Studies Of Phenotypic Selection Research Articles

Natural selection by pollinators on floral attractive and defensive traits did not translate into selection via fruits in common milkweed

Considering both pollinator and herbivore pressures on plant reproductive and defensive traits is key to understanding patterns of selection for plants. However, phenotypic selection studies connecting floral traits and plant defenses with pollinator activity and herbivore damage remain rare. We used the common milkweed, Asclepias syriaca (Apocynaceae), to study phenotypic selection on attractive and defensive traits, and nectar rewards. We measured herbivore (leaf damage) and pollinator activity (pollinia movement) and quantified selection via female (pollinia insertions and fruit number) and male fitness (pollinia removals). We found selection to increase plant and inflorescence size and to decrease floral size (i.e. petal width) via female fitness. We also detected selection to increase floral but not leaf latex. The lack of selection on leaf latex was congruent with the low herbivory observed, however we also did not observe florivory in the population that would explain the advantage of more floral latex. Interestingly, we found selection on attractive traits differed via pollinia insertions and fruits initiated, suggesting that something other than pollinators was driving selection via fruit production. In contrast to female fitness, we did not find selection on any trait through male fitness, suggesting no sexual conflicting selection, at least through these proxies. Our findings reinforce the importance of the direct assessment of pollinator pressures in phenotypic selection studies before assuming pollinators as drivers of floral evolution by natural selection. Further work in southern populations closer to the centre of the species range, where herbivory and plant defense investment are higher, may help elucidate selection on attractive and defensive traits.

Phenotypic selection on an ornamental trait is not modulated by breeding density in a pied flycatcher population

Most studies of phenotypic selection in the wild have focussed on morphological and life‐history traits and looked at abiotic (climatic) variation as the main driver of selection. Consequently, our knowledge of the effects of biotic environmental variation on phenotypic selection on sexual traits is scarce. Population density can be considered a proxy for the intensity of intrasexual and intersexual competition and could therefore be a key factor influencing the covariation between individual fitness and the expression of sexual traits. Here, we used an individual‐based data set from a population of pied flycatchers (Ficedula hypoleuca) monitored over 24 years to analyze the effect of breeding density on phenotypic selection on dorsal plumage colouration, a heritable and sexually selected ornament in males of this species. Using the number of recruits as a fitness proxy, our results showed overall stabilizing selection on male dorsal colouration, with intermediate phenotypes being favoured over extremely dark and dull individuals. However, our results did not support the hypothesis that breeding density mediates phenotypic selection on this sexual trait. We discuss the possible role of other biotic factors influencing selection on ornamental plumage.

Trait-fitness associations do not predict within-species phenotypic evolution over 2 million years.

Long-term patterns of phenotypic change are the cumulative results of tens of thousands to millions of years of evolution. Yet, empirical and theoretical studies of phenotypic selection are largely based on contemporary populations. The challenges in studying phenotypic evolution, in particular trait–fitness associations in the deep past, are barriers to linking micro- and macroevolution. Here, we capitalize on the unique opportunity offered by a marine colonial organism commonly preserved in the fossil record to investigate trait–fitness associations over 2 Myr. We use the density of female polymorphs in colonies of Antartothoa tongima as a proxy for fecundity, a fitness component, and investigate multivariate signals of trait–fitness associations in six time intervals on the backdrop of Pleistocene climatic shifts. We detect negative trait–fitness associations for feeding polymorph (autozooid) sizes, positive associations for autozooid shape but no particular relationship between fecundity and brood chamber size. In addition, we demonstrate that long-term trait patterns are explained by palaeoclimate (as approximated by ∂18O), and to a lesser extent by ecological interactions (i.e. overgrowth competition and substrate crowding). Our analyses show that macroevolutionary outcomes of trait evolution are not a simple scaling-up from the trait–fitness associations.

On the standardization of fitness and traits in comparative studies of phenotypic selection.

