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Multi-type birth–death processes underlie approaches for inferring evolutionary dynamics from phylogenetic trees across biological scales, ranging from deep-time species macroevolution to rapid viral evolution and somatic cellular proliferation. A limitation of current phylogenetic birth–death models is that they require restrictive linearity assumptions that yield tractable message-passing likelihoods, but that also preclude interactions between individuals. Many fundamental evolutionary processes – such as environmental carrying capacity or frequency-dependent selection – entail interactions, and may strongly influence the dynamics in some systems. Here, we introduce a multi-type birth–death process in mean-field interaction with an ensemble of replicas of the focal process. We prove that, under quite general conditions, the ensemble's stochastically evolving interaction field converges to a deterministic trajectory in the limit of an infinite ensemble. In this limit, the replicas effectively decouple, and self-consistent interactions appear as nonlinearities in the infinitesimal generator of the focal process. We investigate a special case that is rich enough to model both carrying capacity and frequency-dependent selection while yielding tractable message-passing likelihoods in the context of a phylogenetic birth–death model. [ABSTRACT FROM AUTHOR]

Explaining the maintenance of genetic variation in fitness‐related traits within populations is a fundamental challenge in ecology and evolutionary biology. Frequency‐dependent selection (FDS) is one mechanism that can maintain such variation, especially when selection favours rare variants (negative FDS). However, our general knowledge about the occurrence of FDS, its strength and direction remain fragmented, limiting general inferences about this important evolutionary process. We systematically reviewed the published literature on FDS and assembled a database of 747 effect sizes from 101 studies to analyse the occurrence, strength, and direction of FDS, and the factors that could explain heterogeneity in FDS. Using a meta‐analysis, we found that overall, FDS is more commonly negative, although not significantly when accounting for phylogeny. An analysis of absolute values of effect sizes, however, revealed the widespread occurrence of modest FDS. However, negative FDS was only significant in laboratory experiments and non‐significant in mesocosms and field‐based studies. Moreover, negative FDS was stronger in studies measuring fecundity and involving resource competition over studies using other fitness components or focused on other ecological interactions. Our study unveils key general patterns of FDS and points in future promising research directions that can help us understand a long‐standing fundamental problem in evolutionary biology and its consequences for demography and ecological dynamics. [ABSTRACT FROM AUTHOR]

Seed predation by rodents relies on the abundance of available seeds and seed traits like sizes and tannin-content. Here, by tracking the fates of 2400 tagged seeds of two sympatric tree species Quercus wutaishanica and Corylus heterophylla with varying size and tannin content, we assessed rodents’ preference for specific seed traits. Meanwhile, we set the ratio of large to small seeds within species as 9:1, 7:3, 5:5, 3:7, and 1:9, respectively, to examine rodent behavior in response to seed relative frequencies. We found that: 1) rodents preferentially hoarded C. heterophylla seeds with lower-tannins after dispersal while Q. wutaishanica seeds with higher-tannins are more often eaten in situ or after dispersal. Peeling seed coat is an adaptive strategy for rodents to avoid high tannins, particularly for Q. wutaishanica. 2) Rodents prioritize small seeds over large ones for Q. wutaishanica whereas significant differences were not observed for C. heterophylla. Rodents commonly eat small seeds after successful dispersal and prefer to hoard larger ones over long distances. 3) Positive frequency-dependent selection was observed for both seed types. Our results suggest that the selection of seeds by rodents is determined by seed tannins and sizes, and changes in seed size frequencies could alter the interaction between trees and rodents from predation towards mutualism.

Mathematical models of genetic evolution often come in pairs, connected by a so-called duality relation. The most seminal example are the Wright–Fisher diffusion and the Kingman coalescent, where the former describes the stochastic evolution of neutral allele frequencies in a large population forwards in time, and the latter describes the genetic ancestry of randomly sampled individuals from the population backwards in time. As well as providing a richer description than either model in isolation, duality often yields equations satisfied by quantities of interest. We employ the so-called Bernoulli factory – a celebrated tool in simulation-based computing – to derive duality relations for broad classes of genetics models. As concrete examples, we present Wright–Fisher diffusions with general drift functions, and Allen–Cahn equations with general, nonlinear forcing terms. The drift and forcing functions can be interpreted as the action of frequency-dependent selection. To our knowledge, this work is the first time a connection has been drawn between Bernoulli factories and duality in models of population genetics. • Bernoulli factories can be used to construct general models of population genetics. • These constructions yield duality relations which can facilitate calculations. • Our approach covers broad classes of Wright–Fisher and Allen–Cahn models. [ABSTRACT FROM AUTHOR]

Traditional norms of human societies in rural China may have changed owing to population expansion, rapid development of the tourism economy and globalization since the 1990s; people from different ethnic groups might adopt cultural traits from outside their group or lose their own culture at different rates. Human behavioural ecology can help to explain adoption of outgroup cultural values. We compared the adoption of four cultural values, specifically speaking outgroup languages/mother tongue and wearing jeans, in two co-residing ethnic groups, the Mosuo and Han. Both groups are learning outgroup traits, including each other's languages through contact in economic activities, education and kin networks, but only the Mosuo are starting to lose their own language. Males are more likely to adopt outgroup values than females in both groups. Females of the two groups are no different in speaking Mandarin and wearing jeans, whereas males do differ, with Mosuo males being keener to adopt them than Han males. The reason might be that Mosuo men experience more reproductive competition over mates, as Mosuo men have larger reproductive skew than others. Moreover, Mosuo men but not others gain fitness benefits from the adoption of Mandarin (they start reproducing earlier than non-speakers). This article is part of the theme issue 'Social norm change: drivers and consequences'. [ABSTRACT FROM AUTHOR]

