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Understanding patterns of diversification necessarily requires accounting for both the generation and the persistence of species. Formal models of speciation genetics, however, focus on the generation of new species without explicitly considering the maintenance of biodiversity (e.g., coexistence, the focus of ecological studies of diversity). Consequently, it remains unclear whether and how new species will coexist following a speciation event, a gap limiting our ability to understand the rate-limiting controls of diversification over macroevolutionary timescales. To connect coexistence and speciation theory and assess the relative importance of ecological versus genetic constraints in diversification events, we develop a deterministic, three-locus, population-genetic model that includes a skewed distribution of available resources (to generate variation in fitness differences), frequency-dependent competition, and assortative mating. Both ecology and genetics play vital and interacting roles in shaping initial speciation events and long-term eco-evolutionary outcomes. Ecological constraints are especially important when fitness differences are large and competition remains strong among dissimilar phenotypes. Ephemeral species can occur in our model and are typically lost because of competitive exclusion, a result demonstrating that species persistence may serve as the rate-limiting control of long-term diversification rates. More broadly, our model adds evidence that the unification of ecological and evolutionary (including genetic) perspectives on biodiversity is needed to predict large-scale patterns. [ABSTRACT FROM AUTHOR]

The social environment is often the most dynamic and fitness-relevant environment animals experience. Here we tested whether plasticity arising from variation in social environments can promote signal-preference divergence—a key prediction of recent speciation theory but one that has proven difficult to test in natural systems. Interactions in mixed social aggregations could reduce, create, or enhance signal-preference differences. In the latter case, social plasticity could establish or increase assortative mating. We tested this by rearing two recently diverged species of Enchenopa treehoppers—sap-feeding insects that communicate with plant-borne vibrational signals—in treatments consisting of mixed-species versus own-species aggregations. Social experience with heterospecifics (in the mixed-species treatment) resulted in enhanced signal-preference species differences. For one of the two species, we tested but found no differences in the plastic response between sympatric and allopatric sites, suggesting the absence of reinforcement in the signals and preferences and their plastic response. Our results support the hypothesis that social plasticity can create or enhance signal-preference differences and that this might occur in the absence of long-term selection against hybridization on plastic responses themselves. Such social plasticity may facilitate rapid bursts of diversification. [ABSTRACT FROM AUTHOR]

Previous theory has shown that assortative mating for plastic traits can maintain genetic divergence across environmental gradients despite high gene flow. Yet these models did not examine how assortative mating affects the evolution of plasticity. We here describe patterns of genetic variation across elevation for plasticity in a trait under assortative mating, using multiple-year observations of budburst date in a common garden of sessile oaks. Despite high gene flow, we found significant spatial genetic divergence for the intercept, but not for the slope, of reaction norms to temperature. We then used individual-based simulations, where both the slope and the intercept of the reaction norm evolve, to examine how assortative mating affects the evolution of plasticity, varying the intensity and distance of gene flow. Our model predicts the evolution of either suboptimal plasticity (reaction norms with a slope shallower than optimal) or hyperplasticity (slopes steeper than optimal) in the presence of assortative mating when optimal plasticity would evolve under random mating. Furthermore, a cogradient pattern of genetic divergence for the intercept of the reaction norm (where plastic and genetic effects are in the same direction) always evolves in simulations with assortative mating, consistent with our observations in the studied oak populations. [ABSTRACT FROM AUTHOR]

The frequency and asymmetry of mixed-species mating set the initial stage for the ecological and evolutionary implications of hybridization. How such patterns of mixed-species mating, in turn, are influenced by the combination of mate choice errors and relative species abundance remains largely unknown. We develop a mathematical model that generates predictions for how relative species abundances and mate choice errors affect hybridization patterns. When mate choice errors are small (<5%), the highest frequency of hybridization occurs when one of the hybridizing species is at low abundance, but when mate choice errors are high (>5%), the highest hybridization frequency occurs when species occur in equal proportions. Furthermore, females of the less abundant species are overrepresented in mixed-species matings. We compare our theoretical predictions with empirical data on naturally hybridizing Ficedula flycatchers and find that hybridization is highest when the two species occur in equal abundance, implying rather high mate choice errors. We discuss ecological and evolutionary implications of our findings and encourage future work on hybrid zone dynamics that take demographic aspects, such as relative species abundance, into account. [ABSTRACT FROM AUTHOR]