Comparisons of the strength and form of phenotypic selection among groups provide a powerful approach for testing adaptive hypotheses. A central and largely unaddressed issue is how fitness and phenotypes are standardized in such studies; standardization across or within groups can qualitatively change conclusions whenever mean fitness differs between groups. We briefly reviewed recent relevant literature, and found that selection studies vary widely in their scale of standardization, but few investigators motivated their rationale for chosen standardization approaches. Here, we propose that the scale at which fitness should be relativized should reflect whether selection is likely to be hard or soft; that is, the scale at which populations (or hypothetical populations in the case of a contrived experiment) are regulated. We argue that many comparative studies of selection are implicitly or explicitly focused on soft selection (i.e., frequency and density-dependent selection). In such studies, relative fitness should preferably be calculated using within-group means, although this approach is taken only occasionally. Related difficulties arise for the standardization of phenotypes. The appropriate scale at which standardization should take place depends on whether groups are considered to be fixed or random. We emphasize that the scale of standardization is a critical decision in empirical studies of selection that should always warrant explicit justification.

An integrated analysis of phenotypic selection on insect body size and development time.

Most studies of phenotypic selection do not estimate selection or fitness surfaces for multiple components of fitness within a unified statistical framework. This makes it difficult or impossible to assess how selection operates on traits through variation in multiple components of fitness. We describe a new generation of aster models that can evaluate phenotypic selection by accounting for timing of life-history transitions and their effect on population growth rate, in addition to survival and reproductive output. We use this approach to estimate selection on body size and development time for a field population of the herbivorous insect, Manduca sexta (Lepidoptera: Sphingidae). Estimated fitness surfaces revealed strong and significant directional selection favoring both larger adult size (via effects on egg counts) and more rapid rates of early larval development (via effects on larval survival). Incorporating the timing of reproduction and its influence on population growth rate into the analysis resulted in larger values for size in early larval development at which fitness is maximized, and weaker selection on size in early larval development. These results illustrate how the interplay of different components of fitness can influence selection on size and development time. This integrated modeling framework can be readily applied to studies of phenotypic selection via multiple fitness components in other systems.

Modelling heterogeneity among fitness functions using random regression.

Summary Statistical approaches for testing hypotheses of heterogeneity in fitness functions are needed to accommodate studies of phenotypic selection with repeated sampling across study units, populations or years. In this study, we tested directly for among‐population variation in complex fitness functions and demonstrate a new approach for locating the region of the trait distribution where variation in fitness and traits is greatest. We modelled heterogeneity in fitness functions among populations by treating regression coefficients of fitness on traits as random variates. We applied random regression using two model specifications, (i) spline‐based curve and (ii) stepwise, to a 2‐year study of selection among 16 populations of the gall wasp, Belonocnema treatae. Log‐likelihood ratio tests of variance components and 10‐fold cross‐validation were used to assess the evidence that selection varied among populations. Ten‐fold cross‐validation prediction error sums of squares (PSS) indicated that spline‐based fitness functions were population specific and that the strength of evidence for heterogeneity in selection differed between years. Hypothesis testing of variance components from both models was consistent with the PSS results. Both the stepwise model and the local prediction error estimates of spline‐based fitness functions identified the region(s) of the phenotype distribution harbouring the greatest heterogeneity among populations. The adopted framework advances our understanding of phenotypic selection in natural populations by extending the analysis of spline‐based fitness functions to testing for heterogeneity among study units and isolating the regions of the phenotypic distribution where this variation is most pronounced.

Sexual selection on flight endurance, flight-related morphology and physiology in a scrambling damselfly

We have limited knowledge on the mechanistic base of sexual selection, especially in scrambling species. This asks for a functional approach that explores the link between each component of the phenotype-performance-fitness axis and that includes both morphological and physiological traits. We explored the phenotype-performance-fitness axis in the scrambling damselfly Coenagrion puella by studying the links between a set of physiological and morphological traits, flight performance (flight speed and flight endurance), and short-term mating success. As expected for scrambling competition, there was sexual selection for increased flight endurance rather than for increased flight speed. For fat content, we could demonstrate the full phenotype-performance-fitness axis, where selection for a higher fat content could be explained by the sexual selection for a higher flight endurance and the positive covariation between fat content and flight endurance. For three other traits (size, relative flight muscle mass and wing loading), however, we detected selection that could not be explained via their effect on flight performance, generating novel testable hypotheses about how the covariation between these traits and mating success is generated. This also urges caution when using morphological traits as proxies for flight speed and flight endurance in phenotypic selection studies.