The persistence of non-neutral trait polymorphism is enigmatic because stabilizing selection is expected to deplete variation. In cryptically coloured prey, negative frequency-dependent selection due to search image formation by predators has been proposed to favour rare variants, promoting polymorphism. However, in a heterogeneous environment, locally varying disruptive selection favours patch type-specific optima, resulting in spatial segregation of colour variants. Here, we address whether negative frequency-dependent selection can overcome selection posed by habitat heterogeneity to promote local polymorphism using an individual-based model. In addition, we compare how prey and predator mobility may modify the outcome. Our model revealed that frequency-dependent predation could strongly promote local prey polymorphism, but only when differences between morphs in patch-specific fitness were small. The effect of frequency-dependent predation depended on the predator adjustment of search image and was hampered by the prey population structure. Gene flow due to prey movement counteracted local selection, promoted local polymorphism to some extent, and relaxed the conditions for polymorphism due to frequency-dependent predation. Importantly, abrupt spatial changes in morph frequencies decreased the probability that mobile frequency-dependent predators could maintain local prey polymorphism. Overall, our study suggests that in a spatially heterogeneous environment, negative frequency-dependent selection may help maintain local polymorphism but only under a limited range of conditions. [ABSTRACT FROM AUTHOR]

While the role of selection in divergence along the speciation continuum is theoretically well understood, defining specific signatures of selection in the genomic landscape of divergence is empirically challenging. Modelling approaches can provide insight into the potential role of selection on the emergence of a heterogenous genomic landscape of divergence. Here, we extend and apply an individual‐based approach that simulates the phenotypic and genotypic distributions of two populations under a variety of selection regimes, genotype–phenotype maps, modes of migration, and genotype‐environment interactions. We show that genomic islands of high differentiation and genomic valleys of similarity may respectively form under divergent and parallel selection between populations. For both types of between‐population selection, negative and positive frequency‐dependent selection within populations generated genomic islands of higher magnitude and genomic valleys of similarity, respectively. Divergence rates decreased under strong dominance with divergent selection, as well as in models including genotype‐environment interactions under parallel selection. For both divergent and parallel selection models, divergence rate was higher under an intermittent migration regime between populations, in contrast to a constant level of migration across generations, despite an equal number of total migrants. We highlight that interpreting a particular evolutionary history from an observed genomic pattern must be done cautiously, as similar patterns may be obtained from different combinations of evolutionary processes. Modelling approaches such as ours provide an opportunity to narrow the potential routes that generate the genomic patterns of specific evolutionary histories. [ABSTRACT FROM AUTHOR]

In patch- or habitat-structured populations, different processes can favor adaptive polymorphism at different scales. While spatial heterogeneity can generate spatially disruptive selection favoring variation between patches, local competition can lead to locally disruptive selection promoting variation within patches. So far, almost all theory has studied these two processes in isolation. Here, we use mathematical modeling to investigate how resource variation within and between habitats influences the evolution of variation in a consumer population where individuals compete in finite patches connected by dispersal. We find that locally and spatially disruptive selection typically act in concert, favoring polymorphism under a wider range of conditions than when in isolation. But when patches are small and dispersal between them is low, kin competition inhibits the emergence of polymorphism, especially when the latter is driven by local competition for resources. We further use our model to clarify what comparisons between trait and neutral genetic differentiation ( Q ST / F ST comparisons) can tell about the nature of selection. Overall, our results help us understand the interaction between two major drivers of polymorphism: locally and spatially disruptive selection, and how this interaction is modulated by the unavoidable effects of kin selection under limited dispersal. [ABSTRACT FROM AUTHOR]

Competitive fitness assays are widely used in evolutionary biology and typically rely on a reference strain to compare different focal genotypes. This approach implicitly relies on the absence of interaction between the competing genotypes. In other words, the performance of the reference strain must not depend on the competitor. This report scrutinized this assumption by competing diverged Drosophila simulans populations against a common reference strain. We detected strong evidence for interaction between the competing genotypes: (1) Frequency‐dependent selection was common with opposite effects in genetically diverged populations. (2) Temporal heterogeneity of fitness estimates, which can be partially attributed to a competitor‐specific delay in the eclosion of the reference strain. We propose that this inconsistent behavior of the reference strain can be considered a specific case of a genotype × environment interaction. Focal populations could modify the environment of the reference strain, either indirectly by altering the microbiome composition and food availability or directly by genotype‐specific cannibalism. Our results provide new insights into the interaction of diverged genotypes and have important implications for the interpretation of competitive fitness assays. [ABSTRACT FROM AUTHOR]

Plant viruses account for enormous agricultural losses worldwide, and the most effective way to combat them is to identify genetic material conferring plant resistance to these pathogens. Aiming to identify genetic associations with responses to infection, we screened a large panel of Arabidopsis thaliana natural inbred lines for four disease-related traits caused by infection by A. thaliana-naïve and -adapted isolates of the natural pathogen turnip mosaic virus (TuMV). We detected a strong, replicable association in a 1.5 Mb region on chromosome 2 with a 10-fold increase in relative risk of systemic necrosis. The region contains several plausible causal genes as well as abundant structural variation, including an insertion of a Copia transposon into a Toll/interleukin receptor (TIR-NBS-LRR) coding for a gene involved in defense, that could be either a driver or a consequence of the disease-resistance locus. When inoculated with TuMV, loss-of-function mutant plants of this gene exhibited different symptoms than wild-type plants. The direction and severity of symptom differences depended on the adaptation history of the virus. This increase in symptom severity was specific for infections with the adapted isolate. Necrosis-associated alleles are found worldwide, and their distribution is consistent with a trade-off between resistance during viral outbreaks and a cost of resistance otherwise, leading to negative frequency-dependent selection.

Abstract Competitive fitness assays are widely used in evolutionary biology and typically rely on a reference strain to compare different focal genotypes. This approach implicitly relies on the absence of interaction between the competing genotypes. In other words, the performance of the reference strain must not depend on the competitor. This report scrutinized this assumption by competing diverged Drosophila simulans populations against a common reference strain. We detected strong evidence for interaction between the competing genotypes: (1) Frequency‐dependent selection was common with opposite effects in genetically diverged populations. (2) Temporal heterogeneity of fitness estimates, which can be partially attributed to a competitor‐specific delay in the eclosion of the reference strain. We propose that this inconsistent behavior of the reference strain can be considered a specific case of a genotype × environment interaction. Focal populations could modify the environment of the reference strain, either indirectly by altering the microbiome composition and food availability or directly by genotype‐specific cannibalism. Our results provide new insights into the interaction of diverged genotypes and have important implications for the interpretation of competitive fitness assays.

