Potential trajectories of old‐growth Neotropical forest functional composition under climate change

Quantifying relationships between plant functional traits and abiotic gradients is valuable for evaluating potential responses of forest communities to climate change. However, the trajectories of change expected to occur in tropical forest functional characteristics as a function of future climate variation are largely unknown. We modeled community level trait values of Costa Rican rain forests as a function of current and future climate, and quantified potential changes in functional composition. We calculated per‐plot community weighted mean (CWM) trait values for leaf area (LA), specific leaf area (SLA), leaf dry matter content (LDMC), leaf nitrogen (N) and phosphorus (P) content, and wood basic specific gravity (WSG), for tree and palm species in 127 0.25 ha plots. We modeled the response of CWM traits to current temperature and precipitation gradients using generalized additive modeling. We then predicted and mapped CWM traits values under current and future climate, and quantified potential changes under a global warming scenario (RCP8.5, year 2050). We calculated the area within the multi trait functional space occupied by forest plots under both current and future climate, and determined potential changes in functional space occupied by forest plots. Overall, precipitation predicted CWM traits better than temperature. Models indicated increases in CWM SLA, N and P, and a decrease in CWM LDMC under climate change. Lowland forest communities converged on a single direction of change towards more acquisitive CWM trait values, indicating a change in forest functional composition resulting from a changed climate. Functional space occupied by forest plots was reduced by 50% under the future climate. Functional composition changes may have further effects on forests ecosystem services. Assessing functional trait spatial‐gradients can help bridge the gap between species‐based biogeography and biogeochemical approaches to strengthen biodiversity and ecosystem services conservation efforts.     

Inclusion of ecologically based trait variation in plant functional types reduces the projected land carbon sink in an earth system model

Earth system models demonstrate large uncertainty in projected changes in terrestrial carbon budgets. The lack of inclusion of adaptive responses of vegetation communities to the environment has been suggested to hamper the ability of modeled vegetation to adequately respond to environmental change. In this study, variation in functional responses of vegetation has been added to an earth system model (ESM) based on ecological principles. The restriction of viable mean trait values of vegetation communities by the environment, called ‘habitat filtering’, is an important ecological assembly rule and allows for determination of global scale trait–environment relationships. These relationships were applied to model trait variation for different plant functional types (PFTs). For three leaf traits (specific leaf area, maximum carboxylation rate at 25 °C, and maximum electron transport rate at 25 °C), relationships with multiple environmental drivers, such as precipitation, temperature, radiation, and CO₂, were determined for the PFTs within the Max Planck Institute ESM. With these relationships, spatiotemporal variation in these formerly fixed traits in PFTs was modeled in global change projections (IPCC RCP8.5 scenario). Inclusion of this environment‐driven trait variation resulted in a strong reduction of the global carbon sink by at least 33% (2.1 Pg C yr^?1) from the 2nd quarter of the 21st century onward compared to the default model with fixed traits. In addition, the mid‐ and high latitudes became a stronger carbon sink and the tropics a stronger carbon source, caused by trait‐induced differences in productivity and relative respirational costs. These results point toward a reduction of the global carbon sink when including a more realistic representation of functional vegetation responses, implying more carbon will stay airborne, which could fuel further climate change.     

Scaling up the diversity–resilience relationship with trait databases and remote sensing data: the recovery of productivity after wildfire

