Genetic distance predicts trait differentiation at the subpopulation but not the individual level in eelgrass,Zostera marina

Ecological studies often assume that genetically similar individuals will be more similar in phenotypic traits, such that genetic diversity can serve as a proxy for trait diversity. Here, we explicitly test the relationship between genetic relatedness and trait distance using 40 eelgrass (Zostera marina) genotypes from five sites within Bodega Harbor, CA. We measured traits related to nutrient uptake, morphology, biomass and growth, photosynthesis, and chemical deterrents for all genotypes. We used these trait measurements to calculate a multivariate pairwise trait distance for all possible genotype combinations. We then estimated pairwise relatedness from 11 microsatellite markers. We found significant trait variation among genotypes for nearly every measured trait; however, there was no evidence of a significant correlation between pairwise genetic relatedness and multivariate trait distance among individuals. However, at the subpopulation level (sites within a harbor), genetic (F_ST) and trait differentiation were positively correlated. Our work suggests that pairwise relatedness estimated from neutral marker loci is a poor proxy for trait differentiation between individual genotypes. It remains to be seen whether genomewide measures of genetic differentiation or easily measured “master” traits (like body size) might provide good predictions of overall trait differentiation.     

Ecological interactions drive evolutionary loss of traits

Loss of traits can dramatically alter the fate of species. Evidence is rapidly accumulating that the prevalence of trait loss is grossly underestimated. New findings demonstrate that traits can be lost without affecting the external phenotype, provided the lost function is compensated for by species interactions. This is important because trait loss can tighten the ecological relationship between partners, affecting the maintenance of species interactions. Here, we develop a new perspective on so-called `compensated trait loss' and how this type of trait loss may affect the evolutionary dynamics between interacting organisms. We argue that: (1) the frequency of compensated trait loss is currently underestimated because it can go unnoticed as long as ecological interactions are maintained; (2) by analysing known cases of trait loss, specific factors promoting compensated trait loss can be identified and (3) genomic sequencing is a key way forwards in detecting compensated trait loss. We present a comprehensive literature survey showing that compensated trait loss is taxonomically widespread, can involve essential traits, and often occurs as replicated evolutionary events. Despite its hidden nature, compensated trait loss is important in directing evolutionary dynamics of ecological relationships and has the potential to change facultative ecological interactions into obligatory ones.     

Trait‐based ecology of terrestrial arthropods

In focusing on how organisms' generalizable functional properties (traits) interact mechanistically with environments across spatial scales and levels of biological organization, trait‐based approaches provide a powerful framework for attaining synthesis, generality and prediction. Trait‐based research has considerably improved understanding of the assembly, structure and functioning of plant communities. Further advances in ecology may be achieved by exploring the trait–environment relationships of non‐sessile, heterotrophic organisms such as terrestrial arthropods, which are geographically ubiquitous, ecologically diverse, and often important functional components of ecosystems. Trait‐based studies and trait databases have recently been compiled for groups such as ants, bees, beetles, butterflies, spiders and many others; however, the explicit justification, conceptual framework, and primary‐evidence base for the burgeoning field of ‘terrestrial arthropod trait‐based ecology’ have not been well established. Consequently, there is some confusion over the scope and relevance of this field, as well as a tendency for studies to overlook important assumptions of the trait‐based approach. Here we aim to provide a broad and accessible overview of the trait‐based ecology of terrestrial arthropods. We first define and illustrate foundational concepts in trait‐based ecology with respect to terrestrial arthropods, and justify the application of trait‐based approaches to the study of their ecology. Next, we review studies in community ecology where trait‐based approaches have been used to elucidate how assembly processes for terrestrial arthropod communities are influenced by niche filtering along environmental gradients (e.g. climatic, structural, and land‐use gradients) and by abiotic and biotic disturbances (e.g. fire, floods, and biological invasions). We also review studies in ecosystem ecology where trait‐based approaches have been used to investigate biodiversity–ecosystem function relationships: how the functional diversity of arthropod communities relates to a host of ecosystem functions and services that they mediate, such as decomposition, pollination and predation. We then suggest how future work can address fundamental assumptions and limitations by investigating trait functionality and the effects of intraspecific variation, assessing the potential for sampling methods to bias the traits and trait values observed, and enhancing the quality and consolidation of trait information in databases. A roadmap to guide observational trait‐based studies is also presented. Lastly, we highlight new areas where trait‐based studies on terrestrial arthropods are well positioned to advance ecological understanding and application. These include examining the roles of competitive, non‐competitive and (multi‐)trophic interactions in shaping coexistence, and macro‐scaling trait–environment relationships to explain and predict patterns in biodiversity and ecosystem functions across space and time. We hope this review will spur and guide future applications of the trait‐based framework to advance ecological insights from the most diverse eukaryotic organisms on Earth.     

