terrestrial and aquatic conditions affect spatial patterns of divergence in species with biphasic life cycles.

North American Enallagma damselflies radiated during the Pleistocene, and species differ mainly by reproductive structures. Although morphologically very different, Enallagma hageni and Enallagma ebrium are genetically very similar. Partitioning of genetic variation (AFLP), isolation by distance and clustering analyses indicate that these morphospecies are locally differentiated genetically. Spatial analyses show that they are rarely sympatric at local sites, and their distributions form a mosaic of patches where one is clearly dominant over hundreds of square kilometers. However, these morphospecies are also not genetically more similar when they are sympatric, indicating that hybridization is probably not occurring. Given that these morphospecies are ecologically equivalent, strong assortative mating, reproductive interference and fast post-glacial recolonization may explain the origin and maintenance of these distributional patches across eastern North America. By limiting opportunities for gene flow, reproductive interference may play an unsuspected role in accelerating genetic differentiation in the early phases of nonecological speciation.

Dispersal distributions are often characterized by many individuals that stay close to their origin and large variation in the distances moved by those that leave. This variation in dispersal distance can strongly influence demographic, ecological, and evolutionary processes. However, a lack of data on the fitness and phenotype of individual dispersers has impeded research on the role of natural selection in maintaining variation in dispersal distance. Six years of spatially explicit capture-mark-recapture data showed that survival increased with dispersal distance in the stream salamander Gyrinophilus porphyriticus. To understand the evolutionary implications of this fitness response, we tested whether variation in dispersal distance has a phenotypic basis. We used photographs of marked individuals to measure head, trunk, and leg morphology. We then tested whether dispersal distances over the six-year study period were predicted by these traits. Dispersal distance was significantly related to leg morphology: individuals with relatively long forelimbs and short hindlimbs dispersed the farthest. These results support the hypothesis that positive fitness consequences maintain phenotypes enabling long-distance dispersal. More broadly, they suggest that natural selection can promote variation in dispersal distance and associated phenotypes, offering an alternative to the view that dispersal distance is driven by stochastic or landscape-specific mechanisms.

Summary 1. Ecological differences among co-occurring taxa are often invoked as an explanation for the maintenance of biodiversity. Whether these differences facilitate coexistence, which allows unequal competitors to remain in systems and thus maintain biodiversity, is still unclear. 2. Here, we used observational and experimental studies to test for ecological partitioning in ways that would promote coexistence among three co-occurring damselfly genera. We evaluated two necessary conditions for coexistence: (i) that the damselfly genera differ in their abilities to engage in interactions with other damselfly genera and environmental conditions such that their relative abundances covary differently along environmental gradients and (ii) that an increase in intrageneric abundance is more detrimental to performance-related demographic features of each genus than increases in intergeneric abundances. 3. Observational studies across 40 lakes showed that relative abundances of each genus covaried differently along an environmental gradient of lake abiotic and biotic features consistent with ecological partitioning. Field experiments in which we manipulated both intra- and intergeneric densities demonstrated that per capita growth rates of each genus are negatively density-dependent and are only limited by increases in intra- not intergeneric densities. 4. Collectively, these results show a clear signature of ecological partitioning among each genus, which should prevent competitive exclusion and maintain each genus in this system. The results do not guarantee local coexistence among the three genera but are consistent with criteria that should promote their coexistence. Our results also suggest that a food web model coupling keystone predation and apparent competition is likely necessary to explain the ecological dynamics of persistence among these genera.

Temporal variation in selection can be generated by temporal variation in either the fitness surface or phenotypic distributions around a static fitness surface, or both concurrently. Here, we use within- and between-generation sampling of fitness surfaces and phenotypic distributions over 2 years to investigate the causes of temporal variation in the form of sexual selection on body size in the damselfly Enallagma aspersum. Within a year, when the average female body size differed substantially from the average male body size, male body size experienced directional selection. In contrast, when male and female size distributions overlapped, male body size experienced stabilizing selection when variances in body size were large, but no appreciable selection when the variances in body size were small. The causes of temporal variation in the form of selection can only be inferred by accounting for changes in both the fitness surface and changes in the distribution of phenotypes.

