Leaf trait integration mediates species richness variation in a species-rich neotropical forest domain

Environmental gradients are known to drive changes in mean trait values, but changes in the trait integration strength across local communities are less well understood, particularly with regard to possible links with species richness variation. Here, we tested if climate, soil, and topography gradients drive species richness indirectly via constraints on trait integration in the Atlantic Forest of South America. We evaluated seven traits (from leaf, wood, seed, and plant size) of 1456 species occurring across 84 local communities. Generalized least square models and a path model were applied to test direct and indirect relationships. Correlations were higher between leaf traits (average r?=?0.28) and lower when other traits were included (average r?=?0.16). In line with this result, species richness was related to a multivariate index of interspecific trait integration (ITI) computed for leaf traits, but not to the ITI for all the seven traits. Abiotic gradients influenced species richness both directly and indirectly through the leaf trait integration. A total of 33% and 26% of the variation in species richness and ITI, respectively, were explained by the models, with climatic conditions showing higher contribution than topographic and edaphic factors. These results support a significant but reduced environmental selection role behind the trait-based community assembly and may suggest that other processes are involved in the constrain of trait integration at larger spatial scales. In addition, different directional trends in trait–trait relationships across local communities suggest that global trait relationships may not necessarily hold at local contexts.

     

Contrasting patterns of trait‐based community assembly in lianas and trees from temperate Australia

Trait variation in plant communities is thought to be constrained by two opposing community assembly processes operating at discrete spatial scales: habitat filtering and limiting similarity between coexisting species. Filtering processes cause convergence in ecological strategy as species are excluded from unsuitable sites, whilst limiting similarity leads to the divergence of trait values between co‐occurring species in order to alleviate competition for finite resources. Levels of alpha (within‐site) and beta (among‐site) trait variation can be indicative of the strength of these community assembly processes. We used trait‐gradient analysis to explicitly compare evidence of community assembly patterns in lianas (woody vines) and trees. These two growth forms exhibit striking differences in carbon capture and regeneration strategies, yet trait‐based mechanisms that maintain their coexistence remain understudied. Using data for four functional traits – leaf mass per area, leaf nitrogen content (N_mass), leaf area and seed mass – we partitioned interspecific trait variation in lianas and trees into alpha and beta components. Our three key findings were: 1) lianas and trees exhibit divergent patterns of trait‐based habitat filtering, due to differences in the relationship between leaf size and the other three traits examined (LMA, N_mass and seed mass), 2) on average, liana species possess smaller seeds, lower LMA and higher N_mass than do trees, but there was no clear difference in leaf area between the two growth forms, and 3) soil fertility was correlated with trait variation (leaf area, seed mass) at the site‐level in trees, but not in lianas. These results provide evidence that dominant growth forms can be filtered into the same habitat on the basis of different combinations of traits. Our findings have important implications for community assembly and co‐existence theory and for more pragmatic matters such as using trait‐based principles to inform community restoration.     

Phylogenetic patterns of trait and trait plasticity evolution: Insights from amphibian embryos

Environmental variation favors the evolution of phenotypic plasticity. For many species, we understand the costs and benefits of different phenotypes, but we lack a broad understanding of how plastic traits evolve across large clades. Using identical experiments conducted across North America, we examined prey responses to predator cues. We quantified five life‐history traits and the magnitude of their plasticity for 23 amphibian species/populations (spanning three families and five genera) when exposed to no cues, crushed‐egg cues, and predatory crayfish cues. Embryonic responses varied considerably among species and phylogenetic signal was common among the traits, whereas phylogenetic signal was rare for trait plasticities. Among trait‐evolution models, the Ornstein–Uhlenbeck (OU) model provided the best fit or was essentially tied with Brownian motion. Using the best fitting model, evolutionary rates for plasticities were higher than traits for three life‐history traits and lower for two. These data suggest that the evolution of life‐history traits in amphibian embryos is more constrained by a species’ position in the phylogeny than is the evolution of life history plasticities. The fact that an OU model of trait evolution was often a good fit to patterns of trait variation may indicate adaptive optima for traits and their plasticities.     