Habitat heterogeneity promotes intraspecific trait variability of shrub species in Australian granite inselbergs

The role of intraspecific trait variability is increasingly recognized as a key factor shaping plant fitness and community assembly worldwide. Studying the direct effects of habitat heterogeneity on trait expression of individual plants of the same species is a useful tool to quantify intraspecific trait variability locally. We investigated how habitat heterogeneity on granite inselbergs affected intraspecific trait variability of 19 functional traits in three shrub species of the family Proteaceae in south western Australia, a global biodiversity hotspot. We used pairwise comparison (single trait) and multivariate analysis (multiple traits, functional space) to detect shifts in trait patterns. Consistent with our predictions, we found that individuals developing in putatively more stressful habitats (highly sun-irradiated, shallow-soil patches on the outcrops) were characterized by trait expressions indicative of more conservative resource-related strategies when compared with plants occurring in the surrounding woodlands that were experiencing more benign ecological conditions. These results were significant for two out of three species. Granite inselbergs promoted plant longevity, a signal that these granite inselbergs might offer refugial conditions defined as protection against fire.     

Distinguishing the signatures of local environmental filtering and regional trait range limits in the study of trait–environment relationships

Understanding the imprint of environmental filtering on community assembly along environmental gradients is a key objective of trait‐gradient analyses. Depending on local constraints, this filtering generally entails that species departing from an optimum trait value have lower abundances in the community. The community‐weighted mean (CWM) and variance (CWV) of trait values are then expected to depict the optimum and intensity of filtering, respectively. However, the trait distribution within the regional species pool and its limits can also affect local CWM and CWV values apart from the effect of environmental filtering. The regional trait range limits are more likely to be reached in communities at the extremes of environmental gradients. Analogous to the mid‐domain effect in biogeography, decreasing CWV values in extreme environments can then represent the influence of regional trait range limits rather than stronger filtering in the local environment. We name this effect the ‘trait‐gradient boundary effect’ (TGBE). First, we use a community assembly framework to build simulated communities along a gradient from a species pool and environmental filtering with either constant or varying intensity while accounting for immigration processes. We demonstrate the significant influence of TGBE, in parallel to environmental filtering, on CWM and CWV at the extremes of the environmental gradient. We provide a statistical tool based on Approximate Bayesian Computation to decipher the respective influence of local environmental filtering and regional trait range limits. Second, as a case study, we reanalyze the functional composition of alpine plant communities distributed along a gradient of snow cover duration. We show that leaf trait convergence found in communities at the extremes of the gradient reflect an influence of trait range limits rather than stronger environmental filtering. These findings challenge correlative trait–environment relationships and call for more explicitly identifying the mechanisms responsible of trait convergence/divergence along environmental gradients.     

Integrating correlation between traits improves spatial predictions of plant functional composition

Functional trait composition is increasingly recognized as key to better understand and predict community responses to environmental gradients. Predictive approaches traditionally model the weighted mean trait values of communities (CWMs) as a function of environmental gradients. However, most approaches treat traits as independent regardless of known tradeoffs between them, which could lead to spurious predictions. To address this issue, we suggest jointly modeling a suit of functional traits along environmental gradients while accounting for relationships between traits. We use generalized additive mixed effect models to predict the functional composition of alpine grasslands in the Guisane Valley (France). We demonstrate that, compared to traditional approaches, joint trait models explain considerable amounts of variation in CWMs, yield less uncertainty in trait CWM predictions and provide more realistic spatial projections when extrapolating to novel environmental conditions. Modeling traits and their co‐variation jointly is an alternative and superior approach to predicting traits independently. Additionally, compared to a ‘predict first, assemble later’ approach that estimates trait CWMs post hoc based on stacked species distribution models, our ‘assemble first, predict later’ approach directly models trait‐responses along environmental gradients, and does not require data and models on species’ distributions, but only mean functional trait values per community plot. This highlights the great potential of joint trait modeling approaches in large‐scale mapping applications, such as spatial projections of the functional composition of vegetation and associated ecosystem services as a response to contemporary global change.     