It is thought that two species can coexist if they use different resources present in the environment, yet this assumes that species are completely reproductively isolated. We simulate coexistence outcomes for two sympatric species that are ecologically differentiated but have incomplete reproductive isolation. The consequences of interbreeding crucially depend on hybrid fitness. When hybrid fitness is high, just a small rate of hybridization can lead to collapse of two species into one. Low hybrid fitness can cause population declines, making extinction of one or both species likely. High intrinsic growth rates result in higher reproductive rates when populations are below carrying capacity, reducing the probability of extinction and increasing the probability of stable coexistence at moderate levels of assortative mating and hybrid fitness. Very strong but incomplete assortative mating can induce low hybrid fitness via a mating disadvantage to rare genotypes, and this can stabilize coexistence of two species at high but incomplete levels of assortative mating. Given these results and evidence that it may take many millions of years of divergence before related species become sympatric, we postulate that coexistence of closely related species is more often limited by insufficient assortative mating than by insufficient ecological differentiation. [ABSTRACT FROM AUTHOR]

Disassortative mating is a rare form of mate preference that promotes the persistence of polymorphism. While the evolution of assortative mating and its consequences for trait variation and speciation have been extensively studied, the conditions enabling the evolution of disassortative mating are still poorly understood. Mate preferences increase the risk of missing mating opportunities, a cost that can be compensated by a greater fitness of offspring. Heterozygote advantage should therefore promote the evolution of disassortative mating, which maximizes the number of heterozygous offspring. From the analysis of a two-locus diploid model with one locus controlling the mating cue under viability selection and the other locus coding for the level of disassortative preference, we show that heterozygote advantage and negative frequency-dependent viability selection acting at the cue locus promote the evolution of disassortative preferences. We predict conditions of evolution of disassortative mating coherent with selection regimes acting on traits observed in the wild. We also show that disassortative mating generates sexual selection, which disadvantages heterozygotes at the cue locus, limiting the evolution of disassortative preferences. Altogether, our results partially explain why this behavior is rare in natural populations. [ABSTRACT FROM AUTHOR]

Students of speciation debate the role of performance trade-offs across different environments early in speciation. We tested for early performance trade-offs with a host shift experiment using a member of the Enchenopa binotata species complex of treehoppers (Hemiptera: Membracidae). In this clade of plant-feeding insects, different species live on different host plants and exhibit strong behavioral and physiological host specialization. After five generations, the experimental host shifts resulted either in no adaptation or in adaptation without specialization. The latter result was more likely in sympatry; in allopatry, populations on novel host plants were more likely to become extinct. We conclude that in the early stages of speciation, adaptation to novel host plants does not necessarily bring about performance trade-offs on ancestral environments. Adaptation may be facilitated rather than hindered by gene flow, which prevents extinction. Additional causes of specialization and assortative mating may be required if colonization of novel environments is to result in speciation. [ABSTRACT FROM AUTHOR]

Sexually selected traits have long been thought to drive diversification, but support for this hypothesis has been persistently controversial. In fishes, sexually dimorphic coloration is associated with assortative mating and speciation among closely related species, as shown in classic studies. However, it is unclear whether these results can generalize to explain diversity patterns across ray-finned fishes, which contain the majority of vertebrate species and 96% of fishes. Here, we use phylogenetic approaches to test for an association between sexual dichromatism and diversification rates (speciation minus extinction) in ray-finned fishes. We assembled dichromatism data for 10,898 species, a data set of unprecedented size. We found no difference in diversification rates between monochromatic and dichromatic species when including all ray-finned fishes. However, at lower phylogenetic scales (within orders and families), some intermediate-sized clades did show an effect of dichromatism on diversification. Surprisingly, dichromatism could significantly increase or decrease diversification rates. Moreover, we found no effect in many of the clades initially used to link dichromatism to speciation in fishes (e.g., cichlids) or an effect only at shallow scales (within subclades). Overall, we show how the effects of dichromatism on diversification are highly variable in direction and restricted to certain clades and phylogenetic scales. [ABSTRACT FROM AUTHOR]

Phenological traits, such as the timing of reproduction, are often influenced by social interactions between paired individuals. Such partner effects may occur when pair members affect each other's prebreeding environment. Partner effects can be environmentally and/or genetically determined, and quantifying direct and indirect genetic effects is important for understanding the evolutionary dynamics of phenological traits. Here, using 26 years of data from a pedigreed population of a migratory seabird, the common tern (Sterna hirundo), we investigate male and female effects on female laying date. We find that female laying date harbors both genetic and environmental variation and is additionally influenced by the environmental and, to a lesser extent, genetic component of its mate. We demonstrate this partner effect to be largely explained by male arrival date. Interestingly, analyses of mating patterns with respect to arrival date show mating to be strongly assortative, and using simulations we show that assortative mating leads to overestimation of partner effects. Our study provides evidence for partner effects on breeding phenology in a long-distance migrant while uncovering the potential causal pathways underlying the observed effects and raising awareness for confounding effects resulting from assortative mating or other common environmental effects. [ABSTRACT FROM AUTHOR]

