Scaling environmental change through the community-level: a trait-based response-and-effect framework for plants

Predicting ecosystem responses to global change is a major challenge in ecology. A critical step in that challenge is to understand how changing environmental conditions influence processes across levels of ecological organization. While direct scaling from individual to ecosystem dynamics can lead to robust and mechanistic predictions, new approaches are needed to appropriately translate questions through the community level. Species invasion, loss, and turnover all necessitate this scaling through community processes, but predicting how such changes may influence ecosystem function is notoriously difficult. We suggest that community‐level dynamics can be incorporated into scaling predictions using a trait‐based response–effect framework that differentiates the community response to environmental change (predicted by response traits) and the effect of that change on ecosystem processes (predicted by effect traits). We develop a response‐and‐effect functional framework, concentrating on how the relationships among species' response, effect, and abundance can lead to general predictions concerning the magnitude and direction of the influence of environmental change on function. We then detail several key research directions needed to better scale the effects of environmental change through the community level. These include (1) effect and response trait characterization, (2) linkages between response‐and‐effect traits, (3) the importance of species interactions on trait expression, and (4) incorporation of feedbacks across multiple temporal scales. Increasing rates of extinction and invasion that are modifying communities worldwide make such a research agenda imperative.     

Plant functional trait responses to grassland succession over 25 years

Abstract. Many grasslands and arable fields have been abandoned in Europe in the last decades. So far, however, ecologists have not been able to establish one general, unifying theory for successionally induced changes in species composition following abandonment. In fact, the course of succession seems to be unique for each site and year. Here we focused on the analysis of plant functional traits in order to detect general trends in trait responses to succession which would prove independent of site characteristics. We studied permanent plot series of 14 grassland sites in SW Germany that had been abandoned for 25 yr. Species composition as well as the course of succession varied significantly among the grasslands. Response to succession was analysed for the following traits: ‘plant height’, ‘canopy structure’, ‘specific leaf area’ (SLA), ‘storage organ’, ‘vegetative spread’, ‘plant persistence’, ‘seed bank longevity’, ‘start of flowering’, ‘duration of flowering’ and ‘seed mass’. We compared a univariate with a multivariate approach. In the univariate approach, attributes of each trait were analysed separately employing GLM whereas in the multivariate approach all traits were handled together in NMS. Both analyses indicated similar trait responses to succession. There was a significant increase in the proportion of species characterized by the following attributes: plant height > 0.6 m, leaves distributed regularly along the stem, vegetative spread > 100 mm, start of flowering later than May, duration of flowering 1–2 months and seeds of either low or high mass.     

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.     

Response of ant functional composition to fire

Little is known about the impact of disturbances on functional diversity and the long‐term provisioning of ecosystem services, especially in animals. In this work we analyze the effect of wildfire on the functional composition of Mediterranean ant communities. In particular, we asked whether a) fire changes functional composition (mean and dissimilarity of trait values) at the community level; and b) such fire‐induced functional modification is driven by changes in the relative abundance‐dominance of species or by a replacement of species with different traits. We sampled ant communities in burned and unburned plots along 22 sites in a western Mediterranean region, and we computed two complementary functional trait composition indices (‘trait average’ and ‘trait dissimilarity’) for 12 functional traits (related to resource exploitation, social structure and reproduction) and with two different datasets varying in the way species abundance is considered (i.e. abundance and occurrence data). Our results suggest a set of functional responses that seem to be related to direct mortality by fire as well as to indirect fire‐induced modifications in environmental conditions relevant for ants. Trait average of colony size, worker size, worker polymorphism and the ratio between queen and worker size, as well as the trait dissimilarity of the proportion of behaviorally dominant species and of liquid food consumption, and overall functional diversity, were higher in burned than in unburned areas. Interestingly, different patterns arise when comparing results from abundance and occurrence data. While the response to fire in trait averages is quite similar, in the case of trait dissimilarity, the higher values in response to fire are much more marked when considering occurrence rather than abundance data. Our results suggest that changes in trait average are driven at the same time by replacement of species with different traits and by changes in the relative abundance‐dominance of species, while fire promotes a higher diversity of functions that is primarily driven by rare species that are functionally unique. Overall, we observed major fire‐induced changes in functional composition in Mediterranean ant communities that might have relevant consequences for ecosystem processes and services.     

