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How, if at all, can scientific progress improve our view of the modal facts? According to rationalist approaches to modal epistemology, science has no substantive role: a priori reflection reveals the structure of modal space, and a posteriori science merely locates us within that modal space, by identifying the actual properties and structures instantiated at our world. According to modal naturalist approaches, science provides evidence about the structure of the underlying modal space. In this paper I distinguish four versions of modal naturalism, with science playing an increasingly robust evidential role, and discuss their plausibility. [ABSTRACT FROM AUTHOR]

Among‐individual variation in cognitive traits, widely assumed to have evolved under adaptive processes, is increasingly being demonstrated across animal taxa. As variation among individuals is required for natural selection, characterizing individual differences and their heritability is important to understand how cognitive traits evolve. Here, we use a quantitative genetic study of wild‐type guppies repeatedly exposed to a 'detour task' to test for genetic variance in the cognitive trait of inhibitory control. We also test for genotype‐by‐environment interactions (GxE) by testing related fish under alternative experimental treatments (transparent vs. semi‐transparent barrier in the detour‐task). We find among‐individual variation in detour task performance, consistent with differences in inhibitory control. However, analysis of GxE reveals that heritable factors only contribute to performance variation in one treatment. This suggests that the adaptive evolutionary potential of inhibitory control (and/or other latent variables contributing to task performance) may be highly sensitive to environmental conditions. The presence of GxE also implies that the plastic response of detour task performance to treatment environment is genetically variable. Our results are consistent with a scenario where variation in individual inhibitory control stems from complex interactions between heritable and plastic components. [ABSTRACT FROM AUTHOR]

Sex differences in lifespan remain an intriguing puzzle in evolutionary biology. While explanations range from sex differences in selection to sex differences in the expression of recessive lifespan‐altering mutations (via X‐linkage), little consensus has been reached. One unresolved issue is the extent to which genetic influences on lifespan dimorphism are modulated by the environment. For example, studies have shown that sex differences in lifespan can either increase or decrease depending upon the social environment. Here, we took an experimental approach, manipulating multiple axes of the social environment across inbred long‐ and short‐lived genotypes and their reciprocal F1s in the fly Drosophila serrata. Our results reveal strong genetic effects and subtle yet significant genotype‐by‐environment interactions for male and female lifespan, specifically due to both population density and mating status. Further, our data do not support the idea that unconditional expression of deleterious X‐linked recessive alleles in heterogametic males accounts for lower male lifespan. [ABSTRACT FROM AUTHOR]

Within host populations, individuals can vary in their susceptibility to infections and in the severity and progression of disease once infected. Though mediated through differences in behaviour, resistance or tolerance, variation in disease outcomes ultimately stems from genetic and environmental (including social) factors. Despite obvious implications for the evolutionary, ecological and epidemiological dynamics of disease traits, the relative importance of these factors has rarely been quantified in naturally infected wild animal hosts. Here, we use a long‐term capture–mark–recapture study of group‐living European badgers (Meles meles) to characterize genetic and environmental sources of variation in host infection status by Mycobacterium bovis, the causative agent of bovine tuberculosis (bTB). We find that genetic factors contribute to M. bovis infection status, whether measured over a lifetime or across repeated captures. In the latter case, the heritability (h2) of infection status is close to zero in cubs and yearlings but increases in adulthood. Overall, environmental influences arising from a combination of social group membership (defined in time and space) and maternal effects appear to be more important than genetic factors. Thus, while genes do contribute to among‐individual variation, they play a comparatively minor role, meaning that rapid evolution of host defences under parasite‐mediated selection is unlikely (especially if selection is on young animals where h2 is lowest). Conversely, our results lend further support to the view that social and early‐life environments are important drivers of the dynamics of bTB infection in badger populations specifically, and of disease traits in wild hosts more generally. [ABSTRACT FROM AUTHOR]

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

The virulence‐transmission trade‐off hypothesis has provided a dominant theoretical basis for predicting pathogen virulence evolution, but empirical tests are rare, particularly at pathogen emergence. The central prediction of this hypothesis is that pathogen fitness is maximized at intermediate virulence due to a trade‐off between infection duration and transmission rate. However, obtaining sufficient numbers of pathogen isolates of contrasting virulence to test the shape of relationships between key pathogen traits, and doing so without the confounds of evolved host protective immunity (as expected at emergence), is challenging. Here, we inoculated 55 isolates of the bacterial pathogen, Mycoplasma gallisepticum, into non‐resistant house finches (Haemorhous mexicanus) from populations that have never been exposed to the disease. Isolates were collected over a 20‐year period from outbreak in disease‐exposed populations of house finches and vary markedly in virulence. We found a positive linear relationship between pathogen virulence and transmission rate to an uninfected sentinel, supporting the core assumption of the trade‐off hypothesis. Further, in support of the key prediction, there was no evidence for directional selection on a quantitative proxy of pathogen virulence and, instead, isolates of intermediate virulence were fittest. Surprisingly, however, the positive relationship between virulence and transmission rate was not underpinned by variation in pathogen load or replication rate as is commonly assumed. Our results indicate that selection favors pathogens of intermediate virulence at disease emergence in a novel host species, even when virulence and transmission are not linked to pathogen load. [ABSTRACT FROM AUTHOR]