Partial prezygotic isolation is often viewed as more important than partial postzygotic isolation (low fitness of hybrids) early in the process of speciation. I simulate secondary contact between two populations (species) to examine effects of assortative mating and low hybrid fitness in preventing blending. A small reduction in hybrid fitness (e.g., by 10%) produces a narrower hybrid zone than a strong but imperfect mating preference (e.g., 10 times stronger preference for conspecific over heterospecific mates). In the latter case, rare F1 hybrids find each other attractive (due to assortative mating), leading to the buildup of a continuum of intermediates. The weakness of assortative mating compared with reduced fitness of hybrids in preventing blending is robust to varying genetic bases of these traits. Assortative mating is most powerful in limiting blending when it is encoded by a single locus or is essentially complete, or when there is a large mate search cost. In these cases assortative mating is likely to cause hybrids to have low fitness, due to frequency-dependent mating disadvantage of individuals of rare mating types. These results prompt a questioning of the concept of partial prezygotic isolation, since it is not very isolating unless there is also postzygotic isolation. [ABSTRACT FROM AUTHOR]

Evolutionary theory predicts that positive assortative mating—the tendency of similar individuals to mate with each other—plays a key role for speciation by generating reproductive isolation between diverging populations. However, comprehensive tests for an effect of assortative mating on species richness at the macroevolutionary scale are lacking. We used a meta-analytic approach to test the hypothesis that the strength of assortative mating within populations is positively related to species richness across a broad range of animal taxa. Specifically, we ran a phylogenetically independent meta-analysis using an extensive database of 1,447 effect sizes for the strength of assortative mating, encompassing 307 species from 130 families and 14 classes. Our results suggest that there is no relationship between the strength of assortative mating and species richness across and within major taxonomic groups and trait categories. Moreover, our analysis confirms an earlier finding that animals typically mate assortatively (global Pearson correlation coefficient: r = 0.36 ; 95% confidence interval: 0.19–0.52) when accounting for phylogenetic nonindependence. We argue that future advances will rely on a better understanding of the evolutionary causes and consequences of the observed intra- and interspecific variation in the strength of assortative mating. [ABSTRACT FROM AUTHOR]

Behavior can strongly influence rates and patterns of hybridization between animal populations and species. Yet few studies have examined reproductive behaviors in natural hybrid zones within the fine-scale social context in which they naturally occur. We use radio-frequency identification tags with social network analyses to test whether phenotypic similarity in plumage and mass correlate with social behavior throughout a breeding season in a California and Gambel's quail hybrid zone. We use a novel approach to partition phenotypic variation in a way that does not confound differences between sexes and species, and we illustrate the complex ways that phenotype and behavior structure the social environment, mating opportunities, and male-male associations. Associations within the admixed population were random with respect to species-specific plumage but showed strong patterns of assortment based on sexually dimorphic plumage, monomorphic plumage, and mass. Weak behavioral reproductive isolation in this admixed population may be the result of complex patterns of phenotypic assortment based on multiple traits rather than a lack of phenotypic discrimination. More generally, our results support the utility of social network analyses for analyzing behavioral factors affecting genetic exchange between populations and species. [ABSTRACT FROM AUTHOR]

Codivergence of mating traits and mate preferences can lead to behavioral isolation among lineages in early stages of speciation. However, mate preferences limit gene flow only when expressed as mate choice, and numerous factors might be more important than preferences in nature. In the extremely color polytypic strawberry poison frog (Oophaga pumilio), female mate preferences have codiverged with color in most allopatric populations tested. Whether these lab-assayed preferences predict mating (gene flow) in the wild remains unclear. We observed courting pairs in a natural contact zone between red and blue lineages until oviposition or courtship termination. We found color-assortative mating in a disturbed habitat with high population density but not in a secondary forest with lower density. Our results suggest color-assortative O. pumilio mate choice in the wild but also mating patterns that do not match those predicted by lab-assayed preferences. [ABSTRACT FROM AUTHOR]

Animal coloration patterns, from zebra stripes to bird egg speckles, are remarkably varied. With research on the perception, function, and evolution of animal patterns growing rapidly, we require a convenient framework for quantifying their diversity, particularly in the contexts of camouflage, mimicry, mate choice, and individual recognition. Ideally, patterns should be defined by their locations in a low-dimensional pattern space that represents their appearance to their natural receivers, much as color is represented by color spaces. This synthesis explores the extent to which animal patterns, like colors, can be described by a few perceptual dimensions in a pattern space. We begin by reviewing biological spatial vision, focusing on early stages during which neurons act as spatial filters or detect simple features such as edges. We show how two methods from computational vision—spatial filtering and feature detection—offer qualitatively distinct measures of animal coloration patterns. Spatial filters provide a measure of the image statistics, captured by the spatial frequency power spectrum. Image statistics give a robust but incomplete representation of the appearance of patterns, whereas feature detectors are essential for sensing and recognizing physical objects, such as distinctive markings and animal bodies. Finally, we discuss how pattern space analyses can lead to new insights into signal design and macroevolution of animal phenotypes. Overall, pattern spaces open up new possibilities for exploring how receiver vision may shape the evolution of animal pattern signals. [ABSTRACT FROM AUTHOR]