Seed predators exert selection on the subindividual variation of seed size

Subindividual variation among repeated organs in plants constitutes an overlooked level of variation in phenotypic selection studies, despite being a major component of phenotypic variation. Animals that interact with plants could be selective agents on subindividual variation. This study examines selective pressures exerted during post-dispersal seed predation and germination on the subindividual variation of seed size in hawthorn (Crataegus monogyna). With a seed offering experiment and a germination test, we estimated phenotypic selection differentials for average and subindividual variation of seed size due to seed predation and germination. Seed size affects germination, growth rate and the probability of an individual seed of escaping predation. Longer seeds showed higher germination rates, but this did not result in significant selection on phenotypes of the maternal trees. On the other hand, seed predators avoided wider seeds, and by doing so exerted phenotypic selection on adult average and subindividual variation of seed size. The detected selection on subindividual variation suggests that the levels of phenotypic variation within individual plants may be, at least partly, the adaptive consequence of animal-mediated selection.

Phenotypic Selection in Natural Populations: What Limits Directional Selection?

Studies of phenotypic selection document directional selection in many natural populations. What factors reduce total directional selection and the cumulative evolutionary responses to selection? We combine two data sets for phenotypic selection, representing more than 4,600 distinct estimates of selection from 143 studies, to evaluate the potential roles of fitness trade-offs, indirect (correlated) selection, temporally varying selection, and stabilizing selection for reducing net directional selection and cumulative responses to selection. We detected little evidence that trade-offs among different fitness components reduced total directional selection in most study systems. Comparisons of selection gradients and selection differentials suggest that correlated selection frequently reduced total selection on size but not on other types of traits. The direction of selection on a trait often changes over time in many temporally replicated studies, but these fluctuations have limited impact in reducing cumulative directional selection in most study systems. Analyses of quadratic selection gradients indicated stabilizing selection on body size in at least some studies but provided little evidence that stabilizing selection is more common than disruptive selection for most traits or study systems. Our analyses provide little evidence that fitness trade-offs, correlated selection, or stabilizing selection strongly constrains the directional selection reported for most quantitative traits.

Which ecologically important traits are most likely to evolve rapidly?

Which ecologically important traits are most likely to evolve rapidly?

Darwin's beautiful contrivances: evolutionary and functional evidence for floral adaptation

Although not 'a professed botanist', Charles Darwin made seminal contributions to understanding of floral and inflorescence function while seeking evidence of adaptation by natural selection. This review considers the legacy of Darwin's ideas from three perspectives. First, we examine the process of floral and inflorescence adaptation by surveying studies of phenotypic selection, heritability and selection responses. Despite widespread phenotypic and genetic capacity for natural selection, only one-third of estimates indicate phenotypic selection. Second, we evaluate experimental studies of floral and inflorescence function and find that they usually demonstrate that reproductive traits represent adaptations. Finally, we consider the role of adaptation in floral diversification. Despite different diversification modes (coevolution, divergent use of the same pollen vector, pollinator shifts), evidence of pollination ecotypes and phylogenetic patterns suggests that adaptation commonly contributes to floral diversity. Thus, this review reveals a contrast between the inconsistent occurrence of phenotypic selection and convincing experimental and comparative evidence that floral traits are adaptations. Rather than rejecting Darwin's hypotheses about floral evolution, this contrast suggests that the tempo of creative selection varies, with strong, consistent selection during episodes of diversification, but relatively weak and inconsistent selection during longer, 'normal' periods of relative phenotypic stasis.

Phenotypic selection in common toad (Bufo bufo)

One of the most important problems for evolutionary biologists is to investigate the patterns and strength of phenotypic selection acting on quantitative traits in natural populations. Measurement of selection is complicated by the presence of correlations between characters; selection on a particular trait produces not only a direct effect, but indirect effects as well. Despite the growing body of phenotypic selection studies in a variety of taxa, studies on amphibians are still sparse. The aim of this study was to estimate patterns and strength of selection acting on a set of correlated characters in a natural population of Bufo bufo from the vicinity of Belgrade, Serbia. Morphological traits (body length, fore- and hind leg length) were measured, while fitness was assayed as fecundity and gonad weight for females and males, respec?tively. The regression approach was used to estimate selection differentials and gradients. Selection patterns differed between the sexes - linear selection differentials showed significant total directional selection for body size in females, but not in males. In males, differentials were significant for both fore- and hind leg lengths. Sample size did not permit identification of significant nonlinear (quadratic) selection.

