Spatially structured intraspecific trait variation can foster biodiversity in disturbed,heterogeneous environments

Trait variation within populations is an important area of research for empirical and theoretical ecologists. While differences between individuals are doubtlessly ubiquitous, their role for species coexistence is much less clear and highly debated. Both unstructured (random) and structured (linked to space, time or inheritance) intraspecific trait variation (ITV) may modify species interactions with nontrivial consequences for emerging community compositions. In many ecosystems, these compositions are further driven by prevalent disturbance regimes. I therefore explored the effects of unstructured as well as spatially structured ITV under disturbances in a generic ecological model of competing sessile species. Using spatially explicit, individual‐based simulations, I studied how intraspecific variation in life history traits together with interspecific tradeoffs and disturbance regimes shape long‐term community composition. I found that 1) unstructured ITV does not affect species coexistence in the given context, 2) spatially structured ITV may considerably increase coexistence, but 3) spatially clumped disturbances reduce this effect of spatially structured ITV, especially if interspecific tradeoffs involve dispersal distance. The findings suggest that spatially structured ITV with individual trait responses to local habitat conditions differing among species may create or expand humps in disturbance–diversity relationships. Hence, if present, these forms of spatially structured ITV should be included in ecological models and will be important for reliably assessing community responses to environmental heterogeneity and change.     

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta‐analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole‐plant (e.g. plant height) vs. organ‐level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait‐based community and ecosystem studies.     

Species divergence and trait convergence in experimental plant community assembly

Despite decades of research, it remains controversial whether ecological communities converge towards a common structure determined by environmental conditions irrespective of assembly history. Here, we show experimentally that the answer depends on the level of community organization considered. In a 9‐year grassland experiment, we manipulated initial plant composition on abandoned arable land and subsequently allowed natural colonization. Initial compositional variation caused plant communities to remain divergent in species identities, even though these same communities converged strongly in species traits. This contrast between species divergence and trait convergence could not be explained by dispersal limitation or community neutrality alone. Our results show that the simultaneous operation of trait‐based assembly rules and species‐level priority effects drives community assembly, making it both deterministic and historically contingent, but at different levels of community organization.     

Functional organization of woody plant assemblages along precipitation and human disturbance gradients in a seasonally dry tropical forest

Chronic anthropogenic disturbances (CAD) and rainfall are important drivers of plant community assembly, but little is known about the role played by inter‐ and intraspecific trait variation as communities respond to these pervasive forces. Here, we examined the hypothesis that lower precipitation and higher CAD reduce both intra‐ and interspecific trait variation in Caatinga dry forests. We sampled woody plants across 15 plots along precipitation and CAD gradients and measured resource‐use traits. The effects of precipitation and CAD on RaoQ functional diversity were decomposed into species turnover and intraspecific variability. We used “T‐statistics” to assess the trait sorting from the regional pool to local communities (i.e., external filtering), and within‐community forces leading to low trait overlap (i.e., internal filtering) at individual and species levels. Intraspecific variability explained at least one‐third of the total trait variation and 46% of variation in multitrait diversity across communities. Increasing disturbance reduced multitrait diversity, while precipitation affected some particular traits, such as wood density. Overall, precipitation determined species sorting across communities, while disturbance relaxed internal filters, leading to higher trait overlap within communities due to higher intraspecific variability. Our results suggest that the woody Caatinga flora contains a substantial amount of both inter‐ and intraspecific trait variation. This variation is not randomly distributed within and across communities, but varies according to rainfall conditions and disturbance intensity. These findings reinforce the emerging idea that human disturbances can reorganize plant communities at multiple scales and highlight trait variability as a key biological asset for the resilience of dry forests.     