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.     

Leaf‐trait plasticity and species vulnerability to climate change in a Mongolian steppe

Climate change is expected to modify plant assemblages in ways that will have major consequences for ecosystem functions. How climate change will affect community composition will depend on how individual species respond, which is likely related to interspecific differences in functional traits. The extraordinary plasticity of some plant traits is typically neglected in assessing how climate change will affect different species. In the Mongolian steppe, we examined whether leaf functional traits under ambient conditions and whether plasticity in these traits under altered climate could explain climate‐induced biomass responses in 12 co‐occurring plant species. We experimentally created three probable climate change scenarios and used a model selection procedure to determine the set of baseline traits or plasticity values that best explained biomass response. Under all climate change scenarios, plasticity for at least one leaf trait correlated with change in species performance, while functional leaf‐trait values in ambient conditions did not. We demonstrate that trait plasticity could play a critical role in vulnerability of species to a rapidly changing environment. Plasticity should be considered when examining how climate change will affect plant performance, species' niche spaces, and ecological processes that depend on plant community composition.     

Functional trait differences and the outcome of community assembly: an experimental test with vernal pool annual plants

Functional trait‐based approaches have seen rapid development in community ecology and biogeography in recent years, as they promise to offer a better mechanistic and predictive understanding of community structure. However, several key challenges remain. First, while many studies have explored connections between functional traits and abiotic gradients, far fewer have directly tested the common assumption that functional trait differences influence interspecific interactions. Second, empirical studies often ignore intraspecific trait variation within communities, even though intraspecific variation has been known to have substantial impacts on community dynamics. Here we present an experiment designed to assess the role of functional trait differences in predicting the outcome of interspecific species interactions among a suite of California vernal pool annual plants. Eight species were grown in pairwise combinations in two levels of inundation in a greenhouse and functional traits were measured on all individuals. Nested models predicting focal plant performance were fit to the data. For seven of the eight species in the experiment, the best model included a functional trait difference term that was consistent with a competitive hierarchy, indicating that focal species tended to do better when they had larger leaf size, lower specific leaf area, and greater investment in lateral canopy spread than their neighbors. Models that included individually measured trait values generally performed better than models using species trait averages. We tested if the same trait measurements predicted tolerance of inundation (a feature of vernal pool habitats), and species depth distributions from extensive field surveys, though we did not find strong relationships. Our results suggest that functional traits can be used to make inferences about the outcome of interspecific interactions, and that greater predictive power can come from considering intraspecific variation in functional traits, particularly in low diversity communities.     

Characterizing the contribution of plasticity and genetic differentiation to community‐level trait responses to environmental change

The match between functional trait variation in communities and environmental gradients is maintained by three processes: phenotypic plasticity and genetic differentiation (intraspecific processes), and species turnover (interspecific). Recently, evidence has emerged suggesting that intraspecific variation might have a potentially large role in driving functional community composition and response to environmental change. However, empirical evidence quantifying the respective importance of phenotypic plasticity and genetic differentiation relative to species turnover is still lacking. We performed a reciprocal transplant experiment using a common herbaceous plant species (Oxalis montana) among low‐, mid‐, and high‐elevation sites to first quantify the contributions of plasticity and genetic differentiation in driving intraspecific variation in three traits: height, specific leaf area, and leaf area. We next compared the contributions of these intraspecific drivers of community trait–environment matching to that of species turnover, which had been previously assessed along the same elevational gradient. Plasticity was the dominant driver of intraspecific trait variation across elevation in all traits, with only a small contribution of genetic differentiation among populations. Local adaptation was not detected to a major extent along the gradient. Fitness components were greatest in O. montana plants with trait values closest to the local community‐weighted means, thus supporting the common assumption that community‐weighted mean trait values represent selective optima. Our results suggest that community‐level trait responses to ongoing climate change should be mostly mediated by species turnover, even at the small spatial scale of our study, with an especially small contribution of evolutionary adaptation within species.     