Abstract Marine gastropods are characterized by an incredible variation in shell color. In this review, we aim to introduce researchers to previous studies of shell color polymorphism in this group of animals, trying to provide an overview of the topic and highlighting some potential avenues for future research. For this, we tackle the different aspects of shell color polymorphism in marine gastropods: its biochemical and genetic basis, its patterns of spatial and temporal distribution, as well as its potential evolutionary causes. In particular, we put special emphasis on the evolutionary studies that have been conducted so far to reveal the evolutionary mechanisms responsible for the maintenance of shell color polymorphism in this group of animals, as it constitutes the least addressed aspect in existing literature reviews. Several general conclusions can be drawn from our review: First, natural selection is commonly involved in the maintenance of gastropod color polymorphism; second, although the contribution of neutral forces (gene flow‐genetic drift equilibrium) to shell color polymorphism maintenance do not seem to be particularly important, it has rarely been studied systematically; third, a relationship between shell color polymorphism and mode of larval development (related to dispersal capability) may exist. As for future studies, we suggest that a combination of both classical laboratory crossing experiments and ‐Omics approaches may yield interesting results on the molecular basis of color polymorphism. We believe that understanding the various causes of shell color polymorphism in marine gastropods is of great importance not only to understand how biodiversity works, but also for protecting such biodiversity, as knowledge of its evolutionary causes may help implement conservation measures in those species or ecosystems that are threatened.

We study ancestral structures for the two-type Moran model with mutation and frequency-dependent selection under the nonlinear dominance or fittest-type-wins scheme. Under appropriate conditions, both lead, in distribution, to the same type-frequency process. Reasoning through the mutations on the ancestral selection graph (ASG), we develop the corresponding killed and pruned lookdown ASGs and use them to determine the present and ancestral type distributions. To this end, we establish factorial moment dualities to the Moran model and a relative. We extend the results to the diffusion limit and present applications for finite populations, as well as for moderate and weak selection limits. [ABSTRACT FROM AUTHOR]

Marine gastropods are characterized by an incredible variation in shell color. In this review, we aim to introduce researchers to previous studies of shell color polymorphism in this group of animals, trying to provide an overview of the topic and highlighting some potential avenues for future research. For this, we tackle the different aspects of shell color polymorphism in marine gastropods: its biochemical and genetic basis, its patterns of spatial and temporal distribution, as well as its potential evolutionary causes. In particular, we put special emphasis on the evolutionary studies that have been conducted so far to reveal the evolutionary mechanisms responsible for the maintenance of shell color polymorphism in this group of animals, as it constitutes the least addressed aspect in existing literature reviews. Several general conclusions can be drawn from our review: First, natural selection is commonly involved in the maintenance of gastropod color polymorphism; second, although the contribution of neutral forces (gene flow‐genetic drift equilibrium) to shell color polymorphism maintenance do not seem to be particularly important, it has rarely been studied systematically; third, a relationship between shell color polymorphism and mode of larval development (related to dispersal capability) may exist. As for future studies, we suggest that a combination of both classical laboratory crossing experiments and ‐Omics approaches may yield interesting results on the molecular basis of color polymorphism. We believe that understanding the various causes of shell color polymorphism in marine gastropods is of great importance not only to understand how biodiversity works, but also for protecting such biodiversity, as knowledge of its evolutionary causes may help implement conservation measures in those species or ecosystems that are threatened. [ABSTRACT FROM AUTHOR]

ABSTRACT Viruses infecting marine prokaryotes have a large impact on the diversity and dynamics of their hosts. Model systems suggest that viral infection is frequency dependent and constrained by the virus-host encounter rate. However, it is unclear whether frequency-dependent infection is pervasive among the abundant prokaryotic populations with different temporal dynamics. To address this question, we performed a comparison of prokaryotic and viral communities using 16S rRNA amplicon and virome sequencing based on samples collected monthly for 2 years at a Japanese coastal site, Osaka Bay. Concurrent seasonal shifts observed in prokaryotic and viral community dynamics indicated that the abundance of viruses correlated with that of their predicted host phyla (or classes). Cooccurrence network analysis between abundant prokaryotes and viruses revealed 6,423 cooccurring pairs, suggesting a tight coupling of host and viral abundances and their “one-to-many” correspondence. Although stable dominant species, such as SAR11, showed few cooccurring viruses, a fast succession of their viruses suggests that viruses infecting these populations changed continuously. Our results suggest that frequency-dependent viral infection prevails in coastal marine prokaryotes regardless of host taxa and temporal dynamics. IMPORTANCE There is little room for doubt that viral infection is prevalent among abundant marine prokaryotes regardless of their taxa or growth strategy. However, comprehensive evaluations of viral infections in natural prokaryotic communities are still technically difficult. In this study, we examined viral infection in abundant prokaryotes by monitoring the monthly dynamics of prokaryotic and viral communities at a eutrophic coastal site, Osaka Bay. We compared the community dynamics of viruses with those of their putative hosts based on genome-based in silico host prediction. We observed frequent cooccurrence among the predicted virus-host pairs, suggesting that viral infection is prevalent in abundant prokaryotes regardless of their taxa or temporal dynamics. This likely indicates that frequent lysis of the abundant prokaryotes via viral infection has a considerable contribution to the biogeochemical cycling and maintenance of prokaryotic community diversity.

MHC genes are extraordinarily polymorphic in most taxa. Host‐pathogen coevolution driven by negative frequency‐dependent selection (NFDS) is one of the main hypotheses for the maintenance of such immunogenetic variation. Here, we test a critical but rarely tested assumption of this hypothesis—that MHC alleles affect resistance/susceptibility to a pathogen in a strain‐specific way, that is, there is a host genotype‐by‐pathogen genotype interaction. In a field study of bank voles naturally infected with the tick‐transmitted bacterium Borrelia afzelii, we tested for MHC class II (DQB) genotype‐by‐B. afzelii strain interactions for infection prevalence between 10 DQB alleles and seven strains. One allele (DQB*37) showed an interaction, such that voles carrying DQB*37 had higher prevalence of two strains and lower prevalence of one strain than individuals without the allele. These findings were corroborated by analyses of strain composition of infections, which revealed an effect of DQB*37 in the form of lower β diversity among infections in voles carrying the allele. Taken together, these results provide rare support at the molecular genetic level for a key assumption of models of antagonistic coevolution through NFDS. [ABSTRACT FROM AUTHOR]

In the target article, Christie, Brusse, et al . argue that selected effect functions do not, in general, explain why a trait exists in a population and, therefore, theories of representational content should not rely on selected effect functions. This response focuses on the claim about functions-for-representation. The role of evolutionary functions in a theory of content is to pick out outcomes that have been systematically stabilized by natural selection. Correctness conditions are conditions involved in explaining how that happened. Selected effect functions can play that role in the complex equilibria that Christie, Brusse, et al . identify. Non-equilibrium cases are also discussed., Competing Interests: No potential conflict of interest was reported by the author(s)., (© 2022 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.)