Speed of Invasion of an Expanding Population by a Horizontally Transmitted Trait

Range expansions are a ubiquitous phenomenon, leading to the spatial spread of genetic, ecological, and cultural traits. While some of these traits are advantageous (and hence selected), other, nonselected traits can also spread by hitchhiking on the wave of population expansion. This requires us to understand how the spread of a hitchhiking trait is coupled to the wave of advance of its host population. Here, we use a system of coupled Fisher-Kolmogorov-Petrovsky-Piskunov (F-KPP) equations to describe the spread of a horizontally transmitted hitchhiking trait within a population as it expands. We extend F-KPP wave theory to the system of coupled equations to predict how the hitchhiking trait spreads as a wave within the expanding population. We show that the speed of this trait wave is controlled by an intricate coupling between the tip of the population and trait waves. Our analysis yields a new speed selection mechanism for coupled waves of advance and reveals the existence of previously unexpected speed transitions.     

Contemporary changes in phenotypic variation,and the potential consequences for eco-evolutionary dynamics

Most studies assessing rates of phenotypic change focus on population mean trait values, whereas a largely overlooked additional component is changes in population trait variation. Theoretically, eco-evolutionary dynamics mediated by such changes in trait variation could be as important as those mediated by changes in trait means. To date, however, no study has comprehensively summarised how phenotypic variation is changing in contemporary populations. Here, we explore four questions using a large database: How do changes in trait variances compare to changes in trait means? Do different human disturbances have different effects on trait variance? Do different trait types have different effects on changes in trait variance? Do studies that established a genetic basis for trait change show different patterns from those that did not? We find that changes in variation are typically small; yet we also see some very large changes associated with particular disturbances or trait types. We close by interpreting and discussing the implications of our findings in the context of eco-evolutionary studies.     

Genetic fatalism and social policy: the implications of behavior genetics research.

Recent advances in molecular genetics methods have provided new means of determining the genetic bases of human behavioral traits. The impetus for the use of these approaches for specific behaviors depends, in large part, on previous familial studies on inheritance of such traits. In the past, a finding of a genetic basis for a trait was often accompanied with the idea that that trait is unchangeable. We discuss the definition of "genetic trait" and heritability and examine the relationship between these concepts and the malleability of traits for both molecular and nonmolecular approaches to behavioral genetics. We argue that the malleability of traits is as much a social and political question as it is a biological one and that whether or not a trait is genetic has little relevance to questions concerning determinism, free will, and individual responsibility for actions. We conclude by noting that "scientific objectivity" should not be used to conceal the social perspectives that underlie proposals regarding social change.     

New method for rapid assessment of the functional composition of herbaceous plant communities

Abstract We propose a rapid sampling method to assess the functional composition of herbaceous plant communities without prior knowledge of the floristic composition. To determine the community‐level value of traits (‘aggregated trait values’) for a plant community, a standardized population‐centred method exists, but requires substantial manpower and reliable botanical knowledge. We tested an alternative method, the trait transect, using four subalpine pastures in the Beaufortain region (Northern French Alps) selected along a fertility gradient. We applied both methods to measure five commonly used ‘soft traits’ known to be responsive to soil nutrient availability: plant vegetative and reproductive height, specific leaf area, leaf dry matter and nitrogen contents. We tested whether the variation of these traits along the gradient detected with the population‐centred method was also detected with the trait transect. Both methods detected expected trends in the traits in response to the fertility gradient. The trait transect method was as efficient as the population‐centred method and is recommended as an appropriate tool for monitoring ecosystem changes in response to environmental conditions and management, especially in species‐rich communities.     

Global to community scale differences in the prevalence of convergent over divergent leaf trait distributions in plant assemblages

Aim The drivers of species assembly, by limiting the possible range of functional trait values, can lead to either convergent or divergent distributions of traits in realized assemblages. Here, to evaluate the strengths of these species assembly drivers, we partition trait variance across global, regional and community scales. We then test the hypothesis that, from global to community scales, the outcome of co‐occurring trait convergence and divergence is highly variable across biomes and communities. Location Global: nine biomes ranging from subarctic highland to tropical rain forest. Methods We analysed functional trait diversity at progressively finer spatial scales using a global, balanced, hierarchically structured dataset from 9 biomes, 58 communities and 652 species. Analyses were based on two key leaf traits (foliar nitrogen content and specific leaf area) that are known to drive biogeochemical cycling. Results While 35% of the global variance in these traits was between biomes, only 15% was between communities within biomes and as much as 50% occurred within communities. Despite this relatively high within‐community variance in trait values, we found that trait convergence dominated over divergence at both global and regional scales through comparisons of functional trait diversity in regional and community assemblages against random (null) models of species assembly. Main conclusions We demonstrate that the convergence of traits occurring from global to regional assemblages can be twice as strong as that from regional to community assemblages, and argue that large differences in the nature and strength of abiotic and biotic drivers of dominant species assembly can, at least partly, explain the variable outcome of simultaneous trait convergence and divergence across sites. Ultimately, these findings stress the urgent need to extend species assembly research to address those scales where trait variance is the highest, i.e. between biomes and within communities.     