We used mtDNA sequence data from the Tana River red colobus and mangabey to determine how their population genetic structure was influenced by dispersal and habitat fragmentation. The colobus and mangabey are critically endangered primates endemic to gallery forests in eastern Kenya. The forests are a Pliocene-Pleistocene refugium that has recently undergone significant habitat loss and fragmentation due to human activities. We expected both primates to exhibit low levels of genetic diversity due to elevated genetic drift in their small populations, and to show a strong correspondence between genetic and geographic distance due to disruption of gene flow between forests by habitat fragmentation. Additionally, because mangabey females are philopatric, we expected their mtDNA variation to be homogeneous within forest patches but to be heterogeneous between patches. In contrast, colobus have a female-biased dispersal and so we expected their mtDNA variation to be homogeneous within and between forest patches. We found high levels of haplotype and nucleotide diversity as well as high levels of sequence divergence between haplotype groups in both species. The red colobus had significantly higher genetic variation than the mangabey did. Most of the genetic variation in both primates was found within forest fragments. Although both species showed strong inter-forest patch genetic structure we found no correspondence between genetic and geographic distances for the two primates. We attributed the high genetic diversity to recent high effective population size, and high sequence divergence and strong genetic structures to long-term habitat changes in the landscape.

Despite the importance of foraging activity for the growth/predation risk trade-off, studies that demonstrated predator-induced survival selection on foraging activity under semi-natural conditions are relatively rare. Here, we tested for fish-induced selection for reduced foraging activity in two larval Enallagma damselflies using a field enclosure experiment. Fish imposed considerable mortality in both damselfly species and survival selection on foraging activity could be detected in Enallagma geminatum. We did not detect selection in Enallagma hageni, probably because this species already was not eating very much in the absence of fish compared to E. geminatum. Both species responded strongly to the presence of predators by reducing their foraging activity. The documented survival selection on foraging activity was detected despite the already low activity levels in fish lake prey species and despite strong predator-induced plasticity in this trait.

Understanding the patterns of diversification in sexual traits and the selection underlying such diversification represents a major unresolved question in evolutionary biology. We examined the phylogenetic diversification for courtship and external genitalic characters across ten species of Timema walking-sticks, to infer the tempos and modes of character change in these sexual traits and to draw inferences regarding the selective pressures underlying speciation and diversification in this clade. Rates of inferred change in male courtship behaviours were proportional to speciation events, but male external genitalic structures showed a pattern of continuous change across evolutionary time, with divergence proportional to branch lengths. These findings suggest that diversification of courtship behaviour is mediated by processes that occur in association with speciation, whereas diversification of genitalia occurs more or less continuously, most likely driven by forces of sexual selection.

Little is known about the ecological correlates and macro-evolution of life history plasticity to time constraints. Here, we compared age and mass at emergence and their plasticity toward combinations of time constraints (delayed larval development), food level and predation risk (caged dragonfly larvae) by rearing two temporary-pond Lestes damselflies (L. congener and L. forcipatus) and the derived vernal-pond L. dryas from the eggs until adult emergence in outdoor tubs. Life history plasticity under time constraints, low food and predation risk was as predicted by optimality models. Delayed larvae in all three species accelerated development and showed a lower fat content and a substantially elevated mortality rate. At low food, all species emerged later at a smaller mass. Also under predation risk adults of all species emerged later, and (at high food) at a smaller mass. Unexpectedly, delayed larvae did not show a smaller life history response to predation risk imposed by dragonfly larvae. Compared to the two temporary-pond Lestes studied, the derived species that invaded more ephemeral vernal ponds showed a faster development rate and a lower deceleration of development to low food. It also showed a lower acceleration of development to time constraints, possibly reflecting that it reached development rates near to its physiological maximum. Unexpectedly, the vernal-pond Lestes did not slow its development less under predation risk. Our results stress the importance of evaluating ecological and evolutionary correlates of life history plasticity under as realistic conditions as possible.