EFFECTS OF NATURAL AND SEXUAL SELECTION ON ADAPTIVE POPULATION DIVERGENCE AND PREMATING ISOLATION IN A DAMSELFLY

The relative strength of different types of directional selection has seldom been compared directly in natural populations. A recent meta‐analysis of phenotypic selection studies in natural populations suggested that directional sexual selection may be stronger in magnitude than directional natural selection, although this pattern may have partly been confounded by the different time scales over which selection was estimated. Knowledge about the strength of different types of selection is of general interest for understanding how selective forces affect adaptive population divergence and how they may influence speciation. We studied divergent selection on morphology in parapatric, natural damselfly (Calopteryx splendens) populations. Sexual selection was stronger than natural selection measured on the same traits, irrespective of the time scale over which sexual selection was measured. Visualization of the fitness surfaces indicated that population divergence in overall morphology is more strongly influenced by divergent sexual selection rather than natural selection. Courtship success of experimental immigrant males was lower than that of resident males, indicating incipient sexual isolation between these populations. We conclude that current and strong sexual selection promotes adaptive population divergence in this species and that premating sexual isolation may have arisen as a correlated response to divergent sexual selection. Our results highlight the importance of sexual selection, rather than natural selection in the adaptive radiation of odonates, and supports previous suggestions that divergent sexual selection promotes speciation in this group.

EFFECTS OF NATURAL AND SEXUAL SELECTION ON ADAPTIVE POPULATION DIVERGENCE AND PREMATING ISOLATION IN A DAMSELFLY

The relative strength of different types of directional selection has seldom been compared directly in natural populations. A recent meta-analysis of phenotypic selection studies in natural populations suggested that directional sexual selection may be stronger in magnitude than directional natural selection, although this pattern may have partly been confounded by the different time scales over which selection was estimated. Knowledge about the strength of different types of selection is of general interest for understanding how selective forces affect adaptive population divergence and how they may influence speciation. We studied divergent selection on morphology in parapatric, natural damselfly (Calopteryx splendens) populations. Sexual selection was stronger than natural selection measured on the same traits, irrespective of the time scale over which sexual selection was measured. Visualization of the fitness surfaces indicated that population divergence in overall morphology is more strongly influenced by divergent sexual selection rather than natural selection. Courtship success of experimental immigrant males was lower than that of resident males, indicating incipient sexual isolation between these populations. We conclude that current and strong sexual selection promotes adaptive population divergence in this species and that premating sexual isolation may have arisen as a correlated response to divergent sexual selection. Our results highlight the importance of sexual selection, rather than natural selection in the adaptive radiation of odonates, and supports previous suggestions that divergent sexual selection promotes speciation in this group.

Variation in carbon isotope discrimination in relation to plant performance in a natural population of Cryptantha flava

Few studies of phenotypic selection have focused on physiological traits, especially in natural populations. The adaptive significance of plant water-use efficiency, the ratio of photosynthesis to water loss through transpiration, has rarely been examined. In this study, carbon isotopic discrimination, Delta, an integrated measure of water-use efficiency, was repeatedly measured in juveniles and adults in a natural population of the herbaceous desert perennial Cryptantha flava over a 4-year period and examined for plasticity in Delta, consistency between years in values of Delta, and evidence for selection on Delta phenotypes. There was significant concordance in Delta values among the 4 years for adult plants and significant correlations in Delta values measured in different years for juveniles and adults combined. The wettest year of the study, 1998, proved an exception because Delta values that year were not correlated with Delta values in any other year of the study. Consistency in Delta measured on the same plants in different years could indicate genotypic variation and/or consistency in the water status of the microhabitats the plants occupied. Two forms of plasticity in Delta were also evident; mean seasonal values were correlated with precipitation the preceding autumn, and Delta values also declined with plant size, indicating increasing water-use efficiency. Phenotypic selection was evident because in the first year of the study juvenile plants that would survive until year five averaged lower Delta values than did those that failed to survive. During the driest year, 2000, Delta was significantly negatively correlated with adult plant size, measured as the number of leaf rosettes, but the negative relationship between Delta and the number of flowering stalks, a more direct measure of fitness, was not significant. These results suggest that the direction of phenotypic selection on Delta changes as plants grow.