Soil conditions and phylogenetic relatedness influence total community trait space during early plant succession

Aims The total space of traits covered by the members of plant communities is an important parameter of ecosystem functioning and complexity. We trace the variability of trait space during early plant succession and ask how trait space co-varies with phylogenetic community structure and soil conditions. Particularly, we are interested in the small-scale variability in trait space and the influence of biotic and abiotic filters.Methods We use data on species richness and soil conditions from the first 7 years of initial succession of an artificial catchment in north-eastern Germany. Total functional attribute diversity serves as a proxy to total trait space.Important findings Total trait space steadily increased during succession. We observed high small-scale variability in total trait space that was positively correlated with species richness and phylogenetic segregation and negatively correlated with total plant cover. Trait space increased with soil carbonate content, while pH and the fraction of sandy material behaved indifferently. Our results indicate that during early succession, habitat filtering processes gain importance leading to a lesser increase in trait space than expected from the increase in species richness alone.     

The dilemma of Fisherian sexual selection: Mate choice for indirect benefits despite rarity and overall weakness of trait‐preference genetic correlation

Fisher's mechanism of sexual selection is a fundamental element of evolutionary theory. In it nonrandom mate choice causes a genetic covariance between a male trait and female preference for that trait and thereby generates a positive feedback process sustaining accelerated coevolution of the trait and preference. Numerous theoretical models of Fisher's mechanism have confirmed its mathematical underpinnings, yet biologists have often failed to find evidence for trait‐preference genetic correlation in populations in which the mechanism was expected to function. We undertook a survey of the literature to conduct a formal meta‐analysis probing the incidence and strength of trait‐preference correlation among animal species. Our meta‐analysis found significant positive genetic correlations in fewer than 20% of the species studied and an overall weighted correlation that is slightly positive. Importantly, a significant positive correlation was not found in any thorough study that included multiple subgroups. We discuss several ways in which the dynamic, multivariate nature of mate choice may reduce the trait‐preference genetic correlation predicted by Fisher's mechanism. We then entertain the possibilities that Fisherian‐like processes sometimes function without genetic correlation, and that mate choice may persist in a population as long as genetic correlation, and therefore Fisher's mechanism, occurs intermittently.     

Trait hierarchies are stronger than trait dissimilarities in structuring spatial co‐occurrence patterns of common tree species in a subtropical forest

1. The dissimilarity and hierarchy of trait values that characterize niche and fitness differences, respectively, have been increasingly applied to infer mechanisms driving community assembly and to explain species co‐occurrence patterns. Here, we predict that limiting similarity should result in the spatial segregation of functionally similar species, while functionally similar species will be more likely to co‐occur either due to environmental filtering or due to competitive exclusion of inferior competitors (hereafter hierarchical competition).
2. We used a fully mapped 50‐ha subtropical forest plot in southern China to explore how pairwise spatial associations between saplings and between adult trees were influenced by trait dissimilarity and hierarchy in order to gain insight into assembly mechanisms. We assessed pairwise spatial associations using two summary statistics of spatial point patterns at different spatial scales and compared the effects of trait dissimilarity and trait hierarchy of different functional traits on the interspecific spatial associations. These comparisons allow us to disentangle the effects of limiting similarity, environmental filtering, and hierarchical competition on species co‐occurrence.
3. We found that trait dissimilarity was generally negatively related to interspecific spatial associations for both saplings and adult trees across spatial scales, meaning that species with similar trait values were more likely to co‐occur and thus supporting environmental filtering or hierarchical competition. We further found that trait hierarchy outweighed trait dissimilarity in structuring pairwise spatial associations, suggesting that hierarchical competition played a more important role in structuring our forest community than environmental filtering across life stages.
4. This study employed a novel method, by offering the integration of pairwise spatial association and trait dissimilarity as well as trait hierarchy, to disentangle the relative importance of multiple assembly mechanisms in structuring co‐occurrence patterns, especially the mechanisms of environmental filtering and hierarchical competition, which lead to indistinguishable co‐occurrence patterns. This study also reinforced the importance of trait hierarchy rather than trait dissimilarity in driving neighborhood competition.