Assortative mating in animals can have substantial evolutionary impact. Numerous reports also make it appear to be pervasive in occurrence. In assortative mating, defined here in behavioral terms, animals select their mates according to a particular shared trait such that mated individuals phenotypically resemble each other more than expected by chance. Body size is a widely studied assortment trait. This is especially relevant for anuran amphibians (frogs and toads), among whom reproductive advantages may accrue to large individuals of both sexes. Anurans also exhibit discrete forms of male mating behavior. Sedentary calling behavior of "sitters" allows for female choice, whereas fighting for possession of mates by "scramblers" precludes female choice. Size-assortative mating in anurans should therefore be a property of sitters, not scramblers. I used meta-analysis to assess the occurrence of true size-assortative mating in relation to mating behavior and other variables in 282 studies of 68 species of anurans. I found publication bias against reporting nonsignificant results and analytical bias resulting from pooling of samples collected at different times or places (Simpson's paradox). Pooled samples significantly inflated the apparent occurrence and strength of size-assortative mating. Controlling for such biases left little credible evidence for size-assortative mating behavior in any anurans. Instead, large-male advantage among scramblers was associated with a secondary pattern of concomitant nonrandom mating. In this disproportionate mating, neither sex behaves according to mate choice rules that could lead to consistently strong assortment. It should thus have relatively little evolutionary impact compared to true assortative mating. [ABSTRACT FROM AUTHOR]

The effect of learned culture (e.g., birdsong dialects and human languages) on genetic divergence is unclear. Previous theoretical research suggests that because oblique learning allows phenotype transmission from individuals with no offspring to an unrelated individual in the next generation, the effect of sexual selection on the learned trait is masked. However, I propose that migration and spatially constrained learning can form statistical associations between cultural and genetic traits, which may allow selection on the cultural traits to indirectly affect the genetic traits. Here, I build a population genetic model that allows such statistical associations to form and find that sexual selection and divergent selection on the cultural trait can indeed help maintain genetic divergence through such statistical associations, while selection against genetic hybrids does not affect cultural trait divergence. Furthermore, I find that even when the cultural trait changes over time due to drift and mutation, it can still help maintain genetic divergence. These results suggest the role of obliquely transmitted traits in evolution may be underrated, and the lack of one-to-one associations between cultural and genetic traits may not be sufficient to disprove the role of culture in genetic divergence. [ABSTRACT FROM AUTHOR]

There is growing evidence that speciation can occur between populations that are not geographically isolated. The emergence of assortative mating is believed to be critical to this process, but how assortative mating arises in diverging populations is poorly understood. The wolf spider genus Schizocosa has become a model system for studying mechanisms of assortative mating. We conducted a series of experiments to identify the factors that control mate pair formation in a Schizocosa population that includes both ornamented and nonornamented males. We show that the population also includes two previously unrecognized female phenotypes. One female phenotype mates mostly or exclusively with ornamented males, and the other mates mostly or exclusively with unornamented males. Assortative mating within these groups is maintained by differences in maturation time, microhabitat use, and female mate preference. We conclude that the population is not a single species, as previously believed, but rather an incipient species pair with multiple overlapping mechanisms of reproductive isolation. The identification of a new incipient species pair in the well-studied and rapidly speciating Schizocosa clade presents new opportunities for the study of speciation without geographic isolation. [ABSTRACT FROM AUTHOR]

The large body of theory on speciation with gene flow has brought to light fundamental differences in the effects of two types of mating rules on speciation: preference/trait rules, in which divergence in both (female) preferences and (male) mating traits is necessary for assortment, and matching rules, in which individuals mate with like individuals on the basis of the presence of traits or alleles that they have in common. These rules can emerge from a variety of behavioral or other mechanisms in ways that are not always obvious. We discuss the theoretical properties of both types of rules and explain why speciation is generally thought to be more likely under matching rather than preference/trait rules. We furthermore discuss whether specific assortative mating mechanisms fall under a preference/trait or matching rule, present empirical evidence for these mechanisms, and propose empirical tests that could distinguish between them. The synthesis of the theoretical literature on these assortative mating rules with empirical studies of the mechanisms by which they act can provide important insights into the occurrence of speciation with gene flow. Finally, by providing a clear framework we hope to inspire greater alignment in the ways that both theoreticians and empiricists study mating rules and how these rules affect speciation through maintaining or eroding barriers to gene flow among closely related species or populations. [ABSTRACT FROM AUTHOR]