A conceptual framework for the spatial analysis of functional trait diversity

Functional trait diversity is a popular tool in modern ecology, mainly used to infer assembly processes and ecosystem functioning. Patterns of functional trait diversity are shaped by ecological processes such as environmental filtering, species interactions and dispersal that are inherently spatial, and different processes may operate at different spatial scales. Adding a spatial dimension to the analysis of functional trait diversity may thus increase our ability to infer community assembly processes and to predict change in assembly processes following disturbance or land‐use change. Richness, evenness and divergence of functional traits are commonly used indices of functional trait diversity that are known to respond differently to large‐scale filters related to environmental heterogeneity and dispersal and fine‐scale filters related to species interactions (competition). Recent developments in spatial statistics make it possible to separately quantify large‐scale patterns (variation in local means) and fine‐scale patterns (variation around local means) by decomposing overall spatial autocorrelation quantified by Moran's coefficient into its positive and negative components using Moran eigenvector maps (MEM). We thus propose to identify the spatial signature of multiple ecological processes that are potentially acting at different spatial scales by contrasting positive and negative components of spatial autocorrelation for each of the three indices of functional trait diversity. We illustrate this approach with a case study from riparian plant communities, where we test the effects of disturbance on spatial patterns of functional trait diversity. The fine‐scale pattern of all three indices was increased in the disturbed versus control habitat, suggesting an increase in local scale competition and an overall increase in unexplained variance in the post‐disturbance versus control community. Further research using simulation modeling should focus on establishing the proposed link between community assembly rules and spatial patterns of functional trait diversity to maximize our ability to infer multiple processes from spatial community structure.     

Seasonality,species richness and poor dispersion mediate intraspecific trait variability in stonefly community responses along an elevational gradient

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On the importance of intraspecific variability for the quantification of functional diversity

Functional diversity (FD) is a key facet of biodiversity used to address central questions in ecology. Despite recent methodological advances, FD remains a complex concept and no consensus has been reached either on how to quantify it, or on how it influences ecological processes. Here we define FD as the distribution of trait values within a community. When and how to account for intraspecific trait variability (ITV) when measuring FD remains one of the main current debates. It remains however unclear to what extent accounting for population‐level ITV would modify FD quantification and associated conclusions. In this paper, we address two critical questions: (1) How sensitive are different components of FD to the inclusion of population‐level ITV? (2) Does the omission of ITV obscure the understanding of ecological patterns? Using a mixture of empirical data and simulation experiments, we conducted a sensitivity analysis of four commonly used FD indices (community weighted mean traits, functional richness, Rao's quadratic entropy, Petchey and Gaston's FD index) and their relationships with environmental gradients and species richness, by varying both the extent (plasticity or not) and the structure (contingency to environmental gradient due to local adaptation) of population‐level ITV. Our results suggest that ITV may strongly alter the quantification of FD and the detection of ecological patterns. Our analysis highlights that 1) species trait values distributions within communities are crucial to the sensitivity to ITV, 2) ITV structure plays a major role in this sensitivity and 3) different indices are not evenly sensitive to ITV, the single‐trait FD from Petchey and Gaston being the most sensitive among the four metrics tested. We conclude that the effects of intraspecific variability in trait values should be more systematically tested before drawing central conclusions on FD, and suggest the use of simulation studies for such sensitivity analyses.     

Past is prologue: host community assembly and the risk of infectious disease over time

Infectious disease risk is often influenced by host diversity, but the causes are unresolved. Changes in diversity are associated with changes in community structure, particularly during community assembly; therefore, by incorporating change over time, host community assembly may provide a framework to resolve causation. In turn, community assembly can be driven by many processes, including resource enrichment. To test the hypothesis that community assembly causally links host diversity to future disease, we experimentally manipulated host diversity and resource supply to hosts, then allowed communities to assemble for 2 years (surveyed 2012–2014). Initially, host diversity increased disease. Subsequently, host diversity did not directly alter disease. However, host diversity determined the trajectory of host community assembly, altering colonisation by exotic host species and richness‐independent host phylogenetic diversity, which together reversed the initial increase in disease. Ultimately, incorporating the temporal dimension of community assembly revealed novel mechanisms linking host diversity to future disease.     

