Individual‐level leaf trait variation and correlation across biological and spatial scales

Even with increasing interest in the ecological importance of intraspecific trait variation (ITV) for better understanding ecological processes, few studies have quantified ITV in seedlings and assessed constraints imposed by trade‐offs and correlations among individual‐level leaf traits. Estimating the amount and role of ITV in seedlings is important to understand tree recruitment and long‐term forest dynamics. We measured ten different size, economics, and whole leaf traits (lamina and petiole) for more than 2,800 seedlings (height ≥ 10 cm and diameter at breast height < 1 cm) in 283 seedling plots and then quantified the amount of ITV and trait correlations across two biological (intraspecific and interspecific) and spatial (within and among plots) scales. Finally, we explored the effects of trait variance and sample size on the strength of trait correlations. We found about 40% (6%–63%) variation in leaf‐level traits was explained by ITV across all traits. Lamina and petiole traits were correlated across biological and spatial scales, whereas leaf size traits (e.g., lamina area) were weakly correlated with economics traits (e.g., specific lamina area); lamina mass ratio was strongly related to the petiole length. Trait correlations varied among species, plots, and different scales but there was no evidence that the strength of trait relationships was stronger at broader than finer biological and spatial scales. While larger trait variance increased the strength of correlations, the sample size was the most important factor that was negatively related to the strength of trait correlations. Our results showed that a large amount of trait variation was explained by ITV, which highlighted the importance of considering ITV when using trait‐based approaches in seedling ecology. In addition, sample size was an important factor that influenced the strength of trait correlations, which suggests that comparing trait correlations across studies should consider the differences in sample size.     

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

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Intraspecific trait variation and reversals of trait strategies across key climate gradients in native Hawaiian plants and non-native invaders

Background and AimsDisplacement of native plant species by non-native invaders may result from differences in their carbon economy, yet little is known regarding how variation in leaf traits influences native–invader dynamics across climate gradients. In Hawaii, one of the most heavily invaded biodiversity hotspots in the world, strong spatial variation in climate results from the complex topography, which underlies variation in traits that probably drives shifts in species interactions.MethodsUsing one of the most comprehensive trait data sets for Hawaii to date (91 species and four islands), we determined the extent and sources of variation (climate, species and species origin) in leaf traits, and used mixed models to examine differences between natives and non-native invasives.Key ResultsWe detected significant differences in trait means, such that invasives were more resource acquisitive than natives over most of the climate gradients. However, we also detected trait convergence and a rank reversal (natives more resource acquisitive than invasives) in a sub-set of conditions. There was significant intraspecific trait variation (ITV) in leaf traits of natives and invasives, although invasives expressed significantly greater ITV than natives in water loss and photosynthesis. Species accounted for more trait variation than did climate for invasives, while the reverse was true for natives. Incorporating this climate-driven trait variation significantly improved the fit of models that compared natives and invasives. Lastly, in invasives, ITV was most strongly explained by spatial heterogeneity in moisture, whereas solar energy explains more ITV in natives.ConclusionsOur results indicate that trait expression and ITV vary significantly between natives and invasives, and that this is mediated by climate. These findings suggest that although natives and invasives are functionally similar at the regional scale, invader success at local scales is contingent on climate.     

Intraspecific trait variation in three common grass species reveals finescale species adjustment to local environmental conditions

Aims Studies along environmental gradients have shown that intraspecific trait variation (ITV) may contribute considerably to community-level trait variation. However, we lack knowledge about how the extent of ITV varies on a local scale and whether a varying extent of ITV is related to differences in local environmental site and plant community characteristics.     

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.     

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

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

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

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

Intraspecific trait variation increases species diversity in a trait‐based grassland model

Intraspecific trait variation (ITV) is thought to play a significant role in community assembly, but the magnitude and direction of its influence are not well understood. Although it may be critical to better explain population persistence, species interactions, and therefore biodiversity patterns, manipulating ITV in experiments is challenging. We therefore incorporated ITV into a trait‐ and individual‐based model of grassland community assembly by adding variation to the plants’ functional traits, which then drive life‐history tradeoffs. Varying the amount of ITV in the simulation, we examine its influence on pairwise‐coexistence and then on the species diversity in communities of different initial sizes. We find that ITV increases the ability of the weakest species to invade most, but that this effect does not scale to the community level, where the primary effect of ITV is to increase the persistence and abundance of the competitively‐average species. Diversity of the initial community is also of critical importance in determining ITV's efficacy; above a threshold of interspecific diversity, ITV does not increase diversity further. For communities below this threshold, ITV mainly helps to increase diversity in those communities that would otherwise be low‐diversity. These findings suggest that ITV actively maintains diversity by helping the species on the margins of persistence, but mostly in habitats of relatively low alpha and beta diversity.     