Evaluating differences in the shape of native and alien plant trait distributions will bring new insights into invasions of plant communities

Failure to quantify differences in the shape of inter‐specific trait distributions (e.g., skew, kurtosis) when comparing co‐occurring alien and native plants hinders the integration of biological invasions and plant community ecology. Within a plant community, understanding the circumstances that lead to the shape of the inter‐specific distribution of one or more functional plant traits being unimodal, bimodal, multimodal or skewed has the potential to shed new light on community vulnerability to invasion, subsequent ecosystem impacts and the selection pressures (e.g., stabilizing, directional or disruptive) acting upon native and alien species. Ignoring differences in the shape of inter‐specific trait distributions of alien and native species could miss important insights into plant invasions, including: the existence of unsaturated native plant communities, empty niches, shifting trait optima of species as a result of environmental change and incomplete colonization–extinction processes following invasion. Future comparisons of functional trait differences between native and alien species should include assessment of the shapes of inter‐specific trait distributions since these may differ even when the mean values of traits are similar for native and alien species. The infrequent application of such approaches may explain the limited generalizations regarding the drivers and consequences of plant invasions in plant communities.     

Modulation of leaf economic traits and trait relationships by climate

Aim Our aim was to quantify climatic influences on key leaf traits and relationships at the global scale. This knowledge provides insight into how plants have adapted to different environmental pressures, and will lead to better calibration of future vegetation–climate models. Location The data set represents vegetation from 175 sites around the world. Methods For more than 2500 vascular plant species, we compiled data on leaf mass per area (LMA), leaf life span (LL), nitrogen concentration (N_mass) and photosynthetic capacity (A_mass). Site climate was described with several standard indices. Correlation and regression analyses were used for quantifying relationships between single leaf traits and climate. Standardized major axis (SMA) analyses were used for assessing the effect of climate on bivariate relationships between leaf traits. Principal components analysis (PCA) was used to summarize multidimensional trait variation. Results At hotter, drier and higher irradiance sites, (1) mean LMA and leaf N per area were higher; (2) average LL was shorter at a given LMA, or the increase in LL was less for a given increase in LMA (LL–LMA relationships became less positive); and (3) A_mass was lower at a given N_mass, or the increase in A_mass was less for a given increase in N_mass. Considering all traits simultaneously, 18% of variation along the principal multivariate trait axis was explained by climate. Main conclusions Trait‐shifts with climate were of sufficient magnitude to have major implications for plant dry mass and nutrient economics, and represent substantial selective pressures associated with adaptation to different climatic regimes.     

Plant invasion alters trait composition and diversity across habitats

Increased globalization has accelerated the movement of species around the world. Many of these nonnative species have the potential to profoundly alter ecosystems. The mechanisms underpinning this impact are often poorly understood, and traits are often overlooked when trying to understand and predict the impacts of species invasions on communities. We conducted an observational field experiment in Canada's first National Urban Park, where we collected trait data for seven different functional traits (height, stem width, specific leaf area, leaf percent nitrogen, and leaf percent carbon) across an abundance gradient of the invasive Vincetoxicum rossicum in open meadow and understory habitats. We assessed invasion impacts on communities, and associated mechanisms, by examining three complementary functional trait measures: community‐weighted mean, range of trait values, and species’ distances to the invader in trait space. We found that V. rossicum invasion significantly altered the functional structure of herbaceous plant communities. In both habitats V. rossicum changed the community‐weighted means, causing invaded communities to become increasingly similar in their functional structure. In addition, V. rossicum also reduced the trait ranges for a majority of traits indicating that species are being deterministically excluded in invaded communities. Further, we observed different trends in the meadow and understory habitats: In the understory, resident species that were more similar to V. rossicum in multivariate trait space were excluded more, however this was not the case in the meadow habitat. This suggests that V. rossicum alters communities uniquely in each habitat, in part by creating a filter in which only certain resident species are able to persist. This filtering process causes a nonrandom reduction in species' abundances, which in turn would be expected to alter how the invaded ecosystems function. Using trait‐based frameworks leads to better understanding and prediction of invasion impacts. This novel framework can also be used in restoration practices to understand how invasion impacts communities and to reassemble communities after invasive species management.     