In recent years, theoretical models have introduced the concept that ongoing hybridization between “good” species can occur because incomplete reproductive isolation can be a selected optimum. They furthermore show that positive frequency-dependent sexual selection, which is naturally generated by some of the underlying processes that lead to assortative mating, plays a key role in the evolution of incomplete reproductive isolation. This occurs, however, through different mechanisms in sympatric versus allopatric scenarios. We investigate the evolution of incomplete reproductive isolation by sexual selection in scenarios ranging from sympatry to allopatry, to examine how these mechanisms interact. We consider an ecological scenario in which there are two habitats used during foraging and individuals can breed either within a habitat or in a common mating pool. We find that when trait divergence is maintained, sexual selection drives the evolution of choosiness in opposite ways in the common mating pool versus within each habitat. Specifically, strong choosiness is favored in the common mating pool, whereas intermediate choosiness is favored within habitat; the interaction of these forces determines whether intermediate reproductive isolation ultimately evolves in the system. We further find cases where the evolution of stronger choosiness occurs but leads to the loss of divergence. Overall, our study shows that contrasting forces on the evolution of reproductive isolation can occur in different mating areas, and we propose a new avenue for understanding the diversity in levels of reproductive isolation within and across species. [ABSTRACT FROM AUTHOR]

Phenotypic plasticity is the ability of one genotype to produce different phenotypes depending on environmental conditions. Several conceptual models emphasize the role of plasticity in promoting reproductive isolation and, ultimately, speciation in populations that forage on two or more resources. These models predict that plasticity plays a critical role in the early stages of speciation, prior to genetic divergence, by facilitating fast phenotypic divergence. The ability to plastically express alternative phenotypes may, however, interfere with the early phase of the formation of reproductive barriers, especially in the absence of geographic barriers. Here, we quantitatively investigate mechanisms under which plasticity can influence progress toward adaptive genetic diversification and ecological speciation. We use a stochastic, individual-based model of a predator-prey system incorporating sexual reproduction and mate choice in the predator. Our results show that evolving plasticity promotes the evolution of reproductive isolation under diversifying environments when individuals are able to correctly select a more profitable habitat with respect to their phenotypes (i.e., adaptive habitat choice) and to assortatively mate with relatively similar phenotypes. On the other hand, plasticity facilitates the evolution of plastic generalists when individuals have a limited capacity for adaptive habitat choice. We conclude that plasticity can accelerate the evolution of a reproductive barrier toward adaptive diversification and ecological speciation through enhanced phenotypic differentiation between diverging phenotypes. [ABSTRACT FROM AUTHOR]

Adaptive speciation can occur when a population undergoes assortative mating and disruptive selection caused by frequency- dependent intraspecific competition. However, other interactions, such as mutualisms based on trait matching, may generate conflicting selective pressures that constrain species diversification. We used individual-based simulations to explore how different types of mutualism affect adaptive diversification. A magic trait was assumed to simultaneously mediate mate choice, intraspecific competition, and mutualisms. In scenarios of intimate, specialized mutualisms, individuals interact with one or few individual mutualistic partners, and diversification is constrained only if the mutualism is obligate. In other scenarios, increasing numbers of different partners per individual limit diversification by generating stabilizing selection. Stabilizing selection emerges from the greater likelihood of trait mismatches for rare, extreme phenotypes than for common intermediate phenotypes. Constraints on diversification imposed by increased numbers of partners decrease if the trait matching degree has smaller positive effects on fitness. These results hold after the relaxation of various assumptions. When trait matching matters, mutualism-generated stabilizing selection would thus often constrain diversification in obligate mutualisms, such as ant-myrmecophyte associations, and in low-intimacy mutualisms, including plant-seed disperser systems. Hence, different processes, such as trait convergence favoring the incorporation of nonrelated species, are needed to explain the higher richness of low-intimacy assemblages—shown here to be up to 1 order of magnitude richer than high-intimacy systems. [ABSTRACT FROM AUTHOR]