Individualistic responses of forest herb traits to environmental change

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

     

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

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

Light and warming drive forest understorey community development in different environments

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

Intraspecific trait variation improves the detection of deterministic community assembly processes in early successional forests,but not in late successional forests

Aims Intraspecific trait variation (ITV) has been increasingly recognized to play an important role in understanding the underlying processes influencing community assembly. However, gaps remain in our understanding of how incorporating ITV will influence the relative importance of deterministic (e.g. habitat filtering, limiting similarity) and stochastic processes in driving community assembly at different successional stages.     

Intraspecific trait variation of woody species reduced in a savanna community,southwest China

Plants deploy various ecological strategies in response to environmental heterogeneity. In many forest ecosystems, plants have been reported to have notable inter- and intra-specific trait variation, as well as clear phylogenetic signals, indicating that these species possess a degree of phenotypic plasticity to cope with habitat variation in the community. Savanna communities, however, grow in an open canopy structure and exhibit little species diversification, likely as a result of strong environmental stress. In this study, we hypothesized that the phylogenetic signals of savanna species would be weak, the intraspecific trait variation (ITV) would be low, and the contribution of intraspecific variation to total trait variance would be reduced, owing to low species richness, multiple stresses and relatively homogenous community structure. To test these hypotheses, we sampled dominant woody species in a dry-hot savanna in southwestern China, focusing on leaf traits related to adaptability of plants to harsh conditions (year-round intense radiation, low soil fertility and seasonal droughts). We found weak phylogenetic signals in leaf traits and low ITV (at both individual and canopy-layer levels). Intraspecific variation (including leaf-, layer- and individual-scales) contributed little to the total trait variance, whereas interspecific variation and variation in leaf phenology explained substantial variance. Our study suggests that intraspecific trait variation is reduced in savanna community. Furthermore, our findings indicate that classifying species by leaf phenology may help better understand how species coexist under similar habitats with strong stresses.     

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.     

A willow drawing from 1786: the earliest depiction of intraspecific trait variation in plants?

Background and AimsThe study of intraspecific trait variation (ITV) in plants has a long history, dating back to the fourth century BC. Its existence was widely acknowledged by the end of the 18th century, although systematic and experimental studies commenced only a century later. However, the historiography of ITV has many gaps, especially with regard to early observations and visual documents. This note identifies an early depiction of plant ITV.MethodsThe botanical works of Johann Wolfgang von Goethe (1749–1832), a German writer and naturalist, were subjected to close reading. This included all publications and unpublished sources related to botany between 1785 and 1832 (e.g. notes, drafts, diaries, letters, drawings). This material is accessible in the multi-volume historical-critical edition of Goethe’s studies in natural science (Leopoldina-Ausgabe).Key ResultsA diary entry from 9 September 1786 described changes in leaf morphology along an elevation gradient in the Bavarian Alps. The leaves of an unidentified species of willow (Spix sp.) and gentian (Gentiana sp.) were said to become narrower with increasing elevation; leaves also stood further apart on twigs, and the latter became thinner. A crude drawing of two willow twigs illustrated the differences. Goethe conjectured that the differences were due to environmental conditions.ConclusionsGoethe’s notes were anecdotal, and it is unclear whether the observed plant individuals actually belonged to the same species. Nevertheless, the notes represent an early and clear articulation of the hypothesis that changes in environmental conditions can cause ITV in a natural plant population. The drawing may be the earliest visual record of environmentally caused plant ITV in the wild.     

Trait means or variance—What determines plant species' local and regional occurrence in fragmented dry grasslands?