Linking intra‐specific trait variation and plant function: seed size mediates performance tradeoffs within species

Substantial intra‐specific trait variation exists within plant communities, and in theory this variation could influence community dynamics. Although recent research has focused on intra‐specific variation in traits themselves, it is the influence of this variation on plant performance that makes intra‐specific trait variation relevant to ecological dynamics within or among species. Understanding the links between trait and performance variation, and the role of traits in mediating relationships among multiple components of performance, is critical for assessing the importance of intra‐specific trait variation for community dynamics. Seed size is thought to affect aspects of plant performance including fecundity, seedling growth, dispersal and tolerance of natural enemies. For two tropical tree species, we assessed how seed size was related to performance variation within each species and determined whether intra‐specific trait variation mediates intra‐specific performance tradeoffs. We used field seed rain collection to characterize size‐dependent outcomes of dispersal, sowed seeds of known size in soil collected near or far from conspecifics to characterize susceptibility to soil pathogens, and monitored growth of seedlings from seeds of known size. We found that intra‐specific seed size variation caused intra‐specific performance variation. The degree of trait‐based performance variation was consistently smaller than the degree of trait variation, and seed size influenced different components of performance for each species. One species exhibited a tradeoff in which small seeds had a fecundity advantage (more seedlings per unit reproductive mass) but produced smaller seedlings, whereas the other species exhibited a tradeoff in which small seeds dispersed to areas of low conspecific density but were less tolerant of density‐responsive natural enemies. Our results indicate that a single trait can influence multiple components of performance and can mediate different tradeoffs in co‐occurring species. Complex and heterogeneous effects of a single trait in multidimensional niche space may favour inter‐specific niche differentiation and coexistence.     

Grazing effects on intraspecific trait variability vary with changing precipitation patterns in Mongolian rangelands

Functional traits are proxies for plant physiology and performance, which do not only differ between species but also within species. In this work, we hypothesized that (a) with increasing precipitation, the percentage of focal species which significantly respond to changes in grazing intensity increases, while under dry conditions, climate‐induced stress is so high that plant species hardly respond to any changes in grazing intensity and that (b) the magnitude with which species change their trait values in response to grazing, reflected by coefficients of variation (CVs), increases with increasing precipitation. Chosen plant traits were canopy height, plant width, specific leaf area (SLA), chlorophyll fluorescence, performance index, stomatal pore area index (SPI), and individual aboveground biomass of 15 species along a precipitation gradient with different grazing intensities in Mongolian rangelands. We used linear models for each trait to assess whether the percentage of species that respond to grazing changes along the precipitation gradient. To test the second hypothesis, we assessed the magnitude of intraspecific trait variability (ITV) response to grazing, per species, trait, and precipitation level by calculating CVs across the different grazing intensities. ITV was most prominent for SLA and SPI under highest precipitation, confirming our first hypothesis. Accordingly, CVs of canopy height, SPI, and SLA increased with increasing precipitation, partly confirming our second hypothesis. CVs of the species over all traits increased with increasing precipitation only for three species. This study shows that it remains challenging to predict how plant performance will shift under changing environmental conditions based on their traits alone. In this context, the implications for the use of community‐weighted mean trait values are discussed, as not only species abundances change in response to changing environmental conditions, but also values of traits considerably change. Including this aspect in further studies will improve our understanding of processes acting within and among communities.     