Evolutionary biology aims to explain the diversity seen in nature. Evolutionary theory provides frameworks to understand how simple polymorphisms or continuous variation are maintained, but phenotypes inherited as discrete suites of quantitative traits are difficult to fit into this framework. Supergenes have been proposed as a solution to this problem-if causal genes are co-located, they can be inherited as if a single gene, thus bridging the gap between simple polymorphisms and continuous traits. We develop models to ask: how are critical supergenes for maintaining phenotypic diversity? In our simplest model, without explicit genetic architectures, three alternative reproductive morphs are maintained in many of the parameter combinations we evaluated. For these same parameter values, models with demographic stochasticity, recombination and mutation (but without explicit genetic architecture) maintained only two of these three morphs, with stochasticity determining which morphs persisted. With explicit genetic architectures, regardless of whether causal loci were co-located in a supergene or distributed randomly, this stochasticity in which morphs are maintained was reduced. Even when phenotypic variation was lost, genetic diversity was maintained. Altogether, categorical traits with polygenic bases exhibited similar evolutionary dynamics to those determined by supergenes. Our work suggests that supergenes are not the only answer to the puzzle of how discrete polygenic phenotypic variation is maintained.

A solitary population of consumers frequently evolves to the middle of a resource gradient and an intermediate mean phenotype compared to a sympatric pair of competing species that diverge to either side via character displacement. The forces governing the distribution of phenotypes in these allopatric populations, however, are little investigated. Theory predicts that the intermediate mean phenotype of the generalist should be maintained by negative frequency‐dependent selection, whereby alternate extreme phenotypes are favored because they experience reduced competition for resources when rare. However, the theory makes assumptions that are not always met, and alternative explanations for an intermediate phenotype are possible. We provide a test of this prediction in a mesocosm experiment using threespine stickleback that are ecologically and phenotypically intermediate between the more specialized stickleback species that occur in pairs. We manipulated the frequency distribution of phenotypes in two treatments and then measured effects on a focal intermediate population. We found a slight frequency‐dependent effect on survival in the predicted direction but not on individual growth rates. This result suggests that frequency‐dependent selection might be a relatively weak force across the range of phenotypes within an intermediate population and we suggest several general reasons why this might be so. We propose that allopatric populations might often be maintained at an intermediate phenotype instead by stabilizing or fluctuating directional selection. [ABSTRACT FROM AUTHOR]

Adaptive radiations are known for rapid niche diversification in response to ecological opportunity. While most resources usually exist prior to adaptive radiation, novel niches associated with novel resources can be created as a clade diversifies. For example, in African lake cichlid radiations some species prey upon other species of the clade (intraclade consumers). Using a trait‐based eco‐evolutionary model, we investigate the evolution of intraclade consumers in adaptive radiations and the effect of this novel trophic interaction on the diversification process of the radiating clade. We find that the evolutionary emergence of intraclade consumers halts the diversification processes of other ecomorphs as a result of increased top‐down control of density. Because high productivity enables earlier evolution of intraclade consumers, highly productive environments come to harbour less species‐rich radiations than comparable radiations in less productive environments. Our results reveal how macroevolutionary and community patterns can emerge from ecological and microevolutionary processes. [ABSTRACT FROM AUTHOR]

A few empirical examples document fixed alternative male mating strategies in animals. Here we focus on the polymorphism of male mating strategies in the ruff (Calidris pugnax, Aves Charadriiformes). In ruffs, three fixed alternative male mating strategies coexist and are signaled by extreme plumage polymorphism. We first present relevant data on the biology of the species. Then we review the available knowledge of the behavioral ecology of ruffs during the breeding season, and we detail the characteristics of each of the three known fixed male mating strategies. We next turn to the results of exceptional quality accumulated on both the structural and functional genomics of the ruff over the past few years. We show how much these genomic data can shed new, mechanistic light on the evolution and maintenance of the three fixed alternative male mating strategies. We then look if there is sufficient indication to support frequency-dependent selection as a key mechanism in maintaining these three strategies. Specifically, we search for evidence of equal fitness among individuals using each of the three strategies. Finally, we propose three lines of research avenues that will help to understand the eco-evolutionary dynamics of phenotypic differences within natural populations of this iconic model species. [ABSTRACT FROM AUTHOR]

Abstract A solitary population of consumers frequently evolves to the middle of a resource gradient and an intermediate mean phenotype compared to a sympatric pair of competing species that diverge to either side via character displacement. The forces governing the distribution of phenotypes in these allopatric populations, however, are little investigated. Theory predicts that the intermediate mean phenotype of the generalist should be maintained by negative frequency‐dependent selection, whereby alternate extreme phenotypes are favored because they experience reduced competition for resources when rare. However, the theory makes assumptions that are not always met, and alternative explanations for an intermediate phenotype are possible. We provide a test of this prediction in a mesocosm experiment using threespine stickleback that are ecologically and phenotypically intermediate between the more specialized stickleback species that occur in pairs. We manipulated the frequency distribution of phenotypes in two treatments and then measured effects on a focal intermediate population. We found a slight frequency‐dependent effect on survival in the predicted direction but not on individual growth rates. This result suggests that frequency‐dependent selection might be a relatively weak force across the range of phenotypes within an intermediate population and we suggest several general reasons why this might be so. We propose that allopatric populations might often be maintained at an intermediate phenotype instead by stabilizing or fluctuating directional selection.