One of the few laws in ecology is that communities consist of few common and many rare taxa. Functional traits may help to identify the underlying mechanisms of this community pattern, since they correlate with different niche dimensions. However, comprehensive studies are missing that investigate the effects of species mean traits (niche position) and intraspecific trait variability (ITV, niche width) on species abundance. In this study, we investigated fragmented dry grasslands to reveal trait‐occurrence relationships in plants at local and regional scales. We predicted that (a) at the local scale, species occurrence is highest for species with intermediate traits, (b) at the regional scale, habitat specialists have a lower species occurrence than generalists, and thus, traits associated with stress‐tolerance have a negative effect on species occurrence, and (c) ITV increases species occurrence irrespective of the scale. We measured three plant functional traits (SLA = specific leaf area, LDMC = leaf dry matter content, plant height) at 21 local dry grassland communities (10 m × 10 m) and analyzed the effect of these traits and their variation on species occurrence. At the local scale, mean LDMC had a positive effect on species occurrence, indicating that stress‐tolerant species are the most abundant rather than species with intermediate traits (hypothesis 1). We found limited support for lower specialist occurrence at the regional scale (hypothesis 2). Further, ITV of LDMC and plant height had a positive effect on local occurrence supporting hypothesis 3. In contrast, at the regional scale, plants with a higher ITV of plant height were less frequent. We found no evidence that the consideration of phylogenetic relationships in our analyses influenced our findings. In conclusion, both species mean traits (in particular LDMC) and ITV were differently related to species occurrence with respect to spatial scale. Therefore, our study underlines the strong scale‐dependency of trait‐abundance relationships.     

Explaining variation in productivity requires intraspecific variability in plant height among communities

解释生产力的变异需要考虑群落间植物高度的种内变异 已有研究表明,种内性状变异在群落构建过程中具有重要作用,但是迄今为止关于种内性状变异对生态系统功能的贡献仍然知之甚少。我们在青藏高原的高寒草甸进行了为期4年的功能群去 除实验,以研究植物高度的种间和种内变异对生产力的相对重要性。将75个控制群落内的株高总 变异分为种间变异(TV_inter)和种内变异(ITV_within),将群落间的群落加权平均高度分解为固定群落加权平 均值(CWM_fixed)和群落间种内变异(ITV_among)。我们通过广义加性混合模型、模型筛选和结构方程模型评估了性状变异(即TV_inter、ITV_within、CWM_fixed和ITV_among)如何间接介导功能群去除后生产力的变化。研究结果表明,功能群去除不仅会直接引起生产力变化,同时还会通过改变种间和种内株高变异间接影响生产力(即CWM_fixed和ITV_among)。“选择效应”和“避荫综合征”都可能导致更高的CWM_fixed和ITV_among,从而起到调节生产力的作用。该研究结果说明,只考虑种间变异可能会低估植物群落功能结构在驱动生态系统过程中的贡献。     

Ecological constraints from incumbent clades drive trait evolution across the tree‐of‐life of freshwater macroinvertebrates

The rates of species and trait diversification vary across the Tree‐of‐Life and over time. Whereas species richness and clade age generally are decoupled, the correlation of accumulated trait diversity of clades (trait disparity) with clade age remains poorly explored. Total trait disparity may be coupled with clade age if the growth of disparity (disparification) within and across clades is continuous with time in an additive niche expansion process (linear‐cumulative model), or alternatively if the rate of trait disparification varies over time and decreases as ecological space becomes gradually saturated (disparity‐dependent model). Using a clock‐calibrated phylogenetic tree for 143 freshwater macroinvertebrate families and richness and trait databases covering > 6400 species, we measured trait disparity in 18 independent clades that successively transitioned to freshwater ecosystems and analyzed its relation with clade age. We found a positive correlation between clade age and total disparity within clades, but no relationship for most individual traits. Traits unique to freshwater lifestyle were highly variable within older clades, while disparity in younger clades shifted towards partially terrestrial lifestyles and saline tolerance to occupy habitats previously inaccessible or underutilized. These results argue that constraints from incumbent lineages limit trait disparity in younger clades that evolved for filling unoccupied regions of the trait space, which suggests that trait disparification may follow a disparity‐dependent model. Overall, we provide an empirical pattern that reveals the potential of the disparity‐dependent model for understanding fundamental processes shaping trait dynamics across the Tree‐of‐Life.     

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.