To understand the evolution of cognitive abilities, we need to understand both how selection acts upon them and their genetic (co)variance structure. Recent work suggests that there are fitness consequences for free-living individuals with particular cognitive abilities. However, our current understanding of the heritability of these abilities is restricted to domesticated species subjected to artificial selection. We investigated genetic variance for, and genetic correlations among four cognitive abilities: inhibitory control, visual and spatial discrimination, and spatial ability, measured on >450 pheasants, Phasianus colchicus , over four generations. Pheasants were reared in captivity but bred from adults that lived in the wild and hence, were subject to selection on survival. Pheasant chicks are precocial and were reared without parents, enabling us to standardize environmental and parental care effects. We constructed a pedigree based on 15 microsatellite loci and implemented animal models to estimate heritability. We found moderate heritabilities for discrimination learning and inhibitory control (h2 = 0.17–0.23) but heritability for spatial ability was low (h2 = 0.09). Genetic correlations among-traits were largely positive but characterized by high uncertainty and were not statistically significant. Principle component analysis of the genetic correlation matrix estimate revealed a leading component that explained 69% of the variation, broadly in line with expectations under a general intelligence model of cognition. However, this pattern was not apparent in the phenotypic correlation structure which was more consistent with a modular view of animal cognition. Our findings highlight that the expression of cognitive traits is influenced by environmental factors which masks the underlying genetic structure. [ABSTRACT FROM AUTHOR]

Genetic factors underpinning phenotypic variation are required if natural selection is to result in adaptive evolution. However, evolutionary and behavioural ecologists typically focus on variation among individuals in their average trait values and seek to characterize genetic contributions to this. As a result, less attention has been paid to if and how genes could contribute towards within‐individual variance or trait 'predictability'. In fact, phenotypic 'predictability' can vary among individuals, and emerging evidence from livestock genetics suggests this can be due to genetic factors. Here, we test this empirically using repeated measures of a behavioural stress response trait in a pedigreed population of wild‐type guppies. We ask (a) whether individuals differ in behavioural predictability and (b) whether this variation is heritable and so evolvable under selection. Using statistical methodology from the field of quantitative genetics, we find support for both hypotheses and also show evidence of a genetic correlation structure between the behavioural trait mean and individual predictability. We show that investigating sources of variability in trait predictability is statistically tractable and can yield useful biological interpretation. We conclude that, if widespread, genetic variance for 'predictability' will have major implications for the evolutionary causes and consequences of phenotypic variation. [ABSTRACT FROM AUTHOR]

Theoretical models of animal contests such as the Hawk-Dove game predict that variation in fighting behavior will persist due to mixed evolutionarily stable strategies (ESS) under certain conditions. However, the genetic basis for this variation is poorly understood and a mixed ESS for fighting can be interpreted in more than one way. Specifically, we do not know whether variation in aggression within a population arises from among-individual differences in fixed strategy (determined by an individual's genotype—direct genetic effects [DGEs]), or from within-individual variation in strategy across contests. Furthermore, as suggested by developments of the original Hawk-Dove model, within-individual variation in strategy may be dependent on the phenotype and thus genotype of the opponent (indirect genetic effects—IGEs). Here we test for the effect of DGEs and IGEs during fights in the beadlet sea anemone Actinia equina. By exploiting the unusual reproductive system of sea anemones, combined with new molecular data, we investigate the role of both additive (DGE + IGE) and non-additive (DGE × IGE) genetic effects on fighting parameters, the latter of which have been hypothesized but never tested for explicitly. We find evidence for heritable variation in fighting ability and that fight duration increases with relatedness. Fighting success is influenced additively by DGEs and IGEs but we found no evidence for non-additive IGEs. These results indicate that variation in fighting behavior is driven by additive indirect genetic effects (DGE + IGE), and support a core assumption of contest theory that strategies are fixed by DGEs. [ABSTRACT FROM AUTHOR]