Leaf trait co‐variation,response and effect in a chronosequence

Question: Is there any generality in terms of leaf trait correlations and the multiple role of leaf traits (response to and/or effect on) during secondary succession? Location: A secondary successional sere was sampled at four different ages since abandonment from several years to nearly 150 years on the Loess Plateau of northwestern China. Method: Specific leaf area (SLA), leaf mass per area (LMA), leaf nitrogen (N_mass, N_area), leaf phosphorus (P_mass, P_area) and leaf dry matter content (LDMC) were measured for all species recorded in the successional sere. Above‐ground net primary productivity (ANPP) and specific rate of litter mass loss (SRLML) were measured as surrogates for ecosystem properties. Soil total carbon (C) and nitrogen (N) were measured in each stage. Leaf traits were related to ecosystem properties and soil nutrient gradients, respectively. Results: LMA is correlated with N_area and P_area' and negatively with N_mass. Correlation between N_area and P_area was higher than between N_mass and P_mass. At the community level, field age, community hierarchy and their interaction explain 64.4 ‐ 93.5% of the variation in leaf traits. At the species level, field age explains 22.4 ‐ 45.5% of the variation in leaf traits (excl. P_area) while plant functional group has a significant effect only for N_mass. LDMC is correlated with ANPP and negatively with SRLML; P_mass is correlated with SRLML. Conclusions: Mean values of LMA, N_mass and N_area are close to the worldwide means, suggesting that large‐scale climate has a profound effect on leaf mass and leaf nitrogen allocation, while environmental gradients represented by succession have little influence on leaf‐trait values. Correlations between leaf traits, such as LMA‐N_area, LMA‐P_area and LMA‐N_mass shown in previous studies, are confirmed here. Although none of the leaf traits is proved to be both a response trait and an effect trait independent of time scale and community hierarchy, mass‐based leaf N is likely a sensitive response trait to soil C and N gradients. In addition, LDMC can be a marker for ANPP and SRLML, while mass‐based leaf P can be a marker for SRLML.     

A unified framework for diversity gradients: the adaptive trait continuum

Aim Adaptive trait continua are axes of covariation observed in multivariate trait data for a given taxonomic group. These continua quantify and summarize life‐history variation at the inter‐specific level in multi‐specific assemblages. Here we examine whether trait continua can provide a useful framework to link life‐history variation with demographic and evolutionary processes in species richness gradients. Taking an altitudinal species richness gradient for Mediterranean butterflies as a study case, we examined a suite of traits (larval diet breadth, adult phenology, dispersal capacity and wing length) and species‐specific habitat measures (temperature and aridity breadth). We tested whether traits and species‐specific habitat measures tend to co‐vary, whether they are phylogenetically conserved, and whether they are able to explain species distributions and spatial genetic variation in a large number of butterfly assemblages. Location Catalonia, Spain. Methods We formulated predictions associated with species richness gradients and adaptive trait continua. We applied principal components analyses (PCAs), structural equation modelling and phylogenetic generalized least squares models. Results We found that traits and species‐specific habitat measures covaried along a main PCA axis, ranging from multivoltine trophic generalists with high dispersal capacity to univoltine (i.e. one generation per year), trophic specialist species with low dispersal capacity. This trait continuum was closely associated with the observed distributions along the altitudinal gradient and predicted inter‐specific differences in patterns of spatial genetic variability (F_ST and genetic distances), population responses to the impacts of global change and local turnover dynamics. Main conclusions The adaptive trait continuum of Mediterranean butterflies provides an integrative and mechanistic framework to: (1) analyse geographical gradients in species richness, (2) explain inter‐specific differences in population abundances, spatial distributions and demographic trends, (3) explain inter‐specific differences in patterns of genetic variation (F_ST and genetic distances), and (4) study specialist–generalist life‐history transitions frequently involved in butterfly diversification processes.     

Plant functional trait response to environmental drivers across European temperate forest understorey communities

