The contribution of intraspecific trait variability to the functional diversity of herbaceous vegetation along a local environmental gradient

We assessed the changes in community-weighted mean (CWM) and variability of specific leaf area (SLA) and leaf area (LA) of different Mediterranean shrub communities along an elevation gradient in the island of Sardinia (Italy). Furthermore, we explored the relative contribution of species turnover and intraspecific variation to shifts in CWM values along the gradient. Forty sampling units (5 × 5 m) were selected in a probabilistic way along a 1300 m elevation gradient which crossed four thermotypes (thermometric belts). Leaf traits were measured in each sampling unit. ANOVA and a trend test for monotonic changes in variance were used to assess, respectively, CWM differences and variability in both the leaf traits across thermotypes. Variance decomposition of CWM values was used to identify the role of inter- and intraspecific variation. SLA and LA responded differently along the studied gradient in terms of abundance-weighted mean values and variability: CWM of SLA showed the lowest values in the driest thermotype, while LA in the more humid one; SLA variability showed a significant increasing trend with increased water availability, while LA variability did not show any pattern. The contribution of intraspecific trait variation was significant for both the leaf traits, but higher for SLA, where negative covariation between inter- and intraspecific variation was detected. We highlight the importance of simultaneously considering measurements of both leaf traits to understand the functional response of communities in Mediterranean environments. Moreover, neglecting intraspecific variation in leaf traits, even along steep gradients with relevant species compositional changes, can result in the underestimation of the amount of trait variation in response to environmental changes.

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.

Summary Climate change is expected to increase the magnitude and the frequency of extreme climatic events such as droughts. Better understanding how plant communities will respond to these droughts is a major challenge. We expect the response to be a shift in functional trait values resulting from both species turnover and intraspecific trait variability, but little research has addressed the relative contribution of both components. We analysed the short‐term functional response of subalpine grassland communities to a simulated drought by focusing on four leaf traits (LDMC: leaf dry matter content, SLA: specific leaf area, LNC: leaf nitrogen concentration and LCC: leaf carbon concentration). After evaluating species turnover and intraspecific variability separately, we determined their relative contribution in the community functional response to drought, reflected by changes in community‐weighted mean traits. We found significant species turnover and intraspecific variability, as well as significant changes in community‐weighted mean for most of the traits. The relative contribution of intraspecific variability to the changes in community mean traits was more important (42–99%) than the relative contribution of species turnover (1–58%). Intraspecific variability either amplified (for LDMC, SLA and LCC) or dampened (for LNC) the community functional response mediated by species turnover. We demonstrated that the small contribution of species turnover to the changes in community mean LDMC and LCC was explained by a lack of covariation between species turnover and interspecific trait differences. Synthesis. These results highlight the need for a better consideration of intraspecific variability to understand and predict the effect of climate change on plant communities. While both species turnover and intraspecific variability can be expected following an extreme drought, we report new evidence that intraspecific variability can be a more important driver of the short‐term functional response of plant communities.

Intraspecific trait variation governs grazing-induced shifts in plant community above- and below-ground functional trait composition

BackgroundInter- and intraspecific variation in plant traits play an important role in grassland community assembly under global change scenarios. However, explorations of how these variations contribute to the responses of community traits to nitrogen (N) addition and drought in different grassland types are lacking. We measured the plant height, leaf area (LA), specific leaf area (SLA), leaf dry matter content (LDMC), leaf N content (LNC) and the ratio of leaf carbon (C) to leaf N (C:N) in a typical and a meadow steppe after three years of N addition, drought and their interaction. We determined the community-weighted means (CWMs) of the six traits to quantify the relative contribution of inter- and intraspecific variation to the responses of community traits to N addition and drought in the two steppes.ResultsThe communities in the two steppes responded to N addition and the interaction by increasing the CWM of LNC and decreasing C:N. The community in the meadow steppe responded to drought through increased CWM of LNC and reduced C:N. Significant differences were observed in SLA, LDMC, LNC and C:N between the two steppes under different treatments. The SLA and LNC of the community in the meadow steppe were greater than those of the typical steppe, and the LDMC and C:N exhibited the opposite results. Moreover, variation in community traits in the typical steppe in response to N addition and drought was caused by intraspecific variation. In contrast, the shifts in community traits in the meadow steppe in response to N addition and drought were influenced by both inter- and intraspecific variation.ConclusionsThe results demonstrate that intraspecific variation contributed more to community functional shifts in the typical steppe than in the meadow steppe. Intraspecific variation should be considered to understand better and predict the response of typical steppe communities to global changes. The minor effects of interspecific variation on meadow steppe communities in response to environmental changes also should not be neglected.