Latent personality traits underpinning observed behavioral variation have been studied in a great many species. However, a lack of standardized behavioral assays, coupled to a common reliance on inferring personality from a single, observed, behavioral trait makes it difficult to determine if, when, and how conclusions can be directly compared across taxa. Here, we estimate the among-individual (co)variance structure (ID) for a set of four behaviors expressed in an open field trial, putatively indicative of boldness, in seven species of small freshwater fish. We show that the ID matrices differ in terms of the total amount of variation present, and crucially the orientation, and as a consequence, biological interpretation of the first eigenvector. Specifically, loading of observed traits on the main axis of variation in ID matched a priori expectations for a shy-bold continuum in only three of the seven cases. Nonetheless, when the "shape" of the matrices was compared in higher dimensions, there was a high level of similarity among species, and weak evidence of phylogenetic signal. Our study highlights the present difficulty of trying to compare empirical inferences about specific personality traits across studies. However, it also shows how multivariate data collection and analysis allows the structure of behavioral variation to be quantitatively compared across populations or species without reliance on ambiguous verbal labels. This suggests that the field may have much to gain from greater uptake of phylogenetically informed comparative approaches when seeking to test evolutionary hypotheses about the origin and maintenance of personality variation. [ABSTRACT FROM AUTHOR]

Why do females of socially monogamous species engage in extra‐pair copulations? This long‐standing question remains a puzzle, because the benefits of female promiscuous behavior often do not seem to outweigh the costs. Genetic constraint models offer an answer by proposing that female promiscuity emerges through selection favoring alleles that are either beneficial for male reproductive success (intersexual pleiotropy hypothesis) or beneficial for female fecundity (intrasexual pleiotropy hypothesis). A previous quantitative genetic study on captive zebra finches, Taeniopygia guttata, reported support for the first, but not for the second hypothesis. Here, we re‐examine both hypotheses based on data from lines selected for high and low male courtship rate. In contrast to previous conclusions, our new analyses clearly reject the hypothesis that male and female promiscuity are genetically homologous traits. We find some support for a positive genetic correlation between female promiscuity and fecundity. This study also shows that the behavioral outcome of extra‐pair courtships primarily depends on individual‐specific female preferences and not on the "attractiveness" of the social mate. In contrast, patterns of paternity are strongly influenced by the social partner and the pair bond, presumably reflecting variation in copulation behavior, fertility, or sperm competitiveness. [ABSTRACT FROM AUTHOR]

Wild quantitative genetic studies have focused on a subset of traits (largely morphological and life history), with others, such as behaviors, receiving much less attention. This is because it is challenging to obtain sufficient data, particularly for behaviors involving interactions between individuals. Here, we explore an indirect approach for pilot investigations of the role of genetic differences in generating variation in parental care. Variation in parental genetic effects for offspring performance is expected to arise from among-parent genetic variation in parental care. Therefore, we used the animal model to predict maternal breeding values for lamb growth and used these predictions to select females for field observation, where maternal and lamb behaviors were recorded. Higher predicted maternal breeding value for lamb growth was associated with greater suckling success, but not with any other measures of suckling behavior. Though our work cannot explicitly estimate the genetic basis of the specific traits involved, it does provide a strategy for hypothesis generation and refinement that we hope could be used to justify data collection costs needed for confirmatory studies. Here, results suggest that behavioral genetic variation is involved in generating maternal genetic effects on lamb growth in Soay sheep. Though important caveats and cautions apply, our approach may extend the ability to initiate more genetic investigations of difficult-to-study behaviors and social interactions in natural populations. [ABSTRACT FROM AUTHOR]

A key element at the heart of the Fisher–Lande–Kirkpatrick model of the sexual selection process is the genetic correlation between (male) trait and (female) preference. The strength of this association is critical in determining a population's evolutionary trajectory, which is why estimating its magnitude is so important. In the Lande model, the trait-preference correlation is solely established and maintained by mate choice, and although it is unclear how exclusively mate choice does this in nature, the experimental designs typically employed to measure trait-preference genetic correlations could be systematically weakening estimates by not allowing free mate choice (similarly with husbandry practices). The precise impact of the problem is unknown, and possibly unknowable, but simple solutions can be applied to ensure the accuracy of trait-preference correlation estimates is maximized. [ABSTRACT FROM AUTHOR]

Understanding how hosts minimize the cost of emerging infections has fundamental implications for epidemiological dynamics and the evolution of pathogen virulence. Despite this, few experimental studies in natural populations have tested whether, in response to disease emergence, hosts evolve resistance, which reduces pathogen load through immune activation, or tolerance, which limits somatic damages without decreasing pathogen load. Further, none has done so accounting for significant natural variation in pathogen virulence, despite known effects on host responses to infection. Here, we investigate whether eastern North American house finches (Haemorhous mexicanus) have evolved resistance and/or tolerance to their emerging bacterial pathogen, Mycoplasma gallisepticum. To do so, we inoculated finches from disease‐exposed and disease‐unexposed populations with 55 distinct isolates of varying virulence. First, although peak pathogen loads, which occurred approximately eight days postinoculation, did not differ between experimentally inoculated finches from disease‐exposed versus unexposed population, pathogen loads subsequently decreased faster and to a greater extent in finches from exposed populations. These results suggest that finches from exposed populations are able to clear the infection through adaptive immune processes. Second, however, finches from exposed populations also displayed lower symptom severity for a given pathogen load, suggesting that a damage‐limitation mechanism, or tolerance, has accompanied the evolution of immune clearance. Our results highlight that resistance and tolerance should be seen as complementary, not alternative, defense strategies: the evolution of resistance benefits from the concomitant evolution of tolerance mechanisms that protect against the damage of immune activation, whereas the evolution of tolerance without resistance will risk runaway selection on pathogen virulence. [ABSTRACT FROM AUTHOR]