• Functional traits respond to environmental drivers, hence evaluating trait‐environment relationships across spatial environmental gradients can help to understand how multiple drivers influence plant communities. Global‐change drivers such as changes in atmospheric nitrogen deposition occur worldwide, but affect community trait distributions at the local scale, where resources (e.g. light availability) and conditions (e.g. soil pH) also influence plant communities.
• We investigate how multiple environmental drivers affect community trait responses related to resource acquisition (plant height, specific leaf area (SLA), woodiness, and mycorrhizal status) and regeneration (seed mass, lateral spread) of European temperate deciduous forest understoreys. We sampled understorey communities and derived trait responses across spatial gradients of global‐change drivers (temperature, precipitation, nitrogen deposition, and past land use), while integrating in‐situ plot measurements on resources and conditions (soil type, Olsen phosphorus (P), Ellenberg soil moisture, light, litter mass, and litter quality).
• Among the global‐change drivers, mean annual temperature strongly influenced traits related to resource acquisition. Higher temperatures were associated with taller understoreys producing leaves with lower SLA, and a higher proportional cover of woody and obligate mycorrhizal (OM) species. Communities in plots with higher Ellenberg soil moisture content had smaller seeds and lower proportional cover of woody and OM species. Finally, plots with thicker litter layers hosted taller understoreys with larger seeds and a higher proportional cover of OM species.
• Our findings suggest potential community shifts in temperate forest understoreys with global warming, and highlight the importance of local resources and conditions as well as global‐change drivers for community trait variation.

     

Intraspecific trait variation and allocation strategies of calcareous grassland species: Results from a restoration experiment

Intra- and interspecific trait variation express the response of plants dealing with different environmental conditions. We measured root and leaf traits on 14 species of calcareous grasslands in a restoration experiment. We aimed at identifying intraspecific differences in biomass allocation and functional plant traits under contrasting soil conditions by comparing plants growing in ancient grassland and two restored grasslands on ex-arable land, one of them with topsoil removal. Relative importance of trait variation within and among species, and among site was assessed by variance partitioning. Interspecific variation was more important than intraspecific variation, but the contribution of the latter to total variation was considerable, especially for specific leaf area. Changes in soil properties due to topsoil removal resulted in lower values of plant height, specific leaf area and specific root length compared to the control (ancient grassland). Soil fertility found in the treatment without top soil removal did not affect plant plasticity compared to the control. The study species showed two allocation strategies in relation to resource stress, while the responses of individual traits to the soil treatments were consistent across species. We conclude that caution must be taken when using mean trait values for plastic species or when working with environmental gradients.     

The acquisitive–conservative axis of leaf trait variation emerges even in homogeneous environments

Background and AimsThe acquisitive–conservative axis of plant ecological strategies results in a pattern of leaf trait covariation that captures the balance between leaf construction costs and plant growth potential. Studies evaluating trait covariation within species are scarcer, and have mostly dealt with variation in response to environmental gradients. Little work has been published on intraspecific patterns of leaf trait covariation in the absence of strong environmental variation.MethodsWe analysed covariation of four leaf functional traits [specific leaf area (SLA) leaf dry matter content (LDMC), force to tear (F_t) and leaf nitrogen content (N_m)] in six Poaceae and four Fabaceae species common in the dry Chaco forest of Central Argentina, growing in the field and in a common garden. We compared intraspecific covariation patterns (slopes, correlation and effect size) of leaf functional traits with global interspecific covariation patterns. Additionally, we checked for possible climatic and edaphic factors that could affect the intraspecific covariation pattern.Key ResultsWe found negative correlations for the LDMC–SLA, F_t–SLA, LDMC–N_m and F_t–N_m trait pairs. This intraspecific covariation pattern found both in the field and in the common garden and not explained by climatic or edaphic variation in the field follows the expected acquisitive–conservative axis. At the same time, we found quantitative differences in slopes among different species, and between these intraspecific patterns and the interspecific ones. Many of these differences seem to be idiosyncratic, but some appear consistent among species (e.g. all the intraspecific LDMC–SLA and LDMC–N_m slopes tend to be shallower than the global pattern).ConclusionsOur study indicates that the acquisitive–conservative leaf functional trait covariation pattern occurs at the intraspecific level even in the absence of relevant environmental variation in the field. This suggests a high degree of variation–covariation in leaf functional traits not driven by environmental variables.     