QuestionsBoth species turnover and intraspecific trait variation can affect plant assemblage dynamics along environmental gradients. Here, we asked how community assemblage patterns in relation to species turnover and intraspecific variation differ between endemic and non‐endemic species. We hypothesized that endemic species show lower intraspecific variation than non‐endemic species because they tend to have high rates of in situ speciation, whereas non‐endemic species are expected to have a larger gene pool and higher phenotypic plasticity.LocationLa Palma, Canary Islands.MethodsWe established 44 sampling sites along a directional gradient of precipitation, heat load, soil nitrogen, phosphorus and pH. Along this gradient, we estimated species abundances and measured three traits (plant height, leaf area and leaf thickness) on perennial endemic and non‐endemic plant species. In total, we recorded traits for 1,223 plant individuals of 43 species. Subsequently, we calculated community‐weighted mean traits to measure the relative contribution of species turnover, intraspecific variation and their covariation along the analysed gradient.ResultsThe contribution of intraspecific variation to total variation was similar in endemic and non‐endemic assemblages. For plant height, intraspecific variation explained roughly as much variation as species turnover. For leaf area and leaf thickness, intraspecific variation explained almost no variation. Species turnover effects mainly drove trait responses along the environmental gradient, but intraspecific variation was important for responses in leaf area to precipitation.ConclusionsDespite their distinct evolutionary history, endemic and non‐endemic plant assemblages show similar patterns in species turnover and intraspecific variation. Our results indicate that species turnover is the main component of trait variation in the underlying study system. However, intraspecific variation can increase individual species’ fitness in response to precipitation. Overall, our study challenges the theory that intraspecific trait variation is more important for the establishment of non‐endemic species compared with endemic species.

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.

Plant community assembly is determined by species turnover and intraspecific trait variations (ITV) controlled by environment changes. However, little is known about how species turnover and ITV affect the responses of plant community to habitat changes and grazing disturbance in semiarid grasslands. Here, we measured five functional plant traits in four typical grassland habitats under fencing and grazing disturbance in a semiarid grassland, Northern China, including plant height, specific leaf area (SLA), leaf dry matter content (LDMC), leaf nitrogen content (LNC) and leaf carbon: nitrogen ratio (C:N). We also calculated the community weighted means (CWM) and non-weighted means (CM) of all traits and examined the relative roles of species turnover and ITV in affecting the responses of community traits to habitat changes and grazing disturbance. Our results showed that the CWM and CM values of five functional traits differed with grassland habitat changes. As compared to other grasslands, the Stipa steppe had the higher plant height, the sandy grassland had the higher SLA and lower LDMC, and the meadow had the lower LNC and higher C:N. Grazing decreased plant height across grassland habitats, as well as decreased SLA and increased LDMC in meadow. The responses of all community-level traits to habitat changes were driven by species turnover, while the responses of phenotypic traits (height, SLA and LDMC) to grazing were determined by both species turnover and ITV. So, we argue that ITV should be considered when understanding plant community assembly under grazing disturbance regime in semiarid grasslands.

Fertilization could influence ecosystem structure and functioning through species turnover (ST) and intraspecific trait variation (ITV), especially in nutrient limited ecosystems. To quantify the relative importance of ITV and ST in driving community functional structure and productivity changes under nitrogen (N) and phosphorous (P) addition in semiarid grasslands. In this regard, we conducted a four-year fertilizer addition experiment in a semiarid grassland on the Loess Plateau, China. We examined how fertilization affects species-level leaf and root trait plasticity to evaluate the ability of plants to manifest different levels of traits in response to different N and P addition. Also, we assessed how ITV or ST dominated community-weighted mean (CWM) traits and functional diversity variations and evaluated their effects on grassland productivity. The results showed that the patterns of plasticity varied greatly among different plant species, and leaf and root traits showed coordinated variations following fertilization. Increasing the level of N and P increased CWM_specific leaf area (CWM_SLA), CWM_leaf N concentration (CWM_LN) and CWM_maximum plant height (CWM_Hmax) and ITV predominate these CWM traits variations. As a results, increased CWM_Hmax, CWM_LN and CWM_SLA positively influenced grassland productivity. In contrast, functional divergence decreased with increasing N and P and showed negative relationships with grassland productivity. Our results emphasized that CWM traits and functional diversity contrastingly drive changes in grassland productivity under N and P addition.