Alternative reproductive tactics, where members of a population exhibit divergent mating approaches, are widespread in fishes. The maintenance of such polymorphisms is often explained using game‐theoretic concepts such as frequency‐dependent selection, which typically assume populations are at equilibrium. However, fish populations exhibit stochastic variability that is inconsistent with the assumption of equilibrium dynamics. We explored how stochastic population dynamics interact with sexual and harvest‐induced selection to shape the frequencies of alternative male phenotypes in Pacific salmon (Oncorhynchus spp., Salmonidae). Males of several salmon species are composed of typical "hooknoses" that compete on the spawning grounds for mating opportunities, and rarer "jacks" that mature younger and at smaller sizes to attempt reproduction by "sneaking." Using stochastic simulations, we found that realistic levels of recruitment variation could increase the prevalence of jacks relative to equilibrium expectations. This result is attributable to the "cohort mismatch" phenomenon, whereby differences in age at maturity between male phenotypes cause jacks from strong recruitments to spawn in the same year as hooknoses from weaker cohorts, elevating the proportion of jacks among spawners. Simulations were supported by data from major populations of sockeye salmon (O. nerka, Salmonidae) throughout their North American range, which indicated a positive association between recruitment variability and jack prevalence in some regions. Moreover, we found that frequency‐dependent mating dampened population responses to fisheries‐induced selection. Our findings emphasize that stochastic ecological processes are important to the maintenance of alternative life histories and interact with selection to produce emergent dynamics that have not previously been recognized. [ABSTRACT FROM AUTHOR]

Although psychopathy is widely conceptualised as a mental disorder, some researchers question the maladaptive nature of psychopathy, and argue that it might be advantageous from an evolutionary point of view. According to this view, psychopathy can be seen as an evolutionary adaptative strategy that relies on deception and manipulation to gain short-term reproductive benefits. Psychopathy is also identified as a fast life strategy in response to early life stress and an adaptation to harsh environments. This paper investigates the evidence that psychopathic traits are adaptive, while also addressing the limitations of current evolutionary models of psychopathy based on frequency-dependent selection and life history theory. We review recent studies on the fitness correlates of psychopathy and find that psychopathic traits present potential adaptive trade-offs between fertility and mortality, and offspring quantity and quality. On a proximate level, individual differences in stress reactivity and environmental risk factors in early development predispose to psychopathy through gene–environment interactions. We propose that environmental, developmental, social and cultural factors can mediate the relationship between psychopathic traits and fitness and therefore should be considered to make accurate predictions on the adaptive potential of psychopathy. We end by outlining gaps in the literature and making recommendations for future evolutionary research on psychopathy.

Hole-nesting tits belonging to the family Paridae produce a hissing display that resembles the exhalatory hiss of a snake. When a predatory animal enters the nest hole of a tit, tits often hiss vigorously, while lunging their head forward and shaking their wings and tail, until the intruder retreats. We assessed the acoustic similarity between such hiss calls from 6 species of tits, snake hisses, and tit syllables used in alarm vocalizations, as well as white noise as a control. Tit hiss calls showed a high degree of similarity with snake hisses from 3 different snake families. Tit hisses had lower similarity to syllable alarm calls, suggesting convergence of tit hisses in their spectral structure. Hiss calls would only be effective in protecting nest boxes if nest predators responded to these calls. In order to test this hypothesis, we trained individual Swinhoe's striped squirrels, Tamiops swinhoei hainanus , a common predator of egg and nestling tits, to feed at feeders in proximity to nest boxes. We compared the aversive response of squirrels to tit's hiss calls and white noise, presented in random order. Squirrels showed a higher degree of avoidance of feeders when hiss calls were played back than when white noise was presented. In conclusion, our study suggests that hole-nesting birds have evolved convergent snake-like hiss calls, and that predators avoid to prey on the contents of nest boxes from which snake-like hisses emerge. [ABSTRACT FROM AUTHOR]

Many species exhibit selective foraging behaviour, where consumers use a nonrandom subset of available food types. Yet little is known about how selective foraging behaviour varies with environmental conditions and the community level consequences of such selection dynamics. We examined selective foraging by wolves preying primarily on elk in Yellowstone National Park (YNP) over a 12-year period and on moose in Isle Royale National Park (IRNP) over a 47-year period. Specifically, we assessed how selection for calves and senescent adults varied with their frequency in the environment, wolf abundance and winter severity. Selection for senescent adults decreased as the relative abundance of senescent prey increased (i.e. negative frequency-dependent selection) in both study sites. In IRNP, selection for calves was also negatively frequency dependent and declined with increasing wolf abundance. These results are inconsistent with the pattern of positive frequency-dependent selection expected under the prey-switching hypothesis. These results suggest that selection is primarily driven by intraspecific differences in prey vulnerability and wolves' interest in minimizing their risk of injury, as opposed to maximizing intake rates. Lastly, we ran simulations to evaluate how predator–prey dynamics were influenced by dynamic patterns of selection, like those observed in YNP and IRNP. The simulations indicated that predators are more efficient (i.e. steeper slope of the numerical response) when selection for calves is negatively frequency dependent, which results in a lower mean abundance of prey. More importantly, predation is a stronger destabilizing force when selection for calves is negatively frequency dependent. That stronger destabilizing force is indicated by greater variability in the abundance of prey and predators, prey populations being less resilient and a steeper negative slope of the relationship between predation rate and prey population growth rate. As such, our simulation analyses suggest that some of the observed patterns of negative-frequency dependent selection may have important consequences for predator–prey dynamics. • Wolves showed negative frequency-dependent selection for calves and senescent prey. • Wolves did not exhibit prey-switching behaviour when preferred prey became rare. • Selection is likely driven by the predator's interest to reduce their risk of injury. • Selection for calves may result in more efficient predators and less resilient prey. [ABSTRACT FROM AUTHOR]

Typically, pistil and pollen specificity determinants are encoded by distinct, but tightly linked and highly polymorphic, genes forming cognate haplotypes at the SI locus (S-locus). This loss will be neutral as long as the pollen remains within the local population, but migrating pollen grains carrying the degenerate S-haplotypes will now be unable to fertilise pistils carrying the original specificity maintained in other subpopulations. However, under collaborative nonself recognition, in addition to these parameters the degree of "completeness", that is the proportion of S-alleles at the pistil gene that can be detoxified by the cumulative action of the products of its pollen-expressed genes present, plays a central role and drastically alters the evolutionary trajectory of the diversification process. The same phenomenon would arise after evolution of a novel S-allele in one subpopulation because immigrant pollen grains would not be able to fertilise pistils carrying this S-allele. [Extracted from the article]