Trait–performance relationships of grassland plant species differ between common garden and field conditions

The way functional traits affect growth of plant species may be highly context‐specific. We asked which combinations of trait values are advantageous under field conditions in managed grasslands as compared to conditions without competition and land‐use. In a two‐year field experiment, we recorded the performance of 93 species transplanted into German grassland communities differing in land‐use intensity and into a common garden, where species grew unaffected by land‐use under favorable conditions regarding soil, water, and space. The plants’ performance was characterized by two independent dimensions (relative growth rates (RGR) of height and leaf length vs. aboveground biomass and survival) that were differently related to the eight focal key traits in our study (leaf dry matter content (LDMC), specific leaf area (SLA), height, leaf anatomy, leaf persistence, leaf distribution, vegetative reproduction, and physical defense). We applied multivariate procrustes analyses to test for the correspondence of the optimal trait–performance relationships between field and common garden conditions. RGRs were species‐specific and species ranks of RGRs in the field, and the common garden were significantly correlated. Different traits explained the performance in the field and the common garden; for example, leaf anatomy traits explained species performance only in the field, whereas plant height was found to be only important in the common garden. The ability to reproduce vegetatively, having leaves that are summer‐persistent and with high leaf dry matter content (LDMC) were traits of major importance under both settings, albeit the magnitude of their influence differed slightly between the field and the common garden experiment. All optimal models included interactions between traits, pointing out the necessity to analyze traits in combination. The differences between field and common garden clearly demonstrate context dependency of trait‐based growth models, which results in limited transferability of favorable trait combinations between different environmental settings.     

Does grazing induce intraspecific trait variation in plants from a sub‐humid mountain ecosystem?

Livestock grazing represents an important human disturbance for vegetation worldwide. We analysed the intraspecific differences in mean trait values between different grazing regimes (ungrazed and grazed) and explored whether these differences are consistent across species in a sub‐humid mountain ecosystem in Central Argentina. We selected 14 species of eight different families, co‐occurring in both regimes and comprising herbaceous (grasses and forbs) and woody (shrubs and trees) plants. For each species and grazing regime we measured 12 traits related to plant size, carbon fixation and water use. We found that plants in the grazed regime had consistently smaller leaves and shorter stature and internodal length than plants of the same species under the ungrazed regime. For the remaining traits the responses were species‐specific. Dry matter content, leaf tensile strength and minimum leaf water potential (Ψ_leaf) showed contrasting responses to grazing. Specific leaf area, wood density and potential water content of wood showed almost no significant responses except for very few species. Neither leaf area per shoot mass nor leaf area per sapwood area differed significantly between grazing regimes. Our study suggested that the intraspecific variation found for the size‐related traits would allow species to respond to grazing without modifying markedly other structural traits, a plastic response that might increase the probability of species success.     

Community trait response to environment: disentangling species turnover vs intraspecific trait variability effects

Ecological communities and their response to environmental gradients are increasingly being described by various measures of trait composition. Aggregated trait averages (i.e. averages of trait values of constituent species, weighted by species proportions) are popular indices reflecting the functional characteristics of locally dominant species. Because the variation of these indices along environmental gradients can be caused by both species turnover and intraspecific trait variability, it is necessary to disentangle the role of both components to community variability. For quantitative traits, trait averages can be calculated from ‘fixed’ trait values (i.e. a single mean trait value for individual species used for all habitats where the species is found) or trait values for individual species specific to each plot, or habitat, where the species is found. Changes in fixed averages across environments reflect species turnover, while changes in specific traits reflect both species turnover and within‐species variability in traits. Here we suggest a practical method (accompanied by a set of R functions) that, by combining ‘fixed’ and ‘specific averages’, disentangles the effect of species turnover, intraspecific trait variability, and their covariation. These effects can be further decomposed into parts ascribed to individual explanatory variables (i.e. treatments or environmental gradients considered). The method is illustrated with a case study from a factorial mowing and fertilization experiment in a meadow in South Bohemia. Results show that the variability decomposition differs markedly among traits studied (height, Specific Leaf Area, Leaf N, P, C concentrations, leaf and stem dry matter content), both according to the relative importance of species turnover and intraspecific variability, and also according to their response to experimental factors. Both the effect of intraspecific trait variability and species turnover must be taken into account when assessing the functional role of community trait structure. Neglecting intraspecific trait variability across habitats often results in underestimating the response of communities to environmental changes.