Community assembly in Lake Tanganyika cichlid fish: quantifying the contributions of both niche‐based and neutral processes

The cichlid family features some of the most spectacular examples of adaptive radiation. Evolutionary studies have highlighted the importance of both trophic adaptation and sexual selection in cichlid speciation. However, it is poorly understood what processes drive the composition and diversity of local cichlid species assemblages on relatively short, ecological timescales. Here, we investigate the relative importance of niche‐based and neutral processes in determining the composition and diversity of cichlid communities inhabiting various environmental conditions in the littoral zone of Lake Tanganyika, Zambia. We collected data on cichlid abundance, morphometrics, and local environments. We analyzed relationships between mean trait values, community composition, and environmental variation, and used a recently developed modeling technique (STEPCAM) to estimate the contributions of niche‐based and neutral processes to community assembly. Contrary to our expectations, our results show that stochastic processes, and not niche‐based processes, were responsible for the majority of cichlid community assembly. We also found that the relative importance of niche‐based and neutral processes was constant across environments. However, we found significant relationships between environmental variation, community trait means, and community composition. These relationships were caused by niche‐based processes, as they disappeared in simulated, purely neutrally assembled communities. Importantly, these results can potentially reconcile seemingly contrasting findings in the literature about the importance of either niche‐based or neutral‐based processes in community assembly, as we show that significant trait relationships can already be found in nearly (but not completely) neutrally assembled communities; that is, even a small deviation from neutrality can have major effects on community patterns.     

Beta‐diversity of central European forests decreases along an elevational gradient due to the variation in local community assembly processes

Beta‐diversity has been repeatedly shown to decline with increasing elevation, but the causes of this pattern remain unclear, partly because they are confounded by coincident variation in alpha‐ and gamma‐diversity. We used 8795 forest vegetation‐plot records from the Czech National Phytosociological Database to compare the observed patterns of beta diversity to null‐model expectations (beta‐deviation) controlling for the effects of alpha‐ and gamma‐diversity. We tested whether β‐diversity patterns along a 1200 m elevation gradient exclusively depend on the effect of varying species pool size, or also on the variation of the magnitude of community assembly mechanisms determining the distribution of species across communities (e.g. environmental filtering, dispersal limitation). The null model we used is a novel extension of an existing null‐model designed for presence/absence data and was specifically designed to disrupt the effect of community assembly mechanisms, while retaining some key features of observed communities such as average species richness and species abundance distribution. Analyses were replicated in ten subregions with comparable elevation ranges. Beta‐diversity declined along the elevation gradient due to a decrease in gamma‐diversity, which was steeper than the decrease in alpha‐diversity. This pattern persisted after controlling for alpha‐ and gamma‐diversity variation, and the results were robust when different resampling schemes and diversity metrics were used. We conclude that in temperate forests the pattern of decreasing beta‐diversity with elevation does not exclusively depend on variation in species pool size, as has been hypothesized, but also on variation in community assembly mechanisms. The results were consistent across resampling schemes and diversity measures, thus supporting the use of vegetation‐plot databases for understanding patterns of beta‐diversity at the regional scale.     