Organisms can affect one another's phenotypes when they socially interact. Indirect genetic effects occur when an individual's phenotype is affected by genes expressed in another individual. These heritable effects can enhance or reduce adaptive potential, thereby accelerating or reversing evolutionary change. Quantifying these social effects is therefore crucial for our understanding of evolution, yet estimates of indirect genetic effects in wild animals are limited to dyadic interactions. We estimated indirect phenotypic and genetic effects, and their covariance with direct effects, for the date of spring breeding in North American red squirrels (Tamiasciurus hudsonicus) living in an array of territories of varying spatial proximity. Additionally, we estimated indirect effects and the strength of selection at low and high population densities. Social effects of neighbours on the date of spring breeding were different from zero at high population densities but not at low population densities. Indirect phenotypic effects accounted for a larger amount of variation in the date of breeding than differences attributable to the among‐individual variance, suggesting social interactions are important for determining breeding dates. The genetic component to these indirect effects was however not statistically significant. We therefore showcase a powerful and flexible method that will allow researchers working in organisms with a range of social systems to estimate indirect phenotypic and genetic effects, and demonstrate the degree to which social interactions can influence phenotypes, even in a solitary species. [ABSTRACT FROM AUTHOR]

Movement of individuals, or their genes, can influence eco-evolutionary processes in structured populations. We have limited understanding of the extent to which spatial behavior varies among groups and individuals within populations. Here, we use genetic pedigree reconstruction in a long-term study of European badgers (Meles meles) to characterize the extent of extra-group paternity, occurring as a consequence of breeding excursions, and to test hypothesized drivers of variation at multiple levels. We jointly estimate parentage and paternity distance (PD; distance between a cub's natal and its father's social group), and test whether population density and sex ratio influence mean annual PD. We also model cub-level PD and extra-group paternity (EGP) to test for variation among social groups and parental individuals. Mean PD varied among years but was not explained by population density or sex ratio. However, cub-level analysis shows strong effects of social group, and parental identities, with some parental individuals being consistently more likely to produce cubs with extra-group partners. Group effects were partially explained by local sex ratio. There was also a strong negative correlation between maternal and paternal social group effects on cub paternity distance, indicating source-sink dynamics. Our analyses of paternity distance and EGP indicate variation in extra-group mating at multiple levels—among years, social groups and individuals. The latter in particular is a phenomenon seldom documented and suggests that gene flow among groups may be disproportionately mediated by a nonrandom subset of adults, emphasizing the importance of the individual in driving eco-evolutionary dynamics. [ABSTRACT FROM AUTHOR]

Individual repeatability (R), defined as the proportion of observed variance attributable to among‐individual differences, is a widely used summary statistic in evolutionarily motivated studies of morphology, life history, physiology and, especially, behaviour. Although statistical methods to estimate R are well known and widely available, there is a growing tendency for researchers to interpret R in ways that are subtly, but importantly, different. Some view R as a property of a dataset and a statistic to be interpreted agnostically with respect to mechanism. Others wish to isolate the contributions of 'intrinsic' and/or 'permanent' individual differences, and draw a distinction between true (intrinsic) and pseudo‐repeatability arising from uncontrolled extrinsic effects. This latter view proposes a narrower, more mechanistic interpretation, than the traditional concept of repeatability, but perhaps one that allows stronger evolutionary inference as a consequence (provided analytical pitfalls are successfully avoided). Neither perspective is incorrect, but if we are to avoid confusion and fruitless debate, there is a need for researchers to recognise this dichotomy, and to ensure clarity in relation to how, and why, a particular estimate of R is appropriate in any case. [ABSTRACT FROM AUTHOR]

Competition for resources including food, physical space, and potential mates is a fundamental ecological process shaping variation in individual phenotype and fitness. The evolution of competitive ability, in particular social dominance, depends on genetic (co)variation among traits causal (e.g., behavior) or consequent (e.g., growth) to competitive outcomes. If dominance is heritable, it will generate both direct and indirect genetic effects (IGE) on resource-dependent traits. The latter are expected to impose evolutionary constraint because winners necessarily gain resources at the expense of losers. We varied competition in a population of sheepshead swordtails, Xiphophorus birchmanni, to investigate effects on behavior, size, growth, and survival. We then applied quantitative genetic analyses to determine (i) whether competition leads to phenotypic and/or genetic integration of behavior with life history and (ii) the potential for IGE to constrain life history evolution. Size, growth, and survival were reduced at high competition. Male dominance was repeatable and dominant individuals show higher growth and survival. Additive genetic contributions to phenotypic covariance were significant, with the G matrix largely recapitulating phenotypic relationships. Social dominance has a low but significant heritability and is strongly genetically correlated with size and growth. Assuming causal dependence of growth on dominance, hidden IGE will therefore reduce evolutionary potential. [ABSTRACT FROM AUTHOR]