Coevolution of plants and pollinators has been suggested as a mechanism driving diversification of plant-pollinator mutualisms. There is increasing recognition that predators or competitors can influence the abundance and behavior of pollinators and indirectly affect the fitness of plants. However, existing theories on plantpollinator diversification focus exclusively on mutualistic interactions between plants and pollinators. Here we used simulations to evaluate whether predation on pollinators promotes coevolutionary diversification of plant-pollinator mutualisms.We developed an individualbased simulation model in which the blooming season of plants and the active seasons of pollinators and predators can evolve. In simulations without predators, plant-pollinator coevolution caused diversification in blooming/active seasons for both plants and pollinators, but this diversification resulted in polymorphisms, not speciation. The introduction of predators promoted a split of plant and pollinator populations into reproductively isolated subpopulations with corresponding blooming and active seasons or a directional shift of blooming and active seasons, increasing the possibility of plant-pollinator cospeciation. This result suggests that predation on pollinators can promote sympatric and allopatric divergence of plant-pollinator mutualisms. Joint action of antagonistic and mutualistic interactions may be fundamentally important for diversification in coevolutionary interactions. [ABSTRACT FROM AUTHOR]

Adaptive speciation with gene flow via the evolution of assortative mating has classically been studied in one of two different scenarios. First, speciation can occur if frequency-dependent competition in sympatry induces disruptive selection, leading to indirect selection for mating with similar phenotypes. Second, if a subpopulation is locally adapted to a specific environment, then there is indirect selection against hybridizing with maladapted immigrants. While both of these mechanisms have been modeled many times, the literature lacks models that allow direct comparisons between them. Here we incorporate both frequency-dependent competition and local adaptation into a single model and investigate whether and how they interact in driving speciation. We report two main results. First, we show that individually, the two mechanisms operate under separate conditions, hardly influencing each other when one of them alone is sufficient to drive speciation. Second, we also find that the two mechanisms can operate together, leading to a third speciation mode in which speciation is initiated by selection against maladapted migrants but completed by within-deme competition in a distinct second phase. While this third mode bears some similarity to classical reinforcement, it is considerably faster, and both newly formed species go on to coexist in sympatry. [ABSTRACT FROM AUTHOR]

Many classic models of speciation incorporate assortative mating based on mating groups, such as plants with different flowering times, and they investigate whether an ecological trait under disruptive natural selection becomes genetically associated with the selectively neutral mating trait. It is well known that this genetic association is potently destroyed by recombination. In this note, we point out a more fundamental difficulty: if a "knife-edge" symmetry assumption of previous models is violated, then the mating trait is no longer neutral and sexual selection eliminates the polymorphism in the mating locus. This result strengthens the growing consensus that magic traits are the more likely route to nonallopatric speciation. We expand the model assuming also ecological selection on the mating trait and investigate the conditions for natural selection to overcome sexual selection and maintain mating polymorphism; we find that the combination of natural and sexual selection can cause also bistability of allele frequencies. [ABSTRACT FROM AUTHOR]

Mating behavior has profound consequences for two phenomena-individual reproductive success and the maintenance of species boundaries-that contribute to evolutionary processes. Studies of mating behavior in relation to individual reproductive success are common in many species, but studies of mating behavior in relation to genetic variation and species boundaries are less commonly conducted in socially complex species. Here we leveraged extensive observations of a wild yellow baboon (Papio cynocephalus) population that has experienced recent gene flow from a close sister taxon, the anubis baboon (Papio anubis), to examine how admixturerelated genetic background affects mating behavior. We identified novel effects of genetic background on mating patterns, including an advantage accruing to anubis-like males and assortative mating among both yellow-like and anubis-like pairs. These genetic effects acted alongside social dominance rank, inbreeding avoidance, and age to produce highly nonrandom mating patterns. Our results suggest that this population may be undergoing admixture-related evolutionary change, driven in part by nonrandom mating. However, the strength of the genetic effect is mediated by behavioral plasticity and social interactions, emphasizing the strong influence of social context on mating behavior in socially complex species. [ABSTRACT FROM AUTHOR]

Human mate choice is central to individuals' lives and to the evolution of the species, but the basis of variation in mate choice is not well understood. Here we looked at a large community-based sample of twins and their partners and parents (N> 20,000 individuals) to test for genetic and family environmental influences on mate choice, while controlling for and not controlling for the effects of assortative mating. Key traits were analyzed, including height, body mass index, age, education, income, personality, social attitudes, and religiosity. This revealed near-zero genetic influences on male and female mate choice over all traits and no significant genetic influences on mate choice for any specific trait. A significant family environmental influence was found for the age and income of females' mate choices, possibly reflecting parental influence over mating decisions. We also tested for evidence of sexual imprinting, where individuals acquire mate-choice criteria during development by using their opposite-sex parent as the template of a desirable mate; there was no such effect for any trait. The main discernible pattern of mate choice was assortative mating; we found that partner similarity was due to initial choice rather than convergence and also at least in part to phenotypic matching. [ABSTRACT FROM AUTHOR]