We present a discrete‐time predator–prey model in which the prey population is assumed to evolve in response to a toxicant. We incorporate frequency‐dependent selection into the prey evolution, assuming that an individual's susceptibility to predation depends on both the trait it possesses and the traits of others in the population. When frequency‐dependent selection is symmetric, we show that the trait equation is unable to track changes in the fitness landscape, that is, the fitness landscape may change while the trait continues to evolve to the same trait value. As a result, evolution may produce unfit prey populations. Meanwhile, we show that asymmetric frequency‐dependence may have a destabilizing effect on the system, resulting in a closed invariant curve via a Neimark–Sacker bifurcation. Recommendations for Resource ManagersWhen considering the impact that a toxicant or other disturbance may have on a system, it is important to consider the impact duration of the toxicant and whether any impacted species have the potential to evolve to reduce the impact of the toxicant.Evolution of toxicant resistance may change system dynamics in a number of ways, such as enabling species persistence at higher toxicant levels. However, here we show that frequency‐dependent evolution has the potential to destabilize system dynamics, resulting in stable closed invariant curves.Destabilization of system dynamics may have important consequences for species persistence, as periods of low abundance may increase the possibility of species extinction. [ABSTRACT FROM AUTHOR]

Polymorphic warning signals in aposematic systems are enigmatic because predator learning should favor the most common form, creating positive frequency-dependent survival. However, many populations exhibit variation in warning signals. There are various selective mechanisms that can counter positive frequency-dependent selection and lead to temporal or spatial warning signal diversification. Examining these mechanisms and their effects requires first confirming whether the most common morphs are favored at both local and regional scales. Empirical examples of this are uncommon and often include potentially confounding factors, such as a lack of knowledge of predator identity and behavior. We tested how bird behavior influences the survival of three coexisting morphs of the aposematic wood tiger moth Arctia plantaginis offered to a sympatric predator (great tit Parus major) at different frequencies. We found that although positive frequency-dependent selection is present, its strength is affected by predator characteristics and varying prey profitability. These results highlight the need to understand predator foraging in natural communities with variable prey defenses in order to better examine how behavioral interactions shape evolutionary outcomes. [ABSTRACT FROM AUTHOR]

Abstract Frequency‐dependent (FD) selection is a central process maintaining genetic variation and mediating evolution of population fitness. FD selection has attracted interest from researchers in a wide range of biological subdisciplines, including evolutionary genetics, behavioural ecology and, more recently, community ecology. However, the implications of frequency dependence for applied biological problems, particularly maladaptation, biological conservation and evolutionary rescue remain underexplored. The neglect of FD selection in conservation is particularly unfortunate. Classical theory, dating back to the 1940s, demonstrated that frequency dependence can either increase or decrease population fitness. These evolutionary consequences of FD selection are relevant to modern concerns about population persistence and the capacity of evolution to alleviate extinction risks. But exactly when should we expect FD selection to increase versus decrease absolute fitness and population growth? And how much of an impact is FD selection expected to have on population persistence versus extinction in changing environments? The answers to these questions have implications for evolutionary rescue under climate change and may inform strategies for managing threatened populations. Here, we revisit the core theory of FD selection, reviewing classical single‐locus models of population genetic change and outlining short‐ and long‐run consequences of FD selection for the evolution of population fitness. We then develop a quantitative genetic model of evolutionary rescue in a deteriorating environment, with population persistence hinging upon the evolution of a quantitative trait subject to both frequency‐dependent and frequency‐independent natural selection. We discuss the empirical literature pertinent to this theory, which supports key assumptions of our model. We show that FD selection can promote population persistence when it aligns with the direction of frequency‐independent selection imposed by abiotic environmental conditions. However, under most scenarios of environmental change, FD selection limits a population's evolutionary responsiveness to changing conditions and narrows the rate of environmental change that is evolutionarily tolerable.

Abstract Males of many fish species exhibit alternative reproductive tactics, which can influence the maturation schedules, fishery productivity, and resilience to harvest of exploited populations. While alternative mating phenotypes can persist in stable equilibria through frequency‐dependent selection, shifts in tactic frequencies have been observed and can have substantial consequences for fisheries. Here, we examine the dynamics of precocious sneaker males called “jacks” in a population of sockeye salmon (Oncorhynchus nerka) from Frazer Lake, Alaska. Jacks, which are of little commercial value due to their small body sizes, have recently been observed at unusually high levels in this stock, degrading the value of regional fisheries. To inform future strategies for managing the prevalence of jacks, we used long‐term monitoring data to identify what regulates the frequencies of alternative male phenotypes in the population over time. Expression of the jack life history could not be explained by environmental factors expected to influence juvenile body condition and maturation probability. Instead, we found a strong positive association between the proportion of individuals maturing as jacks within a cohort and the prevalence of jacks among the males that sired that cohort. Moreover, due to differences in age‐at‐maturity between male phenotypes, and large interannual variability in recruitment strength, jacks from strong year‐classes often spawn among older males from the weaker recruitments of earlier cohorts. Through such “cohort mismatches,” which are amplified by size‐selective harvest on older males, jacks frequently achieve substantial representation in the breeding population, and likely high total fertilizations. The repeated occurrence of these cohort mismatches appears to disrupt the stabilizing influence of frequency‐dependent selection, allowing the prevalence of jacks to exceed what might be expected under equilibrium conditions. These results emphasize that the dynamics of alternative life histories can profoundly influence fishery performance and should be explicitly considered in the management of exploited populations.