     

Evolutionary dynamics and strong Allee effects

We describe the dynamics of an evolutionary model for a population subject to a strong Allee effect. The model assumes that the carrying capacity k(u), inherent growth rate r(u), and Allee threshold a(u) are functions of a mean phenotypic trait u subject to evolution. The model is a plane autonomous system that describes the coupled population and mean trait dynamics. We show bounded orbits equilibrate and that the Allee basin shrinks (and can even disappear) as a result of evolution. We also show that stable non-extinction equilibria occur at the local maxima of k(u) and that stable extinction equilibria occur at local minima of r(u). We give examples that illustrate these results and demonstrate other consequences of an Allee threshold in an evolutionary setting. These include the existence of multiple evolutionarily stable, non-extinction equilibria, and the possibility of evolving to a non-evolutionary stable strategy (ESS) trait from an initial trait near an ESS.     

Within‐species patterns challenge our understanding of the leaf economics spectrum

The utility of plant functional traits for predictive ecology relies on our ability to interpret trait variation across multiple taxonomic and ecological scales. Using extensive data sets of trait variation within species, across species and across communities, we analysed whether and at what scales leaf economics spectrum (LES) traits show predicted trait–trait covariation. We found that most variation in LES traits is often, but not universally, at high taxonomic levels (between families or genera in a family). However, we found that trait covariation shows distinct taxonomic scale dependence, with some trait correlations showing opposite signs within vs. across species. LES traits responded independently to environmental gradients within species, with few shared environmental responses across traits or across scales. We conclude that, at small taxonomic scales, plasticity may obscure or reverse the broad evolutionary linkages between leaf traits, meaning that variation in LES traits cannot always be interpreted as differences in resource use strategy.     

Does environmental heterogeneity drive functional trait variation? A test in montane and alpine meadows

While community‐weighted means of plant traits have been linked to mean environmental conditions at large scales, the drivers of trait variation within communities are not well understood. Local environmental heterogeneity (such as microclimate variability), in addition to mean environmental conditions, may decrease the strength of environmental filtering and explain why communities support different amounts of trait variation. Here, we assess two hypotheses: first, that more heterogeneous local environments and second, that less extreme environments, should support a broader range of plant strategies and thus higher trait variation. We quantified drivers of trait variation across a range of environmental conditions and spatial scales ranging from sub‐meter to tens of kilometers in montane and alpine plant communities. We found that, within communities, both environmental heterogeneity and environmental means are drivers of trait variation. However, the importance of each environmental factor varied depending on the trait. Our results indicate that larger‐scale trait–climate linkages that hold across communities also apply at small spatial scales, suggesting that microclimate variation within communities is a key driver of community functional diversity. Microclimatic variation provides a potential mechanism for helping to maintain diversity in local communities and also suggests that small‐scale environmental heterogeneity should be measured as a better predictor of functional diversity.     

An evolutionary analysis of the relationship between spite and altruism

We investigate the selective pressures on a social trait when evolution occurs in a population of constant size. We show that any social trait that is spiteful simultaneously qualifies as altruistic. In other words, any trait that reduces the fitness of less related individuals necessarily increases that of related ones. Our analysis demonstrates that the distinction between "Hamiltonian spite" and "Wilsonian spite" is not justified on the basis of fitness effects. We illustrate this general result with an explicit model for the evolution of a social act that reduces the recipient's survival ("harming trait"). This model shows that the evolution of harming is favoured if local demes are of small size and migration is low (philopatry). Further, deme size and migration rate determine whether harming evolves as a selfish strategy by increasing the fitness of the actor, or as a spiteful/altruistic strategy through its positive effect on the fitness of close kin.     

Evolutionarily unstable fitness maxima and stable fitness minima of continuous traits

Summary We present models of adaptive change in continuous traits for the following situations: (1) adaptation of a single trait within a single population in which the fitness of a given individual depends on the population's mean trait value as well as its own trait value; (2) adaptation of two (or more) traits within a single population; (3) adaptation in two or more interacting species. We analyse a dynamic model of these adaptive scenarios in which the rate of change of the mean trait value is an increasing function of the fitness gradient (i.e. the rate of increase of individual fitness with the individual's trait value). Such models have been employed in evolutionary game theory and are often appropriate both for the evolution of quantitative genetic traits and for the behavioural adjustment of phenotypically plastic traits. The dynamics of the adaptation of several different ecologically important traits can result in characters that minimize individual fitness and can preclude evolution towards characters that maximize individual fitness. We discuss biological circumstances that are likely to produce such adaptive failures for situations involving foraging, predator avoidance, competition and coevolution. The results argue for greater attention to dynamical stability in models of the evolution of continuous traits.