Assortative mating may result from intrinsic individual mating preferences or from assortment traits not requiring expression of preferences. Assortment traits are phenotypes expressed in both sexes that enhance the probability of encountering individuals possessing similar trait values. In the noctuid moth Spodoptera frugiperda, it has been suggested that nonrandom mating between two host strains is caused by a temporal assortment trait-that is, differential timing of calling and copulation during the night. By experimental manipulation of this trait in controlled mate-choice experiments, we investigated whether mating by same-strain individuals is enhanced mainly by the allochronic shift of mating activity or is also affected by time-independent intrinsic mating preferences. The observed patterns suggest that nonrandom mating between the two host strains in the laboratory is shaped by an interaction of both effects that is dominated by mating preferences during the first encounter night. This interaction changes over time as the preferences become weaker on subsequent nights. Males were less restricted than females with regard to both the time shift in mating activity and mate preferences. Although the nature of the mate-preference mechanism remains elusive, its restriction to females suggests that maleproduced close-range pheromones emitted during courtship play a role. [ABSTRACT FROM AUTHOR]

Both males and females vary in fitness. While high-fitness males typically have greater siring success, it is not clear whether these males sire an equal fraction of offspring from all females or a disproportionately large fraction with high-fitness females. The latter nonrandom reproductive pattern can arise as the result of sexual selection and creates a positive correlation in fitness between mates. Such a correlation, if it reflects a positive genetic correlation between mates with respect to fitness, increases the efficiency of selection, reducing mutation load and speeding adaptation. While there is evidence from many taxa that assortative mating for fitness may occur, these studies typically focus on observed matings rather than realized reproductive output. Here, we examine assortative mating for fitness in Drosophila tnelanogaster, first in the context of virgin matings and then using a measure of realized reproduction that incorporates re- mating and postcopulatory processes. We find evidence for positive assortative mating among virgins but no evidence of assortative mating using the more complete measure of reproduction. [ABSTRACT FROM AUTHOR]

The theory of mate choice posits that intensified competition for mates can generate variation in either the strength or the direction of mate preferences within the competing sex. Here, we show that intensified male competition, manipulated through the operational sex ratio, induced differential mate choosiness among threespine stickleback males Gasterosteus aculeatus. In the absence of male competition, males were choosy independent of their condition when presented sequentially with a large and a small female. However, in a male-biased social setting, males in poor condition became indiscriminate, whereas good-condition males continued to be selective. Hence, competition induced condition-dependent mate choosiness. This was probably due to mating opportunities decreasing more for poor-condition than for good-condition males when competition intensified, resulting in condition-dependent cost of choice. Variation in condition and cost of choice could thus allow the persistence of male mate choosiness in populations experiencing intense male competition. [ABSTRACT FROM AUTHOR]

Male mate selection during polygyny traditionally has been eclipsed in the literature by its female counterpart. Existing models that have studied male mate choice have concluded that males with genetically inherited preferences for females exhibiting particular traits are often less fit than males without such a preference, leading to preference loss. In this article, we explore the consequences of a fundamental difference between male and female mate choice, the way in which the opposite sex acts as a resource during mating. By incorporating a strategic process at the ecological level, we show that if males are allowed to actively adjust the distribution of their courtship efforts over the available classes of females, male preference can be maintained as a polymorphism. Further, the resulting coexistence induces a reproductive segregation within the population that, when coupled with genetic control of female traits, can lead to strong linkage disequilibrium between the alleles for trait and preference. These processes can cause complete assortative mating to emerge in the model. [ABSTRACT FROM AUTHOR]

Speciation with gene flow may be driven by a combination of positive assortative mating and disruptive selection, particularly if selection and assortative mating act on the same trait, eliminating recombination between ecotype and mating type. Phenotypically unimodal populations of threespine stickleback (Gasterosteus aculeatus) are commonly subject to disruptive selection due to competition for alternate prey. Here we present evidence that stickleback also exhibit assortative mating by diet. Among-individual diet variation leads to variation in stable isotopes, which reflect prey use. We find a significant correlation between the isotopes of males and eggs within their nests. Because egg isotopes are derived from females, this correlation reflects assortative mating between males and females by diet. In concert with disruptive selection, this assortative mating should facilitate divergence. However, the stickleback population remains phenotypically unimodal, highlighting the fact that assortative mating and disruptive selection do not guarantee evolutionary divergence and speciation. [ABSTRACT FROM AUTHOR]

Many animals and plants show a correlation between the traits of the individuals in the mating pair, implying assortative mating. Given the ubiquity of assortative mating in nature, why and how it has evolved remain open questions. Here we attempt to answer these questions in those cases where the trait under assortment is the same in males and females. We consider the most favorable scenario for assortment to evolve, where the same trait is under assortment and viability selection. We find conditions for assortment to evolve using a multilocus formalism in a haploid population. Our results show how epistasis in fitness between the loci that control the focal trait is crucial for assortment to evolve. We then assume specific forms of assortment in haploids and diploids and study the limiting cases of selective and nonselective mating. We find that selection for increased assortment is weak and that where increased assortment is costly, it does not invade. [ABSTRACT FROM AUTHOR]