There is often large divergence in the effects of key nutrients on life span (LS) and reproduction in the sexes, yet nutrient intake is regulated in the same way in males and females given dietary choice. This suggests that the sexes are constrained from feeding to their sex-specific nutritional optima for these traits. Here, we examine the potential for intralocus sexual conflict (IASC) over optimal protein and carbohydrate intake for LS and reproduction to constrain the evolution of sex-specific nutrient regulation in the field cricket, Teleogryllus commodus. We show clear sex differences in the effects of protein and carbohydrate intake on LS and reproduction and strong positive genetic correlations between the sexes for the regulated intake of these nutrients. However, the between-sex additive genetic covariance matrix had very little effect on the predicted evolutionary response of nutrient regulation in the sexes. Thus, IASC appears unlikely to act as an evolutionary constraint on sex-specific nutrient regulation in T. commodus. This finding is supported by clear sexual dimorphism in the regulated intake of these nutrients under dietary choice. However, nutrient regulation did not coincide with the nutritional optima for LS or reproduction in either sex, suggesting that IASC is not completely resolved in T. commodus. [ABSTRACT FROM AUTHOR]

Having recognized that variation around the population-level "Golden Mean" of labile traits contains biologically meaningful information, behavioural ecologists have focused increasingly on exploring the causes and consequences of individual variation in behaviour. These are exciting new directions for the field, assisted in no small part by the adoption of mixed-effects modelling techniques that enable the partitioning of among- and within-individual behavioural variation. It has become commonplace to extract predictions of individual random effects from such models for use in subsequent analyses (for example, between a personality trait and other individual traits such as cognition, physiology, or fitness-related measures). However, these predictions are made with large amounts of error that is not carried forward, rendering further tests susceptible to spurious P values from these individual-level point estimates. We briefly summarize the problems with such statistical methods that are used regularly by behavioural ecologists, and highlight the robust solutions that exist within the mixed model framework, providing tutorials to aid in their implementation. [ABSTRACT FROM AUTHOR]

Sexual selection can target many different types of traits. However, the relative influence of different sexually selected traits during evolutionary divergence is poorly understood. We used the field cricket Teleogryllus oceanicus to quantify and compare how five traits from each of three sexual signal modalities and components diverge among allopatric populations: male advertisement song, cuticular hydrocarbon (CHC) profiles and forewing morphology. Population divergence was unexpectedly consistent: we estimated the among-population (genetic) variance-covariance matrix, D, for all 15 traits, and Dmax explained nearly two-thirds of its variation. CHC and wing traits were most tightly integrated, whereas song varied more independently. We modeled the dependence of among-population trait divergence on genetic distance estimated from neutral markers to test for signatures of selection versus neutral divergence. For all three sexual trait types, phenotypic variation among populations was largely explained by a neutral model of divergence. Our findings illustrate how phenotypic integration across different types of sexual traits might impose constraints on the evolution of mating isolation and divergence via sexual selection. [ABSTRACT FROM AUTHOR]

Some species actively change color and pattern for camouflage on a range of background types. Such dynamic camouflage may be particularly advantageous for species inhabiting heterogeneous habitats, such as intertidal zones, where individuals are exposed to both terrestrial and marine predators depending on tides and wave action. Most studies of dynamic pattern camouflage have focused on relatively few species, and rarely species inhabiting the intertidal zone. We used image analysis and predator (avian) vision modelling to determine if rock gobies (Gobius paganellus) change their body pattern in response to their background, and to explore how background marking size influence pattern change. Rock gobies rapidly (within 1 min) changed their pattern when placed on checkerboards with different sized squares, and on backgrounds resembling natural substrates. On backgrounds resembling natural substrates, those with a small grain size, such as sand, elicited a larger degree of pattern change than those with a larger grain size. However, despite this, the majority of fish showed little or no improvement in background matching over time. Instead, the markings elicited are characteristic of disruptive coloration and may function primarily through breaking up the body outline rather than via improved match to the background pattern itself. [ABSTRACT FROM AUTHOR]