Despite increasing evidence of the importance of intraspecific trait variation in plant communities, its role in community trait responses to environmental variation, particularly along broad-scale climatic gradients, is poorly understood. We analyzed functional trait variation among early-successional herbaceous plant communities (old fields) across a 1200-km latitudinal extent in eastern North America, focusing on four traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We determined the contributions of species turnover and intraspecific variation to between-site functional dissimilarity at multiple spatial scales and community trait responses to edaphic and climatic factors. Among-site variation in community mean trait values and community trait responses to the environment were generated by a combination of species turnover and intraspecific variation, with species turnover making a greater contribution for all traits. The relative importance of intraspecific variation decreased with increasing geographic and environmental distance between sites for SLA and leaf area. Intraspecific variation was most important for responses of vegetative height and responses to edaphic compared to climatic factors. Individual species displayed strong trait responses to environmental factors in many cases, but these responses were highly variable among species and did not usually scale up to the community level. These findings provide new insights into the role of intraspecific trait variation in plant communities and the factors controlling its relative importance. The contribution of intraspecific variation to community trait responses was greatest at fine spatial scales and along edaphic gradients, while species turnover dominated at broad spatial scales and along climatic gradients.

Summary 1. Trait‐based approaches applied to community ecology have led to a considerable advance in understanding the effect of environmental filters on species assembly. Although plant traits are known to vary both between and within species, little is known about the role of intraspecific trait variability in the non‐random assembly mechanisms controlling the coexistence of species, including habitat filtering and niche differentiation. 2. We investigate the role of intraspecific variability in three key functional traits – specific leaf area (SLA), leaf dry matter content (LDMC) and height – in structuring grassland communities distributed along a flooding gradient. We quantified the contribution of intraspecific variability relative to interspecific differences in the trait–gradient relationship, and we used a null model approach to detect patterns of habitat filtering and niche differentiation, with and without intraspecific variability. 3. Community mean SLA and height varied significantly along the flooding gradient and intraspecific variability accounted for 44% and 32%, respectively, of these trait–gradient relationships. LDMC did not vary along the gradient, with and without accounting for intraspecific variability. Our null model approach revealed significant patterns of habitat filtering and niche differentiation for SLA and height, but not for LDMC. More strikingly, considering intraspecific trait variability greatly increased the detection of habitat filtering and was necessary to detect niche differentiation processes. 4. Synthesis. Our study provides evidence for a strong role of intraspecific trait variability in community assembly. Our findings suggest that intraspecific trait variability promotes species coexistence, by enabling species to pass through both abiotic and biotic filters. We argue that community ecology would benefit from more attention to intraspecific variability.

Traditional approaches in trait-based community ecology typically expect that trait filtering across broad environmental gradients is largely due to replacement of species, rather than intraspecific trait adjustments. Recently, the role of intraspecific trait variability has been largely highlighted as an important contributor mediating the ability of communities to persist under changing conditions and determining the community-level trait variation, particularly across limited environmental gradients. Unfortunately, few studies quantify the relative importance of species turnover versus intraspecific variability mediating the response of communities different from vascular plants. Here, we studied the functional changes in epiphytic lichen communities within 23 beech forests across large latitudinal (ca. 3,000km) and environmental gradients in Europe to quantify the relative contribution of species turnover and intraspecific variability and the role of climate controlling community-level trait changes. For 58 lichen species, we focused on a set of 10 quantitative functional traits potentially affected by climatic conditions and related to photosynthetic performance (n=1,184 thalli), water use strategy (n=1,018 thalli), and nutrient uptake (n=1,179 thalli). Our results showed that intraspecific trait variability explained most of the functional changes in lichen communities in response to the latitudinal gradient. Further, such functional changes were determined by the covariation between intraspecific trait variability and species turnover, which varied in sign depending on the trait considered. Finally, different climatic predictors explained functional variation due to both intraspecific trait variability and species turnover. We propose that lichen communities cope with contrasting climatic conditions by adjusting the functional trait values of the most abundant species within the communities rather than by the replacement of the species. Consequently, intraspecific variability should be explicitly incorporated to understand the effect of environmental changes on lichen communities, even over large environmental variations, better. Our results challenge the universality of the hypothesis that species turnover chiefly drives functional trait changes across large environmental gradients and call for a wider test of such important assumptions in trait ecology in different organism types and ecosystems.