Multilevel selection on offspring size occurs when offspring fitness depends on both absolute size (hard selection) and size relative to neighbors (soft selection). We examined multilevel selection on egg size at two biological scales—within clutches and among clutches from different females—using an external fertilizing tube worm. We exposed clutches of eggs to two sperm environments (limiting and saturating) and measured their fertilization success. We then modeled environmental (sperm-dependent) differences in hard and soft selection on individual eggs as well as selection on clutch-level traits (means and variances). Hard and soft selection differed in strength and form depending on sperm availability—hard selection was consistently stabilizing; soft selection was directional and favored eggs relatively larger (sperm limitation) or smaller (sperm saturation) than the clutch mean. At the clutch level, selection on mean egg size was largely concave, while selection on within-clutch variance was weak but generally negative—although some correlational selection occurred between these two traits. Importantly, we found that the optimal clutch mean egg size differed for mothers and offspring, suggesting some antagonism between the levels of selection. We thus identify several pathways that may maintain offspring size variation: environmentally (sperm-) dependent soft selection, antagonistic multilevel selection, and correlational selection on clutch means and variances. Multilevel approaches are powerful but seldom-used tools for studies of offspring size, and we encourage their future use. [ABSTRACT FROM AUTHOR]

Maintenance of genetic polymorphism remains one of the big questions of evolutionary biology, which for a long time tended to be explained by balancing selection. This explanation was later criticized, but now is again accepted as an important mechanism in evolution. Human blood group systems seem affected by balancing selection especially strongly. In this preregistered study, we focused on stable coexistence of RhD‐positive and RhD‐negative subjects in a population. This is an evolutionary conundrum, because carriers of the less frequent negative allele suffer from lower fecundity due to haemolytic disease of the newborn affecting RhD‐positive infants born to RhD‐negative women. One explanation of persisting stability of RhD polymorphism points to heterozygote advantage. Over the past decade, numerous studies demonstrated that RhD‐positive subjects score better than RhD‐negative homozygotes in psychomotor tests and physical health‐related variables. Still, evidence of better health and performance of heterozygotes is scarce and merely indirect. We compared the physical and mental health of 2,539 subjects whose RhD genotype was estimated based on their and their parents' RhD phenotype. We confirmed that RhD‐negative homozygotes fare worse in terms of physical and mental health than subjects with RhD‐positive phenotype and that RhD‐positive heterozygotes enjoy better health than both homozygotes. For the first time, we demonstrated that RhD‐positive homozygotes might suffer from worse health than RhD‐negative homozygotes. Our results strongly support the hypothesis that RhD polymorphism is maintained by heterozygote advantage and that balancing selection may have played an important role in human evolution in this context and in general. [ABSTRACT FROM AUTHOR]

Intrasexual polychromatism has generally been linked to alternative reproductive and ecological tactics, where different morphs may exhibit differences in their behavioural, morphological, physiological, ecological, and life history characteristics. Therefore, these differences might favour the fitness of a certain morph over others. If multiple morphs are able to persist in the population however, they must have had similar fitness levels over a long period of time. This balance can be achieved via two processes: niche partitioning and negative frequency-dependent selection. The present study describes the aggressive sexual behaviour of the lizard Sceloporus minor and evaluates its possible relationships to ecological and behavioural aspects of two different male morphs (yellow and red) in a population from central Mexico. The results reveal that both the yellow and the red morphs are territorial. The home range (HR) of yellow males was 17 times larger than that of females, whereas the HR of red males was 13 times larger than that of females; however, there was no significant difference in HR size between morphs. Spatial distribution analysis indicated a pattern of regular distribution for adult S. minor. Our results also demonstrated that red males had more females within their HRs than yellow males had; however, yellow males were more aggressive than red males. In addition, both red and yellow morphs were similar in some other behavioural, ecological, and morphological traits, disproving the hypothesis that morphs in this population are maintained by niche partitioning. This leaves open the possibility that frequency-dependent selection maintains the polychromatism in this population. [ABSTRACT FROM AUTHOR]

Many animals mimic the behavior or the appearance of venomous snakes. When humans or other potential predators place their hand near the nest of tits belonging to the family Paridae (and a few other species), the incubating female performs a hissing display that mimics the inhalation hiss of a viper or another snake. They hiss vigorously while lunging their head forward and shaking their wings and tail, repeating this behavior several times. The structure of the hiss in tits is similar to that of the inhalation hiss of a snake, providing evidence of significant convergence of the mimic toward the model. The behavior of individual females is repeatable among trials. Individuals that flew away from their nest box only performed the hissing display on 6% of later trials, when present at their box, whereas individuals that did not fly away hissed on 28% of occasions, consistent with great tits Parus major either cautiously flying away or staying put on their nest while actively defending it. Individuals that flew away produced fewer chicks than individuals that stayed and hissed. The hissing display was more common when snakes were more abundant: 1) When breeding late during the season; 2) when breeding at sites with more snakes; and 3) when breeding in subtropical and tropical China with a higher abundance of snakes than in Denmark with a lower abundance. The frequency of nest predation was higher in sites with no snakes, and the frequency of predation increased with decreasing frequency of hissing display. These findings are consistent with expectations for frequency-dependent selection acting on snake mimicry. [ABSTRACT FROM AUTHOR]

Cultural diversity is disappearing quickly. Whilst a phylogenetic approach makes explicit the continuous extinction of cultures, and the generation of new ones, cultural evolutionary changes such as the rise of agriculture or more recently colonisation can cause periods of mass cultural extinction. At the current rate, 90% of languages will become extinct or moribund by the end of this century. Unlike biological extinction, cultural extinction does not necessarily involve genetic extinction or even deaths, but results from the disintegration of a social entity and discontinuation of culture-specific behaviours. Here we propose an analytical framework to examine the phenomenon of cultural extinction. When examined over millennia, extinctions of cultural traits or institutions can be studied in a phylogenetic comparative framework that incorporates archaeological data on ancestral states. Over decades or centuries, cultural extinction can be studied in a behavioural ecology framework to investigate how the fitness consequences of cultural behaviours and population dynamics shift individual behaviours away from the traditional norms. Frequency-dependent costs and benefits are key to understanding both the origin and the loss of cultural diversity. We review recent evolutionary studies that have informed cultural extinction processes and discuss avenues of future studies.