In animal contests, individuals respond plastically to the phenotypes of the opponents that they confront. These "opponent"—or "indirect"—effects are often repeatable, for example, certain opponents consistently elicit more or less aggressiveness in others. "Personality" (repeatable among-individual variation in behavior) has been proposed as an important source of indirect effects. Here, we repeatedly assayed aggressiveness of wild-caught adult male field crickets Gryllus campestris in staged dyadic fights, measuring aggressiveness of both contestants. Measurements of their personality in nonsocial contexts (activity and exploration behavior) enabled us to ask whether personality caused indirect effects on aggressiveness. Activity, exploration, and aggressiveness were positively associated into a behavioral syndrome eliciting aggressiveness in conspecifics, providing direct evidence for the role of personality in causing indirect effects. Our findings imply that a multivariate view of phenotypes that includes indirect effects greatly improves our ability to understand the ecology and evolution of behavior. [ABSTRACT FROM AUTHOR]

Debates about how coevolution of sexual traits and preferences might promote evolutionary diversification have permeated speciation research for over a century. Recent work demonstrates that the expression of such traits can be sensitive to variation in the social environment. Here, we examined social flexibility in a sexually selected male trait-cuticular hydrocarbon (CHC) profiles-in the field cricket Teleogryllus oceanicus and tested whether population genetic divergence predicts the extent or direction of social flexibility in allopatric populations. We manipulated male crickets' social environments during rearing and then characterized CHC profiles. CHC signatures varied considerably across populations and also in response to the social environment, but our prediction that increased social flexibility would be selected in more recently founded populations exposed to fluctuating demographic environments was unsupported. Furthermore, models examining the influence of drift and selection failed to support a role of sexual selection in driving population divergence in CHC profiles. Variation in social environments might alter the dynamics of sexual selection, but our results align with theoretical predictions that the role social flexibility plays in modulating evolutionary divergence depends critically on whether responses to variation in the social environment are homogeneous across populations, or whether gene by social environment interactions occur. [ABSTRACT FROM AUTHOR]

Many reproductive traits that have evolved under sexual conflict may be influenced by both sexes. Investigation of the genetic architecture of such traits can yield important insight into their evolution, but this entails that the heritable component of variation is estimated for males and females-as an interacting phenotype. We address the lack of research in this area through an investigation of egg production and copula duration in the fruit fly, Drosophila melanogaster. Despite egg production rate being determined by both sexes, which may cause sexual conflict, an assessment of this trait as an interacting phenotype is lacking. It is currently unclear whether copula duration is determined by males and/or females. We found significant female, but not male, genetic variance for egg production rate that may indicate reduced potential for ongoing sexually antagonistic coevolution. In contrast, copula duration was determined by significant genetic variance in both sexes. We also identified genetic variation in egg retention among virgin females. Although previously identified in wild populations, it is unclear why this should be present in a laboratory stock. This study provides a novel insight into the shared genetic architecture of reproductive traits that are the subject of sexual conflict. [ABSTRACT FROM AUTHOR]

Adaptive evolution occurs when fitness covaries with genetic merit for a trait (or traits). The breeder's equation (BE), in both its univariate and multivariate forms, allows us to predict this process by combining estimates of selection on phenotype with estimates of genetic (co)variation. However, predictions are only valid if all factors causal for trait-fitness covariance are measured. Although this requirement will rarely (if ever) be met in practice, it can be avoided by applying Robertson's secondary theorem of selection (STS). The STS predicts evolution by directly estimating the genetic basis of trait-fitness covariation without any explicit model of selection. Here we apply the BE and STS to four morphological traits measured in Soay sheep ( Ovis aries) from St. Kilda. Despite apparently positive selection on heritable size traits, sheep are not getting larger. However, although the BE predicts increasing size, the STS does not, which is a discrepancy that suggests unmeasured factors are upwardly biasing our estimates of selection on phenotype. We suggest this is likely to be a general issue, and that wider application of the STS could offer at least a partial resolution to the common discrepancy between naive expectations and observed trait dynamics in natural populations. [ABSTRACT FROM AUTHOR]

Simon Saunders and David Wallace have proposed an attractive semantics for interpreting linguistic communities embedded in an Everettian multiverse. It provides a charitable interpretation of our ordinary talk about the future, and allows us to retain a principle of bivalence for propositions and to retain the law of excluded middle in the logic of propositions about the future. But difficulties arise when it comes to providing an appropriate account of the metaphysics of macroscopic objects and events. I evaluate various metaphysical frameworks which might be combined with the Saunders-Wallace semantics. I conclude that the most appropriate metaphysics to underwrite the semantics renders Everettian quantum mechanics a theory of non-overlapping worlds. [ABSTRACT FROM AUTHOR]