Environmental changes are expected to shift the distribution of functional trait values in plant communities through a combination of species turnover and intraspecific variation. The strength of these shifts may depend on the availability of individuals with trait values adapted to new environmental conditions, represented by the functional diversity (FD) of existing community residents or dispersal from the regional species pool. We conducted a 3-year nutrient- and seed-addition experiment in old-field plant communities to examine the contributions of species turnover and intraspecific variation to community trait shifts, focusing on four key plant functional traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We further examined the influence of initial FD and seed availability on the strength of these shifts. Community mean height, leaf area, and SLA increased in response to fertilization, and these shifts were driven almost entirely by intraspecific variation. The strength of intraspecific shifts in height and leaf area was positively related to initial intraspecific FD in these traits. Intraspecific trait responses to fertilization varied among species, with species of short stature displaying stronger shifts in SLA and LDMC but weaker shifts in leaf area. Trait shifts due to species turnover were generally weak and opposed intraspecific responses. Seed addition altered community taxonomic composition but had little effect on community trait shifts. These results highlight the importance of intraspecific variation for short-term community functional responses and demonstrate that the strength of these responses may be mediated by community FD.

A community functional structure may respond to environmental changes such as nitrogen (N) enrichment by altering intraspecific and interspecific trait variations. However, the relative contributions of both components in determining the community response to N enrichment are unclear. In this study, we measured the plant height (H), leaf area (LA), leaf dry matter content (LDMC), and specific leaf area (SLA) based on a nine-year N addition gradient experiment in an alpine meadow on the Tibetan Plateau. We examined the intraspecific and interspecific variations within and among the communities, the responses of traits in terms of community weighted mean (CWM) and non-weighted mean (CM) to N addition, and the effects of these trait variations on aboveground net primary productivity (ANPP). Our results show that N addition increased the interspecific variation in H while decreasing that of LA within the community, whereas it had no significant effects on the intraspecific variations in the four traits within the community. In contrast, N addition significantly increased the intraspecific variation in H and decreased that of LA among the communities. Moreover, the contribution of intraspecific variation was greater than that of the interspecific variation in terms of CWM for all traits, while the opposite contribution was observed in terms of CM, suggesting that the dominant species would have greater resilience while subdominant species would become less resistant to N addition. Further, intraspecific variations of LA and LDMC within the community played an important role in explaining community productivity. Our results highlight the importance of both intraspecific and interspecific variations in mediating functional trait responses to N enrichment, and intraspecific variation within the communities has important implications for community functioning that should be considered to better understand and predict the responses of the alpine grasslands to N enrichment.

Summary Despite widespread focus on interspecific variation in trait‐based ecology, there is growing evidence that intraspecific trait variability can play a fundamental role in plant community responses to environmental change and community assembly. Here, we quantify the strength and direction of inter‐ and intraspecific plant community trait responses along a 900 m elevation gradient spanning alpine and subalpine plant communities in southern New Zealand. We measured five commonly used leaf traits (i.e. dry matter content, N and P concentrations, leaf area and specific leaf area) on all 31 dominant and subordinate species recorded along the gradient, and examined their species‐specific and community‐level responses to elevation using both abundance‐weighted and nonweighted averages of trait values. By decomposing the variance of community‐level measures of these traits across the gradient, we showed that the contribution of interspecific variation to the response of plant assemblages to elevation was stronger than that of intraspecific variation, for all traits except specific leaf area. Further, the relative contributions of interspecific effects were greater when abundance‐weighted rather than nonweighted measures were used. We also observed contrasting intraspecific trait responses to the gradient among species (particularly for leaf N and P concentrations), and found both positive and negative covariation between inter‐ and intraspecific effects on community‐level trait values. The weak community‐average trait responses to elevation, as found for specific leaf area (SLA) and leaf N and P concentrations, resulted from strong but opposing responses among vs. within species, which are not typically accounted for in species‐based measures of plant community responses. For instance, increasing elevation (and associated factors such as a decrease in soil nutrient availability) favoured the dominance of species with relatively high leaf nutrient concentrations while simultaneously triggering an intraspecific decrease in the leaf nutrient concentrations of these species. The context dependency of positive and negative covariation between inter‐ and intraspecific trait variability, and the species‐specific nature of intraspecific shifts in functional trait values, reveal highly complex plastic responses of plants to environmental changes, and highlights the need for greater consideration of the role that intraspecific variation plays in community‐level processes.