BryForTrait – A life‐history trait database of forest bryophytes

In ecological research, plant functional trait analyses are widely applied to understand to what extent the inter‐specific variation in trait attributes has an adaptive value or to predict ecosystem processes and changes. Compared to vascular plants, trait studies using bryophytes are scarce, which is likely due to missing trait information for bryophyte species. With the BryForTrait database, we want to reduce this deficit. Our database represents a compilation of autecological information and morphological and regenerative trait data on different stages of the life cycle of bryophytes occurring in forest ecosystems. The database contains information for 35 traits and 721 Central European bryophyte species; in total more than 23,000 trait values. The BryForTrait database will enable future trait studies, providing new insights into bryophyte‐dominated ecosystems.     

Environmentally and behaviourally mediated co‐occurrence of functional traits in bird communities of tropical forest fragments

Two major theories of community assembly – based on the assumption of ‘limiting similarity’ or ‘habitat filtering’, respectively – predict contrasting patterns in the spatial arrangement of functional traits. Previous analyses have made progress in testing these predictions and identifying underlying processes, but have also pointed to theoretical as well as methodological shortcomings. Here we applied a recently developed methodology for spatially explicit analysis of phylogenetic meta‐community structure to study the pattern of co‐occurrence of functional traits in Afrotropical and Neotropical bird species inhabiting forest fragments. Focusing separately on locomotory, dietary, and dispersal traits, we tested whether environmental filtering causes spatial clustering, or competition leads to spatial segregation as predicted by limiting similarity theory. We detected significant segregation of species co‐occurrences in African fragments, but not in the Neotropical ones. Interspecific competition had a higher impact on trait co‐occurrence than filter effects, yet no single functional trait was able to explain the observed degree of spatial segregation among species. Despite high regional variability spanning from spatial segregation to aggregation, we found a consistent tendency for a clustered spatial patterning of functional traits among communities in fragmented landscapes, particularly in non‐territorial species. Overall, we show that behavioural effects, such as territoriality, and environmental effects, such as the area of forest remnants or properties of the landscape matrix in which they are embedded, can strongly affect the pattern of trait co‐occurrence. Our findings suggest that trait‐based analyses of community structure should include behavioural and environmental covariates, and we here provide an appropriate method for linking functional traits, species ecology and environmental conditions to clarify the drivers underlying spatial patterns of species co‐occurrence.     

A functional assessment of the response of grassland vegetation to reduced grazing and abandonment

Question: Predicting the impact of land‐use change on vegetation is vital to understanding how biodiversity and ecosystem function may respond. Is it correct to assume that abandonment is an extreme form of grazing reduction? Location: Borders and central Scotland. Methods: The analysis used data sets from two identical experiments where the impacts of two unfertilized, extensively grazed treatments and one unfertilized abandoned treatment were compared against the species dynamics of a pasture subject to normal, productive grazing management over a 16‐year period. Initial multivariate analysis using Principal Response Curves was used to assess if particular traits were associated with either extensive or abandoned treatments, and was checked using univariate tests of individual traits. RLQ analysis followed by clustering into response groups was used to assess if species behaved in a similar manner between sites. Results: For many traits/attributes the shift in value or proportion was approximately linear across the extensification treatments as grazing was removed. However, certain traits showed step changes and quadratic responses. Leaf dry matter content, an important effect trait, was in the latter group. Most traits/attributes and species behaved similarly at the two sites. However, traits such as regenerative strategy, seed length, longevity and mass and seed bank type behaved differently, indicating that they are not predictable response traits. Conclusion: The results indicate that responses to grazing removal during extensification are largely straightforward and largely independent of species pool. However, there are discrepancies that suggest that simple analyses of the impacts of land‐use changes such as grazing reduction may hide more complex responses.     

Differential effects of functional traits on aboveground biomass in semi-natural grasslands