Floral scent, a key mediator in plant–pollinator interactions, varies not only among plant species, but also within species. In deceptive plants, it is assumed that variation in floral scents and other traits involved in pollinator attraction is maintained by negative frequency-dependent selection, i.e., rare phenotypes are more attractive to pollinators and hence, have a higher fitness than common phenotypes. So far, it is unknown whether the rarity of multivariate and/or continuous floral scent traits influences the pollination success of flowers. Here, we tested in the deceptive orchid Cypripedium calceolus, whether flowers with rarer scent bouquets within a population have a higher chance to getting pollinated than flowers with more common scents. We collected the scent of more than 100 flowers in two populations by dynamic headspace and analyzed the samples by gas chromatography coupled to mass spectrometry (GC/MS). From the same flowers we also recorded whether they set a fruit or not. We introduced rarity measures of uni- and multivariate floral scent traits for single flowers, which allowed us to finally test for frequency-dependent pollination, a prerequisite for negative frequency-dependent selection. Our results do not show rarity has an effect on the likelihood to set fruits in neither of the two populations and in none of the scent characteristics analyzed. Hence, there is no evidence of negative frequency-dependent pollination mediated by the floral scent of C. calceolus. We discuss that our approach to determine rarity of a scent is applicable to any univariate or multivariate (semi)quantitative trait.

Recent work incorporating demographic-genetic interactions indicates the importance of population size, gene flow, and selection in influencing local adaptation. This work typically assumes that density-dependent survival affects individuals equally, but individuals in natural population rarely compete equally. Among-individual differences in resource use generate stronger competition between more similar phenotypes (frequency-dependent competition) but it remains unclear how this additional form of selection changes the interactions between population size, gene flow, and local stabilizing selection. Here, we integrate migration-selection dynamics with frequency-dependent competition. We developed a coupled demographic-quantitative genetic model consisting of two patches connected by dispersal and subject to local stabilizing selection and competition. Our model shows that frequency-dependent competition slightly increases local adaptation, greatly increases genetic variance within patches, and reduces the amount that migration depresses population size, despite the increased genetic variance load. The effects of frequency-dependence depend on the strength of divergent selection, trait heritability, and when mortality occurs in the life cycle in relation to migration and reproduction. Essentially, frequency-dependent competition reduces the density-dependent interactions between migrants and residents, the extent to which depends on how different and common immigrants are compared to residents. Our results add new dynamics that illustrate how competition can alter the effects of gene flow and divergent selection on local adaptation and population carrying capacities.

Floral scent, a key mediator in plant–pollinator interactions, varies not only among plant species, but also within species. In deceptive plants, it is assumed that variation in floral scents and other traits involved in pollinator attraction is maintained by negative frequency-dependent selection, i.e., rare phenotypes are more attractive to pollinators and hence, have a higher fitness than common phenotypes. So far, it is unknown whether the rarity of multivariate and/or continuous floral scent traits influences the pollination success of flowers. Here, we tested in the deceptive orchid Cypripedium calceolus , whether flowers with rarer scent bouquets within a population have a higher chance to getting pollinated than flowers with more common scents. We collected the scent of more than 100 flowers in two populations by dynamic headspace and analyzed the samples by gas chromatography coupled to mass spectrometry (GC/MS). From the same flowers we also recorded whether they set a fruit or not. We introduced rarity measures of uni- and multivariate floral scent traits for single flowers, which allowed us to finally test for frequency-dependent pollination, a prerequisite for negative frequency-dependent selection. Our results do not show rarity has an effect on the likelihood to set fruits in neither of the two populations and in none of the scent characteristics analyzed. Hence, there is no evidence of negative frequency-dependent pollination mediated by the floral scent of C. calceolus. We discuss that our approach to determine rarity of a scent is applicable to any univariate or multivariate (semi)quantitative trait. [ABSTRACT FROM AUTHOR]

We construct a multitype constant-size population model allowing for general selective interactions as well as extreme reproductive events. Our multidimensional model aims for the generality of adaptive dynamics and the tractability of population genetics. It generalises the idea of Krone and Neuhauser [39] and González Casanova and Spanò [29], who represented the selection by allowing individuals to sample several potential parents in the previous generation before choosing the 'strongest' one, by allowing individuals to use any rule to choose their parent. The type of the newborn can even not be one of the types of the potential parents, which allows modelling mutations. Via a large population limit, we obtain a generalisation of $\Lambda$ -Fleming–Viot processes, with a diffusion term and a general frequency-dependent selection, which allows for non-transitive interactions between the different types present in the population. We provide some properties of these processes related to extinction and fixation events, and give conditions for them to be realised as unique strong solutions of multidimensional stochastic differential equations with jumps. Finally, we illustrate the generality of our model with applications to some classical biological interactions. This framework provides a natural bridge between two of the most prominent modelling frameworks of biological evolution: population genetics and eco-evolutionary models. [ABSTRACT FROM AUTHOR]

Warning signals are predicted to develop signal monomorphism via positive frequency‐dependent selection (+FDS) albeit many aposematic systems exhibit signal polymorphism. To understand this mismatch, we conducted a large‐scale predation experiment in four countries, among which the frequencies of hindwing warning coloration of the aposematic moth, Arctia plantaginis, differ. Here we show that selection by avian predators on warning colour is predicted by local morph frequency and predator community composition. We found +FDS to be the strongest in monomorphic Scotland and lowest in polymorphic Finland, where the attack risk of moth morphs depended on the local avian community. +FDS was also found where the predator community was the least diverse (Georgia), whereas in the most diverse avian community (Estonia), hardly any models were attacked. Our results support the idea that spatial variation in predator communities alters the strength or direction of selection on warning signals, thus facilitating a geographic mosaic of selection. [ABSTRACT FROM AUTHOR]

Understanding how environmental variation affects phenotypic evolution requires models based on ecologically realistic assumptions that include variation in population size and specific mechanisms by which environmental fluctuations affect selection. Here we generalize quantitative genetic theory for environmentally induced stochastic selection to include general forms of frequency‐ and density‐dependent selection. We show how the relevant fitness measure under stochastic selection relates to Fisher's fundamental theorem of natural selection, and present a general class of models in which density regulation acts through total use of resources rather than just population size. In this model, there is a constant adaptive topography for expected evolution, and the function maximized in the long run is the expected factor restricting population growth. This allows us to generalize several previous results and to explain why apparently "K‐selected" species with slow life histories often have low carrying capacities. Our joint analysis of density‐ and frequency‐dependent selection reveals more clearly the relationship between population dynamics and phenotypic evolution, enabling a broader range of eco‐evolutionary analyses of some of the most interesting problems in evolution in the face of environmental variation. [ABSTRACT FROM AUTHOR]