Various evolutionary forces may shape the evolution of traits that influence the mating decisions of males and females. Phenotypic traits that males and females use to judge the species identify of potential mates should evolve in a punctuated fashion, changing significantly at the time of speciation but changing little between speciation events. In contrast, traits experiencing sexual selection or sexually antagonistic interactions are generally expected to change continuously over time because of the directional selection pressures imposed on one sex by the actions of the other. To test these hypotheses, we used spherical harmonic representations of the shapes of male mating structures in reconstructions of the evolutionary tempo of these structures across the history of the Enallagma damselfly clade. Our analyses show that the evolution of these structures is completely consistent with a punctuated model of evolutionary change and a constant evolutionary rate throughout the clade's history. In addition, no interpopulation variation in shape was detected across the range of one species. These results indicate that male mating structures in this genus are used primarily for identifying the species of potential mates and experience little or no selection from intraspecific sexual selection or sexual antagonism. The implications of these results for speciation are discussed.

We know little about the macroevolution of life-history traits along environmental gradients, especially with regard to the directionality compared to the ancestral states and the associated costs to other functions. Here we examine how age and size at maturity evolved when Lestes damselflies shifted from their ancestral temporary pond habitat (i.e., ponds that may dry once every decade or so) to extremely ephemeral vernal ponds (ponds that routinely dry completely each year). Larvae of three species were reared from eggs until emergence under different levels of photoperiod and transient starvation stress. Compared to the two temporary-pond Lestes, the phylogenetically derived vernal-pond Lestes dryas developed more rapidly across photoperiod treatments until the final instar, and only expressed plasticity in development time in the final instar under photoperiod levels that simulated a later hatching date. The documented change in development rate can be considered adaptive and underlies the success of the derived species in vernal ponds. Results suggest associated costs of faster development are lower mass at maturity and lower immune function after transient starvation stress. These costs may not only have impeded further evolution of the routine development rate to what is physiologically maximal, but also maintained some degree of plasticity to time constraints when the habitat shift occurred.

Compensatory growth where animals compensate for time stress or transient nutritional or thermal stress by accelerating their growth rate is widespread. We know, however, relatively little about the evolution and ecological correlates of compensatory growth. For this we need studies on congeneric species with known phylogenetic relationships that also focus on the associated largely understudied costs. Here we tested for compensatory growth and associated costs in response to time stress (manipulated by photoperiod) and a transient period of starvation or cooling in larvae of the permanent-pond damselfly Lestes eurinus, and compare the results with former studies on temporary-pond Lestes. Larvae showed full compensation in body mass at emergence for all combinations of time stress and starvation or cooling. Unexpectedly, compensatory growth to starvation or cooling was not stronger under time stress. Instead, males under time stress delayed emergence after these transient stressors. In line with a stronger compensatory growth response to time stress than to the other stressors, physiological costs in terms of a reduced investment in immune response (measured as phenoloxidase activity) and energy storage (measured as fat content) were detected only under time stress. Compared to temporary-pond Lestes, L. eurinus showed stronger compensatory growth to time stress. We hypothesize that the stronger compensatory (growth) response in permanent-pond Lestes co-evolved with their derived slower lifestyle when they invaded permanent ponds.

Despite the survival value of high energy reserves during winter, animals often face energy deficits when entering winter. Compensatory growth in energy reserves during the winter period to buffer such deficits may increase winter survival and alleviate the need for costly compensatory mechanisms before or after winter when predation risk is much higher. However, such compensatory responses in energy reserves during winter have not been demonstrated under field conditions. We explored if Lestes eurinus damselfly larvae can compensate for suboptimal energy reserves during winter at 4°C when their ponds are covered with ice. In a field enclosure experiment, we demonstrated compensatory growth in terms of body mass and energy reserves in larvae whose energy status was previously manipulated in the laboratory. These results were supported by patterns in body mass and energy reserves over winter in two natural unmanipulated populations. Winter survival was high overall and not affected by compensatory growth. We hypothesize that the observed compensatory growth in energy reserves during winter may shape life history decisions in autumn and spring, and may make resource availability during winter as or more important than energy reserves before winter.