Despite increasing evidence on the importance of species functional characteristics for ecosystem processes, two major hypotheses suggest different mechanisms: the ‘mass ratio hypothesis’ assumes that functional traits of the dominant species determine ecosystem processes, while the ‘complementarity hypothesis’ predicts that resource niches may be used more completely when a community is functionally more diverse. Here, we present a method which uses two different groups of biotic predictor variables being (1) abundance‐weighted mean (=aggregated) trait values and (2) functional trait diversity based on Rao's quadratic diversity (FD_Q) to test the competing hypotheses on biodiversity–ecosystem functioning relationships after accounting for co‐varying abiotic factors. We applied this method to data recorded on biodiversity–biomass relationships and environmental variables in 35 semi‐natural temperate grasslands and used a literature‐based matrix of fourteen plant functional traits to assess the explanatory power of models including different sets of predictor variables. Aboveground community biomass did not correlate with species richness. Abiotic factors, in particular soil nitrogen concentration, explained about 50% of variability in aboveground biomass. The best model incorporating functional trait diversity explained only about 30%, while the best model based on aggregated trait values explained about 54% of variability in aboveground biomass. The inclusion of all predictor variable groups in a combined model increased the predictive power to about 75%. This model comprised soil nitrogen concentration as abiotic factor, aggregated traits being indicative for species competitive dominance (rooting depth, leaf distribution, specific leaf area, perennial life cycle) and functional trait diversity in vegetative plant height, leaf area and life cycle. Our study strongly suggests that abiotic factors, trait values of the dominant species and functional trait diversity in combination may best explain differences in aboveground community biomass in natural ecosystems and that their isolated consideration may be misleading.     

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.     

Contrasting effects of plant inter‐ and intraspecific variation on community trait responses to restoration of a sandy grassland ecosystem

Changes in plant community traits along an environmental gradient are caused by interspecific and intraspecific trait variation. However, little is known about the role of interspecific and intraspecific trait variation in plant community responses to the restoration of a sandy grassland ecosystem. We measured five functional traits of 34 species along a restoration gradient of sandy grassland (mobile dune, semi‐fixed dune, fixed dune, and grassland) in Horqin Sand Land, northern China. We examined how community‐level traits varied with habitat changes and soil gradients using both abundance‐weighted and non‐weighted averages of trait values. We quantified the relative contribution of inter‐ and intraspecific trait variation in specific leaf area (SLA), leaf dry matter content (LDMC), leaf carbon content (LCC), leaf nitrogen content (LNC), and plant height to the community response to habitat changes in the restoration of sandy grassland. We found that five weighted community‐average traits varied significantly with habitat changes. Along the soil gradient in the restoration of sandy grassland, plant height, SLA, LDMC, and LCC increased, while LNC decreased. For all traits, there was a greater contribution of interspecific variation to community response in regard to habitat changes relative to that of intraspecific variation. The relative contribution of the interspecific variation effect of an abundance‐weighted trait was greater than that of a non‐weighted trait with regard to all traits except LDMC. A community‐level trait response to habitat changes was due largely to species turnover. Though the intraspecific shift plays a small role in community trait response to habitat changes, it has an effect on plant coexistence and the maintenance of herbaceous plants in sandy grassland habitats. The context dependency of positive and negative covariation between inter‐ and intraspecific variation further suggests that both effects of inter‐ and intraspecific variation on a community trait should be considered when understanding a plant community response to environmental changes in sandy grassland ecosystems.     

Intra-Specific Leaf Trait Variation: Management and Fertility Matter More than the Climate at Continental Scales

Plant functional traits have contributed to the understanding of how vegetation responds to management, community assembly and how vegetation controls ecosystem processes. As traits are time consuming to measure – and it may not always be feasible or practical to measure all traits within a study – great reliance has been put on using trait values from databases. This ignores intraspecific trait variability and the traits’ responses to the environment. This study uses trait values measured as part of a large-scale investigation of land use impacts on vegetation to assess how environmental factors, specifically climate, soil and management, control inter-population level intra-specific variation in key traits. There was clear evidence that intra-specific variation in leaf carbon content, leaf dry matter content, leaf nitrogen content and specific leaf area were all sensitive to edaphic factors and to management, but only leaf dry matter content was linearly affected by the climate (rainfall). There were also significant interactions between climate and species identity for leaf dry matter content and leaf nitrogen content, suggesting that species responses to the climate are not uniform or simple. The results of this study suggest that site fertility and management ought to be associated with trait data in databases to allow for the incorporation of intra-specific variation in analyses and that more research is needed to identify the shape of trait:climate relationships.     

Trait‐based tests of coexistence mechanisms

Recent functional trait studies have shown that trait differences may favour certain species (environmental filtering) while simultaneously preventing competitive exclusion (niche partitioning). However, phenomenological trait‐dispersion analyses do not identify the mechanisms that generate niche partitioning, preventing trait‐based prediction of future changes in biodiversity. We argue that such predictions require linking functional traits with recognised coexistence mechanisms involving spatial or temporal environmental heterogeneity, resource partitioning and natural enemies. We first demonstrate the limitations of phenomenological approaches using simulations, and then (1) propose trait‐based tests of coexistence, (2) generate hypotheses about which plant functional traits are likely to interact with particular mechanisms and (3) review the literature for evidence for these hypotheses. Theory and data suggest that all four classes of coexistence mechanisms could act on functional trait variation, but some mechanisms will be stronger and more widespread than others. The highest priority for future research is studies of interactions between environmental heterogeneity and trait variation that measure environmental variables at within‐community scales and quantify species' responses to the environment in the absence of competition. Evidence that similar trait‐based coexistence mechanisms operate in many ecosystems would simplify biodiversity forecasting and represent a rare victory for generality over contingency in community ecology.     