Cryptic evolution has been defined as adaptive evolutionary change being masked by concurrent environmental change. Empirical studies of cryptic evolution have usually invoked a changing climate and/ or increasing population density as the form of detrimental environmental change experienced by a population undergoing cryptic evolution. However, Fisher (1958) emphasized that evolutionary change in itself is likely to be an important component of "environmental deterioration," a point restated by Cooke et al. (1990) in the context of intraspecific competition. In this form, environmental deterioration arises because a winning lineage has to compete against more winners in successive generations as the population evolves. This "evolutionary environmental deterioration" has different implications for the selection and evolution of traits influenced by resource competition than general environmental change. We reformulate Cooke's model as a quantitative genetic model to show that it is identical in form to more recent developments proposed by quantitative geneticists. This provides a statistical framework for discriminating between the alternative hypotheses of environmental change and environmental deterioration caused by evolutionary change. We also demonstrate that in systems where no phenotypic change has occurred, there are many reasonable biological processes that will generate patterns in predicted breeding values that are consistent with what has been interpreted as cryptic evolution, and care needs to be taken when interpreting these patterns. These processes include mutation, sib competition, and invisible fractions. [ABSTRACT FROM AUTHOR]

Phenotypic plasticity is an important mechanism via which populations can respond to changing environmental conditions, but we know very little about how natural populations vary with respect to plasticity. Here we use random-regression animal models to understand the multivariate phenotypic and genetic patterns of plasticity variation in two key life-history traits, laying date and clutch size, using data from long-term studies of great tits in The Netherlands (Hoge Veluwe [HV]) and UK (Wytham Woods [WW]). We show that, while population-level responses of laying date and clutch size to temperature were similar in the two populations, between-individual variation in plasticity differed markedly. Both populations showed significant variation in phenotypic plasticity (IxE) for laying date, but IxE was significantly higher in HV than in WW. There were no significant genotype-by-environment interactions (GxE) for laying date, yet differences in GxE were marginally nonsignificant between HV and WW. For clutch size, we only found significant IxE and GxE in WW but no significant difference between populations. From a multivariate perspective, plasticity in laying date was not correlated with plasticity in clutch size in either population. Our results suggest that generalizations about the form and cause of any response to changing environmental conditions across populations may be difficult. [ABSTRACT FROM AUTHOR]

The independent evolution of the sexes may often be constrained if male and female homologous traits share a similar genetic architecture. Thus, cross-sex genetic covariance is assumed to play a key role in the evolution of sexual dimorphism (SD) with consequent impacts on sexual selection, population dynamics, and speciation processes. We compiled cross-sex genetic correlations ( rMF) estimates from 114 sources to assess the extent to which the evolution of SD is typically constrained and test several specific hypotheses. First, we tested if rMF differed among trait types and especially between fitness components and other traits. We also tested the theoretical prediction of a negative relationship between rMF and SD based on the expectation that increases in SD should be facilitated by sex-specific genetic variance. We show that rMF is usually large and positive but that it is typically smaller for fitness components. This demonstrates that the evolution of SD is typically genetically constrained and that sex-specific selection coefficients may often be opposite in sign due to sub-optimal levels of SD. Most importantly, we confirm that sex-specific genetic variance is an important contributor to the evolution of SD by validating the prediction of a negative correlation between rMF and SD. [ABSTRACT FROM AUTHOR]

This work demonstrates that environmental conditions experienced by individuals can shape their development and affect the stability of genetic associations. The implication of this observation is that the environmental response may influence the evolution of traits in the wild. Here, we examined how the genetic architecture of a suite of sexually dimorphic traits changed as a function of environmental conditions in an unmanaged population of Soay sheep (Ovis aries) on the island of Hirta, St. Kilda, northwest Scotland. We examined the stability of phenotypic, genetic, and environmental (residual) covariance in males during the first year of life between horn length, body weight, and parasite load in environments of different quality. We then examined the same covariance structures across environments within and between the adult sexes. We found significant genotype-by-environment interactions for lamb male body weight and parasite load, leading to a change in the genetic correlation among environments. Horn length was genetically correlated with body weight in males but not females and the genetic correlation among traits within and between the sexes was dependent upon the environmental conditions experienced during adulthood. Genetic correlations were smaller in more favorable environmental conditions, suggesting that in good environ- merits, loci are expressed that have sex-specific effects. The reduction in genetic correlation between the sexes may allow independent evolutionary trajectories for each sex. This study demonstrates that the genetic architecture of traits is not stable under temporally varying environments and highlights the fact that evolutionary processes may depend largely upon ecological conditions. [ABSTRACT FROM AUTHOR]

FST and RST estimates for Arctic charr from six microsatelite markers collected from two neighbouring Scottish lakes, Loch Maree and Loch Stack, confirm the presence of two distinct genetic groupings representing separate populations within each lake. In both lakes, there was also a clear body size dimorphism, with large and small body size forms that segregated according to genetic grouping. There was evidence of only subtle foraging ecology differences between morphs, with the small body size morph in both lakes being more generalist in its foraging in the summer (consuming mostly plankton but also some macrobenthos) than the large body size morph, which specialized on planktonic prey. Trophic morphology (head and mouth shape) did not differ significantly between morphs (although the small sample size for Maree makes this a preliminary finding). Cluster analysis of the microsatelite data and the presence of private alleles showed that morphologically similar forms in different lakes were not genetically similar, as would be expected under a multiple invasion hypothesis. Thus, the data do not support a hypothesis of a dual invasion of both lakes by two common ancestors but instead suggest an independent origin of the two forms in each lake. Thus parallel sympatric divergence as a result of common selection pressures in both lakes is the most parsimonious explanation of the evolutionary origin of these polymorphisms. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 748–757. [ABSTRACT FROM AUTHOR]