Several recent models have shown that frequency-dependent disruptive selection created by intraspecific competition can lead to the evolution of assortative mating and, thus, to competitive sympatric speciation. However, since most of these results rely on limited numerical analyses, their generality has been debated. Here,we consider one of the standard models (the so-called Roughgarden model) with a simplified genetics where the selected trait is determined by a single diallelic locus. This model is sufficiently complex to maintain key properties of the general multilocus case but simple enough to allow for comprehensive analytical treatment by means of invasion fitness arguments. Depending on (1) the strength and (2) the shape of stabilizing selection, (3) the strength and (4) the shape of pairwise competition, (5) the shape of the mating function, and (6) whether assortative mating leads to sexual selection, we find five different evolutionary regimes. In one of these regimes, complete reproductive isolation can evolve through arbitrarily small steps in the strength of assortative mating. Our approach provides a mechanistic understanding of several phenomena that have been found in previous models. The results demonstrate how even in a simple model, the evolutionary outcome depends in a complex way on ecological and genetic parameters. [ABSTRACT FROM AUTHOR]

Ecologically driven sympatric speciation has received much attention recently. We investigate a multilocus model of a quantitative trait that is under frequency-dependent selection caused by intraspecific competition and acts as mating character for assortment. We identify the conditions that lead to the establishment of reproductively isolated clusters. This may be interpreted as evolutionary splitting or sympatric speciation. In our model, there are parameters that independently determine the strength of assortment, the costs for being choosy, and the strength of frequency-dependent natural selection. Sufficiently strong frequency dependence leads to disruptive selection on the phenotypes. The population consists of (sexual) haploid individuals. If frequency dependence is strong enough to induce disruptive selection and costs are absent or low, the result of evolution depends in a distinctive nonlinear way on the strength of assortment: under moderately strong assortment, less genetic variation is maintained than under weak or strong assortment, and sometimes there is none at all. Evolutionary splitting occurs only if frequency dependence and assortment are both strong enough and costs are low. Even then, the evolutionary outcome depends on the genetics and the initial conditions. The roles of the number of loci, of linkage, and of asymmetric selection are also explored. [ABSTRACT FROM AUTHOR]

The reinforcement model of speciation holds an important place in modern evolutionary thought. A closer examination of the model using a computer simulation reveals that there are a number of implicit assumptions necessary for reinforcement to work. 1. It requires that reproductive characters form part of an open genetic system with high heritability and that there is considerable variability within this system. 2. The two strains must coexist throughout the period in which selection is acting. This requires intense frequency-dependent selection to counteract the inherently unstable selection regime of negative heterosis. The necessary intensity suggests that extinction of one of the strains is the more likely outcome unless there are substantial initial differences in reproductive characters. Furthermore, chance extinction of one of the strains cannot occur in this selection period. 3. The net cost of assortative characters must be less than the cost of a greater probability of producing hybrid offspring. This is a complex matter with several opposing factors. 4. The fitness of the hybrids must be zero or close to it and must remain so. 5. The selective pressure for assortative reproductive characters becomes weaker as these characters are established. It appears unlikely that selection alone could complete the process, as is hypothesized in some formulations of the model. Further data from natural populations and laboratory experiments are required to clarify several of these points, especially the first. These problems do not mean that reinforcement can never occur, but they do imply that speciation by reinforcement is not nearly as prevalent as much of the biological literature suggests. [ABSTRACT FROM AUTHOR]

In central Europe, the hybridogenetic waterfrog Rana esculenta, a hybrid between Rana ridibunda and Rana lessonae, lives in sympatry with one of its parental species, the poolfrog Rana lessonae. As R. esculenta has to backcross constantly with R. lessonae in order to produce viable offspring, this coexistence is obligatory for R. esculenta. Since R. esculenta has a higher primary fitness than R. lessonae, a mechanism is required that prevents the hybrid from driving the parental species, and hence itself, to extinction. Here, we present an analytical model and a computer simulation that investigate whether assortative mating can operate as a such a control mechanism. Our results show that assortative mating is very effective in regulating coexistence in such a hybrid-host system. This is particularly true when choice is affected by the proportion of the two male types in the population. Furthermore, we could show that even if the species composition in a mixed hybrid-host population may be largely influenced by differences in life-history parameters, assortative mating still plays a very important role by stabilizing coexistence. Thus, mating behavior turns out to be more important for the populations dynamics of hybridogenetic waterfrog systems than previously assumed.