Using phylogeny and functional traits for assessing community assembly along environmental gradients: A deterministic process driven by elevation

Community assembly processes is the primary focus of community ecology. Using phylogenetic‐based and functional trait‐based methods jointly to explore these processes along environmental gradients are useful ways to explain the change of assembly mechanisms under changing world. Our study combined these methods to test assembly processes in wide range gradients of elevation and other habitat environmental factors. We collected our data at 40 plots in Taibai Mountain, China, with more than 2,300 m altitude difference in study area and then measured traits and environmental factors. Variance partitioning was used to distinguish the main environment factors leading to phylogeny and traits change among 40 plots. Principal component analysis (PCA) was applied to colligate other environment factors. Community assembly patterns along environmental gradients based on phylogenetic and functional methods were studied for exploring assembly mechanisms. Phylogenetic signal was calculated for each community along environmental gradients in order to detect the variation of trait performance on phylogeny. Elevation showed a better explanatory power than other environment factors for phylogenetic and most traits’ variance. Phylogenetic and several functional structure clustered at high elevation while some conserved traits overdispersed. Convergent tendency which might be caused by filtering or competition along elevation was detected based on functional traits. Leaf dry matter content (LDMC) and leaf nitrogen content along PCA 1 axis showed conflicting patterns comparing to patterns showed on elevation. LDMC exhibited the strongest phylogenetic signal. Only the phylogenetic signal of maximum plant height showed explicable change along environmental gradients. Synthesis. Elevation is the best environment factors for predicting phylogeny and traits change. Plant's phylogenetic and some functional structures show environmental filtering in alpine region while it shows different assembly processes in middle‐ and low‐altitude region by different trait/phylogeny. The results highlight deterministic processes dominate community assembly in large‐scale environmental gradients. Performance of phylogeny and traits along gradients may be independent with each other. The novel method for calculating functional structure which we used in this study and the focus of phylogenetic signal change along gradients may provide more useful ways to detect community assembly mechanisms.     

Going beyond limitations of plant functional types when predicting global ecosystem–atmosphere fluxes: exploring the merits of traits‐based approaches

Aim Despite their importance for predicting fluxes to and from terrestrial ecosystems, dynamic global vegetation models have insufficient realism because of their use of plant functional types (PFTs) with constant attributes. Based on recent advances in community ecology, we explore the merits of a traits‐based vegetation model to deal with current shortcomings. Location Global. Methods A research review of current concepts and information, providing a new perspective, supported by quantitative analysis of a global traits database. Results Continuous and process‐based trait–environment relations are central to a traits‐based approach and allow us to directly calculate fluxes based on functional characteristics. By quantifying community assembly concepts, it is possible to predict trait values from environmental drivers, although these relations are still imperfect. Through the quantification of these relations, effects of adaptation and species replacement upon environmental changes are implicitly accounted for. Such functional links also allow direct calculation of fluxes, including those related to feedbacks through the nitrogen and water cycle. Finally, a traits‐based model allows the prediction of new trait combinations and no‐analogue ecosystem functions projected to arise in the near future, which is not feasible in current vegetation models. A separate calculation of ecosystem fluxes and PFT occurrences in traits‐based models allows for flexible vegetation classifications. Main conclusions Given the advantages described above, we argue that traits‐based modelling deserves consideration (although it will not be easy) if one is to aim for better climate projections.     