Temporal and microspatial variation in the intensities of natural and sexual selection in the yellow dung fly Scathophaga stercoraria

Abstract Studies of phenotypic selection in natural populations often concentrate only on short time periods and do not quantify selection intensities. We quantified temporal and microspatial variation in the intensities of natural and sexual selection for body size in the yellow dung fly over 2 years. Female fecundity selection intensity remained approximately constant over the season with an overall mean ± SE of 0.187 ± 0.014. Selection intensity for male reproductive success, defined as eggs obtained by mating males, did not differ from zero, indicating there was no assortative mating by size. Sexual selection intensity for male mating success favouring large males was variable but overall strong in the two years (0.499 ± 0.053 and 0.510 ± 0.051). As theoretically expected for male–male competition, sexual selection intensity increased with competitor density and reached an asymptote at about 250 males per pat; it also decreased with time in spring and increased again in autumn as a function of density. Small males had the best chance of obtaining a female at very low male densities. Greater selection intensity for large size in males than females is consistent with, and might be responsible for, the observed sexual size dimorphism in this species, as males are larger. The seasonal pattern of mean male body size (smallest at the beginning and end of the season) most likely reflects mere environmental (primarily temperature) influences on phenotypic size.

Measuring the Mating Skew

Charles Darwin (1871) defined sexual selection as a difference inreproductive success due to competition over mates, a definition that is still valid today. Being able to quantify such differences is thus essential for explaining the consequences of sexual selection, say, the ornamentation rdisplay behavior of males. In the specific ase of lekking (aggregated, non-resource-based display of males), attempts to do so have proven difficult. Any measure that does not take into account the possibility of random mating to produce the observed pattern of mating success is unsatisfactory as a measure of the intensity of sexual selection (Sutherland 1987; Mackenzie et al. 1995b). That variation in male mating success on leks is systematically higher than expected from pure random mating has been verified by Bradbury et al. (1985) and Mackenzie et al. (1995a). However, these studies do not yet provide means to estimate the expected mating success of individuals displaying together. Since a lek is not itself a trait but an assemblage of several males, each trying to optimize their own fitness (Hoglund and Alatalo 1995), fitness considerations of single individuals are required in explaining the evolution of lekking. The skew index, defined as the ratio of observed to maximum skew (Keller and Vargo 1993; Reeve and Ratnieks 1993), has been gaining popularity as a measure of skewed reproductive success (e.g., Bourke and Heinze 1994; Keller and Reeve 1994; Reeve and Keller 1995), and it has also been used to estimate the expected fitness of lekking individuals (Widemo and Owens 1995). However, this measure has been shown to have undesirable discontinuities (Pamilo and Crozier 1996). Moreover, it interprets random variation as real differences in mating success. This is most clearly seen in models of pure random mating, which nevertheless lead to positive values of the skew measure (Mackenzie et al. 1995b). Recently, Keller and Krieger (1996) proposed a method for correcting the skew index for random variation. We shall show, however, that the corrected measure still is not satisfactory, and we suggest a novel, statistically rigorous method. We give two criteria for a proper measure of skewed mating success. First, it should allow determining the most probable underlying distribution of mating

SPATIAL VARIATION IN SELECTIVE REGIMES: SEXUAL SELECTION IN THE WATER STRIDER, GERRIS ODONTOGASTER.

Studies of phenotypic selection in natural populations are often concerned with simply detecting selection. In adopting a more mechanistic approach, this study compares the sexual selection regimes in natural populations of the water strider Gerris odontogaster with a priori predictions of selection, based on a number of previous field and laboratory studies of the behavioral mechanisms of selection. In this species, a general reluctance of females to mate allows for intersexual selection for ability to subdue reluctant females in males. Female reluctance to mate has been shown to decrease with increasing population density, suggesting that sexual selection should be weaker in high density populations. Three different populations with large differences in population density were studied. A number of traits including parasite load, body mass, body size and male abdominal process length were found to experience significant sexual selection. The investigated populations differed considerably with regard to the total strength of selection on the measured traits and the form of selection on single traits. In general, males in the population with the highest density experienced the weakest selection for grasping ability. This pattern is ascribed to density-related alterations of female mating behavior. Selection for male grasping ability, as reflected by selection on male abdominal process length, is reduced in high-density populations where reluctant females are more easily subdued. Further, the studied populations differed significantly in mean phenotype and phenotypic variance for male abdominal process length. It is suggested that interpopulational differences in selective regimes may generate local adaptations with respect to male abdominal process length, and that gene flow may contribute to the maintenance of the high genetic variation in this trait. It is further suggested that more empirical effort should be made in quantifying and understanding spatial and temporal variation in selection in natural populations, since this may provide information on the prevalence of local adaptations in metric traits and on the mechanisms of selection.