Individualistic responses of forest herb traits to environmental change

• Intraspecific trait variation (ITV; i.e. variability in mean and/or distribution of plant attribute values within species) can occur in response to multiple drivers. Environmental change and land‐use legacies could directly alter trait values within species but could also affect them indirectly through changes in vegetation cover. Increasing variability in environmental conditions could lead to more ITV, but responses might differ among species. Disentangling these drivers on ITV is necessary to accurately predict plant community responses to global change.
• We planted herb communities into forest soils with and without a recent history of agriculture. Soils were collected across temperate European regions, while the 15 selected herb species had different colonizing abilities and affinities to forest habitat. These mesocosms (384) were exposed to two‐level full‐factorial treatments of warming, nitrogen addition and illumination. We measured plant height and specific leaf area (SLA).
• For the majority of species, mean plant height increased as vegetation cover increased in response to light addition, warming and agricultural legacy. The coefficient of variation (CV) for height was larger in fast‐colonizing species. Mean SLA for vernal species increased with warming, while light addition generally decreased mean SLA for shade‐tolerant species. Interactions between treatments were not important predictors.
• Environmental change treatments influenced ITV, either via increasing vegetation cover or by affecting trait values directly. Species’ ITV was individualistic, i.e. species responded to different single resource and condition manipulations that benefited their growth in the short term. These individual responses could be important for altered community organization after a prolonged period.

     

Incorporating intraspecific variation in tests of trait-based community assembly

Environmental filtering and niche differentiation are processes proposed to drive community assembly, generating nonrandom patterns in community trait distributions. Despite the substantial intraspecific trait variation present in plant communities, most previous studies of trait-based community assembly have used species mean trait values and therefore not accounted for intraspecific variation. Using a null model approach, I tested for environmental filtering and niche differentiation acting on three key functional traits??vegetative height, specific leaf area (SLA), and leaf dry matter content (LDMC)??in old-field plant communities. I also examined how accounting for intraspecific variation at the among-plot and individual levels affected the detection of nonrandom assembly patterns. Tests using fixed species mean trait values provided evidence of environmental filtering acting on height and SLA and niche differentiation acting on SLA. Including plot-level intraspecific variation increased the strength of these patterns, indicating an important role of intraspecific variation in community assembly. Tests using individual trait data indicated strong environmental filtering acting on all traits, but provided no evidence of niche differentiation, although these signals may have been obscured by the effects of dispersal limitation and spatial aggregation of conspecific individuals. There was also strong evidence of nonrandom assembly of individuals within single species, with the strength of environmental filtering varying among species. This study demonstrates that, while analyses using fixed species mean trait values can provide insights into community assembly processes, accounting for intraspecific variation provides a more complete view of communities and the processes driving their assembly.     

Gap pattern and colonization opportunities in plant communities: effects of species richness, mortality, and spatial aggregation

We investigate how perturbations that induce mortality transform original spatial patterns in plant communities into binary spatial patterns of survivors and perished individuals. By means of computer simulation, we analyse effects of average mortality, interspecific variation of mortality around the mean, spatial distribution of the species (clumping degree), and species richness. Gap spatial pattern is quantified by four spatial indices or landscape metrics (gap area, density, shape and coherence). In single‐species communities, the emerging gap patterns are subject to critical phenomena: opportunities for colonizers to establish increase with mortality, but more rapidly at specific mortality thresholds. In multi‐species communities, neither species richness nor interspecific variation of mortality influences gap spatial pattern when community assembly is random. Colonization opportunities would therefore not be affected by local species extinction in such a system, nor by the presence of species with divergent sensitivities to perturbation. In a community that is highly spatially aggregated, increases in interspecific mortality variation shift the pattern towards fewer gaps that are larger and more isodiametric, which suggests increased establishment chances for colonizers. Similar changes are induced in communities characterized by large interspecific mortality differences if clumping degree is increased. Loss of species richness only modifies gap spatial pattern to a substantial extent if mortality variation is high: in this case, depauperate communities exhibit a wider variety of colonization opportunities (more gaps which are on average smaller, but the largest gap is larger) than species‐richer ones. These findings may explain the contrast between the negative diversity‐invasibility relationship often found in small‐scale experimental studies and the positive diversity‐invasibility relationship found in observational studies at larger scale. They also demonstrate that the pre‐disturbance spatial structure of a community significantly affects colonization opportunities for alien species, and is therefore a likely determinant of the trajectory of secondary succession following perturbation.     