Genetic structure among arable populations of Capsella bursa‐pastoris is linked to functional traits and in‐field conditions

Wild arable plants can be an economic burden but they also support diverse arable food webs and contribute to valuable ecosystem functions. These benefits may have been compromised over recent decades by declining weed diversity. The decline in wild arable plant diversity has been viewed predominantly in terms of species shifts a view that ignores the genetic and functional variation existing within species and the impact on ecological and evolutionary processes which this may have. To examine within‐species diversity, ISSR markers were used in parallel with environmental and phenotypic characterisation, to investigate the population structure and diversity of Capsella bursa‐pastoris (shepherd's purse) from arable fields in the UK. Analysis of 338 ISSR products for 109 individuals from 51 accessions obtained from the seed banks of 33 arable fields showed that in‐field populations of shepherd's purse were genetically differentiated between individuals, and among accessions and fields. In addition, cluster analysis identified three genetically distinct regional‐scale populations. Phenotypic variation was present at all scales of genetic differentiation, including the regional scale where populations differed in their key life‐history traits: flowering time, fecundity and dormancy. Genetic drift is proposed as a contributor to differentiation among genetically isolated but locally co‐occurring accessions. In addition, the genetic and phenotypic variation in shepherd's purse exhibited large scale, spatial trends and showed statistically significant associations with cropping intensity and soil‐pH. These results suggest that adaptation as a result of selection by cropping practise and soil‐pH has played a role in the ability of shepherd's purse to colonise and persist in arable fields.     

Functional diversity in nematode communities across terrestrial ecosystems

Functional diversity can be defined as the distribution of trait values within a community. Hence, functional diversity can be an indicator of habitat filtering and a reliable environmental predictor of ecosystem functioning. However, there is a serious lack of studies that test how functional diversity indices change depending on the environmental conditions. The aim of this study is to provide such evidence by analyzing the distribution and variation of continuous body-mass values (i.e. functional diversity) and related shifts in body length and width in a nematode community.We used a large online dataset on nematode traits to analyze: (i) the distribution of body mass using three functional diversity indices, i.e. functional richness, functional divergence and functional evenness; (ii) the shifts in body-size traits (length and width); and (iii) the body-mass distributions of five trophic groups and of the entire nematode community.Managed grasslands exhibited the widest range of body-mass values while body-mass distribution in arable fields covered the greatest area in comparison to the other ecosystem types. The shift in body size revealed environmental filters that could not have been identified by the study of functional diversity indices per se. We found low values of functional evenness to be associated with high values of functional richness. We provide novel empirical evidence that body-mass distribution within a trophic group mirrors the effects of habitat filtering more than the distribution in the community as a whole. Hence, our trait-based approach, more than functional diversity itself, disclosed soil food-web structure and identified community responses.     

Functional diversity: a tool for answering challenging ecological questions

Functional trait diversity provides a powerful means of addressing ecology's persistent questions, through its dual role as an indicator of mechanisms driving differences in species composition between communities and as a predictor of ecosystem‐level processes. Functional traits provide a means of testing mechanisms behind species turnover between communities because environmental heterogeneity, competition and disturbance influence species fitness via their traits. Functional traits also provide a link between species and multiple ecosystem‐level processes, such as primary productivity, nutrient fluxes and resilience, since species influence these processes via their traits. This special issue demonstrates that functional diversity offers a practical means of investigating ecology's persistent questions.     

Land‐use effects on the functional distinctness of arthropod communities

Land‐use change is a major driver of the global loss of biodiversity, but it is unclear to what extent this also results in a loss of ecological traits. Therefore, a better understanding of how land‐use change affects ecological traits is crucial for efforts to sustain functional diversity. To this end we tested whether higher species richness or taxonomic distinctness generally leads to increased functional distinctness and whether intensive land use leads to functionally more narrow arthropod communities. We compiled species composition and trait data for 350 species of terrestrial arthropods (Araneae, Carabidae and Heteroptera) in different land‐use types (forests, grasslands and arable fields) of low and high land‐use intensity. We calculated the average functional and taxonomic distinctness and the rarified trait richness for each community. These measures reflect the range of traits, taxonomic relatedness and number of traits that are observed in local communities. Average functional distinctness only increased significantly with species richness in Carabidae communities. Functional distinctness increased significantly with taxonomic distinctness in communities of all analyzed taxa suggesting a high functional redundancy of taxonomically closely related species. Araneae and Heteroptera communities had the expected lower functional distinctness at sites with higher land‐use intensity. More frequently disturbed land‐use types such as managed grasslands or arable fields were characterized by species with smaller body sizes and higher dispersal abilities and communities with lower functional distinctness or trait richness. Simple recommendations about the conservation of functional distinctness of arthropod communities in the face of future land‐use intensification and species loss are not possible. Our study shows that these relationships depend on the studied taxa and land‐use type. However, for some arthropod groups functional distinctness is threatened by intensification and conversion from less to more frequently disturbed land‐uses.