A population's potential for evolutionary change depends on the amount of genetic variability expressed in traits under selection. Studies attempting to measure this variability typically do so over the life span of individuals, but theory suggests that the amount of additive genetic variance can change during the course of individuals' lives. Here we use pedigree data from historical Finns and a quantitative genetic framework to investigate how female fecundity, throughout an individual's reproductive life, is influenced by “maternal” versus additive genetic effects. We show that although maternal effects explain variation in female fecundity early in life, these effects wane with female age. Moreover, this decline in maternal effects is associated with a concomitant increase in additive genetic variance with age. Our results thus highlight that single over-lifetime estimates of trait heritability may give a misleading view of a trait's potential to respond to changing selection pressures. [ABSTRACT FROM AUTHOR]

Reproductive and early life-history traits can be considered aspects of either offspring or maternal phenotype, and their evolution will therefore depend on selection operating through offspring and maternal components of fitness. Furthermore, selection at these levels may be antagonistic, with optimal offspring and maternal fitness occurring at different phenotypic values. We examined selection regimes on the correlated traits of birth weight, birth date, and litter size in Soay sheep (Ovis aries) using data from a long-term study of a free-living population on the archipelago of St. Kilda, Scotland. We tested the hypothesis that selective constraints on the evolution of the multivariate phenotype arise through antagonistic selection, either acting at offspring and maternal levels, or on correlated aspects of phenotype. All three traits were found to be under selection through variance in short-term and lifetime measures of fitness. Analysis of lifetime fitness revealed strong positive directional selection on birth weight and weaker selection for increased birth date at both levels. However, there was also evidence for stabilizing selection on these traits at the maternal level, with reduced fitness at high phenotypic values indicating lower phenotypic optima for mothers than for offspring. Additionally, antagonistic selection was found on litter size. From the offspring's point of view it is better to be born a singleton, whereas maternal fitness increases with average litter size. The decreased fitness of twins is caused by their reduced birth weight; therefore, this antagonistic selection likely results from trade- offs between litter size and birth weight that have different optimal resolutions with respect to offspring and maternal fitness. Our results highlight how selection regimes may vary depending on the assignment of reproductive and early life-history traits to either offspring or maternal phenotype. [ABSTRACT FROM AUTHOR]

The article offers the author's insights on the study "Indirect genetic effects in behavioral ecology: does behavior play a special role in evolution?" by N. W. Bailey and colleagues. Topics include the possible indirect genetic effects (IGEs) on the behavioral traits, the issues on IGE variance partitioning such as dyadic interactions, and the plasticity in the social environment.

BIJMA et al. (2007a,b) presented a quantitative genetic theory of multilevel selection and showed how to estimate the relevant parameters using standard restricted maximum-likelihood (REML) methodology. Extending their results we develop a wider class of models that provide a more realistic framework for capturing the effects of interacting individuals. These models also make use of standard REML techniques and include the original model as a special case. [ABSTRACT FROM AUTHOR]

How do environmental conditions influence selection and genetic variation in wild populations? There is widespread evidence for selection-by-environment interactions (S*E), but we reviewed studies of natural populations estimating the extent of genotype-by-environment interactions (G*E) in response to natural variation in environmental conditions and found that evidence for G*E appears to be rare within single populations in the wild. Studies estimating the simultaneous impact of environmental variation on both selection and genetic variation are especially scarce. Here, we used 24 years of data collected from a wild Soay sheep population to quantify how an important environmental variable, population density, impacts upon (1) selection through annual contribution to fitness and (2) expression of genetic variation, in six morphological and life history traits: body weight, hind leg length, parasite burden, horn length, horn growth, and testicular circumference. Our results supported the existence of S*E: selection was stronger in years of higher population density for all traits apart from horn growth, with directional selection being stronger under more adverse conditions. Quantitative genetic models revealed significant additive genetic variance for body weight, leg length, parasite burden, horn length, and testes size, but not for horn growth or our measure of annual fitness. However, random regression models found variation between individuals in their responses to the environment in only three traits, and did not support the presence of G*E for any trait. Our analyses of St Kilda Soay sheep data thus concurs with our cross-study review that, while natural environmental variation within a population can profoundly alter the strength of selection on phenotypic traits, there is less evidence for its effect on the expression of genetic variance in the wild. [ABSTRACT FROM AUTHOR]