In this paper, I examine the dynamics of spe- cies richness in a model system in which multiple species compete in a metacommunity (multiple patches linked by dispersal). Patches lie along an environmental gradient, and new species are introduced into the system by speci- ation of existing species. This model is used to explore how the ecological similarity of species influences the pat- terns in community structure that result and to determine whether patterns in fossil and systematics data may be sig- natures for different types of community structure. Making species more similar overall along the entire gradient or making new species that have more similar optimal posi- tions along the gradient to their progenitor both increase the time required to drive species extinction. As a result, making species more similar ecologically to one another increases overall species richness because of an increased frequency of transient species in the system. Having more transient species in systems shifted the longevity distribu- tions of species in the fossil record towards having a greater frequency of shorter duration species, and the age distribution of extant species that would be estimated from molecular phylogenies also had a higher frequency of younger aged species. Comparisons of these results with species longevity distributions extracted from two data sets and with species ages derived from species-level molecular phylogenies strongly suggest that transient species are an important component of real biological communities.

Reconstructing evolutionary patterns of species and populations provides a framework for asking questions about the impacts of climate change. Here we use a multilocus dataset to estimate gene trees under maximum likelihood and Bayesian models to obtain a robust estimate of relationships for a genus of North American damselflies, Enallagma. Using a relaxed molecular clock, we estimate the divergence times for this group. Furthermore, to account for the fact that gene tree analyses can overestimate ages of population divergences, we use a multi-population coalescent model to gain a more accurate estimate of divergence times. We also infer diversification rates using a method that allows for variation in diversification rate through time and among lineages. Our results reveal a complex evolutionary history of Enallagma, in which divergence events both predate and occur during Pleistocene climate fluctuations. There is also evidence of diversification rate heterogeneity across the tree. These divergence time estimates provide a foundation for addressing the relative significance of historical climatic events in the diversification of this genus.

The Enallagma and Lestes damselflies have both diversified and adapted over the past 10-15 million years to the various ecological milieus found along the pond permanence gradient among North American ponds and lakes. Previous articles have explored this diversification process for Enallagma. In this article, we present a phylogenetic hypothesis for the North American Lestes, use this hypothesis to reconstruct Lestes diversification, and compare the diversification processes inferred for Lestes and Enallagma. The results of this study suggest that Lestes began in temporary ponds where large dragonflies are the top predators, while Enallagma began in permanent lakes where fish are the top predators. Starting from these different ancestral habitats, both genera have invaded and adapted to habitats already occupied by the other genus. Moreover, these adaptive habitat shifts involved substantial convergence on the behaviors used to deal with fish and dragonfly predation in both genera and a major life-history shift from diapausing to directly developing eggs in Lestes. However, in Lestes lineages invading fish lakes, swimming speed and morphology did not change to match those of Enallagma species, illustrating that reciprocal shifts between alternative selection regimes are not necessarily evolutionary opposites. Also, the greater sizes and growth rates of Lestes species compared to Enallagma species, which should impart substantial ecological advantages in competition between the genera, were shown to result from phylogenetic inheritance and not from adaptive diversification. This historical analysis of diversification raises new questions about the relationship between the macroevolutionary mechanisms driving lineage diversification and the ecological mechanisms structuring local food webs and regional species assemblages. ispartof: The American naturalist vol:168 issue:6 pages:S50-S72 ispartof: location:United States status: published

The ecological trade-off between foraging gains and predation risk shapes the ecology of many taxa, including larval amphibians and fishes. Is this ecological trade-off reflected at the population level as a negative genetic correlation between traits that contribute to growth and survival? We addressed this question for a single population of Green Frog tadpoles (Rana clamitans) by estimating heritabilities and correlations of traits that mediate the trade-off. Burst speed, growth rate, and size varied among sires and were significantly heritable, but swimming activity in the presence and absence of predator odors was not. Differences among traits in heritability generally accorded with differences in genetic coefficients of variation, with the exception of activity. Growth rate was not correlated with any measure of predator avoidance or escape ability. Likewise, predator escape ability (burst speed) was not correlated with predator avoidance ability (activity and its plasticity). Our results i...