Partitioning of functional diversity reveals the scale and extent of trait convergence and divergence

Questions: Trait differentiation among species occurs at different spatial scales within a region. How does the partitioning of functional diversity help to identify different community assembly mechanisms? Location: Northeastern Spain. Methods: Functional diversity can be partitioned into within‐community (α) and among‐communities (β) components, in analogy to Whittaker's classical α and β species diversity concept. In light of ecological null models, we test and discuss two algorithms as a framework to measure α and β functional diversity (the Rao quadratic entropy index and the variance of trait values). Species and trait (specific leaf area) data from pastures under different climatic conditions in NE Spain are used as a case study. Results: The proposed indices show different mathematical properties but similarly account for the spatial components of functional diversity. For all vegetation types along the climatic gradient, the observed α functional diversity was lower than expected at random, an observation consistent with the hypothesis of trait convergence resulting from habitat filtering. On the other hand, our data exhibited a remarkably higher functional diversity within communities compared to among communities (α?β). In contrast to the high species turnover, there was a limited functional diversity turnover among communities, and a large part of the trait divergence occurred among coexisting species. Conclusions: Partitioning functional diversity within and among communities revealed that both trait convergence and divergence occur in the formation of assemblages from the local species pool. A considerable trait convergence exists at the regional scale in spite of changes in species composition, suggesting the existence of ecological redundancy among communities.     

A functional trait-based approach to understand community assembly and diversity–productivity relationships over 7 years in experimental grasslands

Several multi-year biodiversity experiments have shown positive species richness–productivity relationships which strengthen over time, but the mechanisms which control productivity are not well understood. We used experimental grasslands (Jena Experiment) with mixtures containing different numbers of species (4, 8, 16 and 60) and plant functional groups (1–4; grasses, legumes, small herbs, tall herbs) to explore patterns of variation in functional trait composition as well as climatic variables as predictors for community biomass production across several years (from 2003 to 2009). Over this time span, high community mean trait values shifted from the dominance of trait values associated with fast growth to trait values suggesting a conservation of growth-related resources and successful reproduction. Increasing between-community convergence in means of several productivity-related traits indicated that environmental filtering and exclusion of competitively weaker species played a role during community assembly. A general trend for increasing functional trait diversity within and convergence among communities suggested niche differentiation through limiting similarity in the longer term and that similar mechanisms operated in communities sown with different diversity. Community biomass production was primarily explained by a few key mean traits (tall growth, large seed mass and leaf nitrogen concentration) and to a smaller extent by functional diversity in nitrogen acquisition strategies, functional richness in multiple traits and functional evenness in light-acquisition traits. Increasing species richness, presence of an exceptionally productive legume species (Onobrychis viciifolia) and climatic variables explained an additional proportion of variation in community biomass. In general, community biomass production decreased through time, but communities with higher functional richness in multiple traits had high productivities over several years. Our results suggest that assembly processes within communities with an artificially maintained species composition maximize functional diversity through niche differentiation and exclusion of weaker competitors, thereby maintaining their potential for high productivity.     

Light and warming drive forest understorey community development in different environments

Plant community composition and functional traits respond to chronic drivers such as climate change and nitrogen (N) deposition. In contrast, pulse disturbances from ecosystem management can additionally change resources and conditions. Community responses to combined environmental changes may further depend on land‐use legacies. Disentangling the relative importance of these global change drivers is necessary to improve predictions of future plant communities. We performed a multifactor global change experiment to disentangle drivers of herbaceous plant community trajectories in a temperate deciduous forest. Communities of five species, assembled from a pool of 15 forest herb species with varying ecological strategies, were grown in 384 mesocosms on soils from ancient forest (forested at least since 1850) and postagricultural forest (forested since 1950) collected across Europe. Mesocosms were exposed to two‐level full‐factorial treatments of warming, light addition (representing changing forest management) and N enrichment. We measured plant height, specific leaf area (SLA) and species cover over the course of three growing seasons. Increasing light availability followed by warming reordered the species towards a taller herb community, with limited effects of N enrichment or the forest land‐use history. Two‐way interactions between treatments and incorporating intraspecific trait variation (ITV) did not yield additional inference on community height change. Contrastingly, community SLA differed when considering ITV along with species reordering, which highlights ITV’s importance for understanding leaf morphology responses to nutrient enrichment in dark conditions. Contrary to our expectations, we found limited evidence of land‐use legacies affecting community responses to environmental changes, perhaps because dispersal limitation was removed in the experimental design. These findings can improve predictions of community functional trait responses to global changes by acknowledging ITV, and subtle changes in light availability. Adaptive forest management to impending global change could benefit the restoration and conservation of understorey plant communities by reducing the light availability.     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.     

Underdispersion and overdispersion of traits in terrestrial snail communities on islands

Understanding and disentangling different processes underlying the assembly and diversity of communities remains a key challenge in ecology. Species can assemble into communities either randomly or due to deterministic processes. Deterministic assembly leads to species being more similar (underdispersed) or more different (overdispersed) in certain traits than would be expected by chance. However, the relative importance of those processes is not well understood for many organisms, including terrestrial invertebrates. Based on knowledge of a broad range of species traits, we tested for the presence of trait underdispersion (indicating dispersal or environmental filtering) and trait overdispersion (indicating niche partitioning) and their relative importance in explaining land snail community composition on lake islands. The analysis of community assembly was performed using a functional diversity index (Rao's quadratic entropy) in combination with a null model approach. Regression analysis with the effect sizes of the assembly tests and environmental variables gave information on the strength of under‐ and overdispersion along environmental gradients. Additionally, we examined the link between community weighted mean trait values and environmental variables using a CWM‐RDA. We found both trait underdispersion and trait overdispersion, but underdispersion (eight traits) was more frequently detected than overdispersion (two traits). Underdispersion was related to four environmental variables (tree cover, habitat diversity, productivity of ground vegetation, and location on an esker ridge). Our results show clear evidence for underdispersion in traits driven by environmental filtering, but no clear evidence for dispersal filtering. We did not find evidence for overdispersion of traits due to diet or body size, but overdispersion in shell shape may indicate niche differentiation between snail species driven by small‐scale habitat heterogeneity. The use of species traits enabled us to identify key traits involved in snail community assembly and to detect the simultaneous occurrence of trait underdispersion and overdispersion.     

The interplay among intraspecific leaf trait variation,niche breadth and species abundance along light and soil nutrient gradients

It is assumed that widespread, generalist species have high phenotypic variation, but we know little about how intraspecific trait variation (ITV) relates to species abundance and niche breadth. In the temperate rainforest of southern Chile, we hypothesized that species with wide niche breadth would exhibit 1) high among‐plot ITV, 2) a strong relationship between trait values and the environment, and 3) a close fit between traits and local environment trait optima. We measured leaf functional traits (leaf area, LMA, leaf N and P concentrations) of saplings in woody species, and compared the relative abundance of each species with its niche breadth, measured as the range of light, soil N and P availability. We used the slope of the linear regression of species’ trait–environment relationships to assess the strength and direction of these relationships, and measured the degree to which species’ trait values track the environmental optimum across plots. In some cases, species having wide niche breadth had high ITV in leaf N and also matched traits (LMA and leaf P) to local optima along the light gradient; they also had high ITV in general and matched leaf P to local optima along the soil P gradient. The relationship between species with wide niche breadth and the strength of intraspecific trait–environment relationships was generally weak and varied depending on the niche dimension and trait in question. Species varied considerably in the strength of trait–environment relationships and total magnitude of ITV, and this variation was not generally strongly related to species abundances or niche breadth patterns. In conclusion, trait variation at the community level is not driven by a few abundant, widely distributed species, but depends on the aggregate trait responses of both abundant and rare species. This makes it difficult to scale individual species trait responses up to the community level.