Understanding the mechanisms underlying ecosystem resilience – why some systems have an irreversible response to disturbances while others recover – is critical for conserving biodiversity and ecosystem function in the face of global change. Despite the widespread acceptance of a positive relationship between biodiversity and resilience, empirical evidence for this relationship remains fairly limited in scope and localized in scale. Assessing resilience at the large landscape and regional scales most relevant to land management and conservation practices has been limited by the ability to measure both diversity and resilience over large spatial scales. Here, we combined tools used in large‐scale studies of biodiversity (remote sensing and trait databases) with theoretical advances developed from small‐scale experiments to ask whether the functional diversity within a range of woodland and forest ecosystems influences the recovery of productivity after wildfires across the four‐corner region of the United States. We additionally asked how environmental variation (topography, macroclimate) across this geographic region influences such resilience, either directly or indirectly via changes in functional diversity. Using path analysis, we found that functional diversity in regeneration traits (fire tolerance, fire resistance, resprout ability) was a stronger predictor of the recovery of productivity after wildfire than the functional diversity of seed mass or species richness. Moreover, slope, elevation, and aspect either directly or indirectly influenced the recovery of productivity, likely via their effect on microclimate, while macroclimate had no direct or indirect effects. Our study provides some of the first direct empirical evidence for functional diversity increasing resilience at large spatial scales. Our approach highlights the power of combining theory based on local‐scale studies with tools used in studies at large spatial scales and trait databases to understand pressing environmental issues.     

Remotely sensed between‐individual functional trait variation in a temperate forest

1. Trait‐based ecology holds the promise to explain how plant communities work, for example, how functional diversity may support community productivity. However, so far it has been difficult to combine field‐based approaches assessing traits at the level of plant individuals with limited spatial coverage and approaches using remote sensing (RS) with complete spatial coverage but assessing traits at the level of vegetation pixels rather than individuals. By delineating all individual‐tree crowns within a temperate forest site and then assigning RS‐derived trait measures to these trees, we combine the two approaches, allowing us to use general linear models to estimate the influence of taxonomic or environmental variation on between‐ and within‐species variation across contiguous space.
2. We used airborne imaging spectroscopy and laser scanning to collect individual‐tree RS data from a mixed conifer‐angiosperm forest on a mountain slope extending over 5.5 ha and covering large environmental gradients in elevation as well as light and soil conditions. We derived three biochemical (leaf chlorophyll, carotenoids, and water content) and three architectural traits (plant area index, foliage‐height diversity, and canopy height), which had previously been used to characterize plant function, from the RS data. We then quantified the contributions of taxonomic and environmental variation and their interaction to trait variation and partitioned the remaining within‐species trait variation into smaller‐scale spatial and residual variation. We also investigated the correlation between functional trait and phylogenetic distances at the between‐species level. The forest consisted of 13 tree species of which eight occurred in sufficient abundance for quantitative analysis.
3. On average, taxonomic variation between species accounted for more than 15% of trait variation in biochemical traits but only around 5% (still highly significant) in architectural traits. Biochemical trait distances among species also showed a stronger correlation with phylogenetic distances than did architectural trait distances. Light and soil conditions together with elevation explained slightly more variation than taxonomy across all traits, but in particular increased plant area index (light) and reduced canopy height (elevation). Except for foliage‐height diversity, all traits were affected by significant interactions between taxonomic and environmental variation, the different responses of the eight species to the within‐site environmental gradients potentially contributing to the coexistence of the eight abundant species.
4. We conclude that with high‐resolution RS data it is possible to delineate individual‐tree crowns within a forest and thus assess functional traits derived from RS data at individual level. With this precondition fulfilled, it is then possible to apply tools commonly used in field‐based trait ecology to partition trait variation among individuals into taxonomic and potentially even genetic variation, environmental variation, and interactions between the two. The method proposed here presents a promising way of assessing individual‐based trait information with complete spatial coverage and thus allowing analysis of functional diversity at different scales. This information can help to better understand processes shaping community structure, productivity, and stability of forests.

     

吉林蛟河针阔混交林功能性状分布格局及其对地形因素的响应

森林群落的构建过程及其内在机制是生态学研究的热点问题。植物功能性状是指能够代表植物的生活史策略,反映植物对环境变化响应的一系列植物属性。通过植物功能性状的分布格局及其对环境因素的响应有助于推测群落的构建过程及其内在作用机制。以吉林蛟河21.12hm²温带针阔混交林样地为研究对象,采集并测量了样地内34种木本植物的6种不同的功能性状。以20m×20m的样方为研究单元,通过计算平均成对性状距离指数（mean pairwise trait distance;PW）和平均最近邻体性状距离指数（mean nearest neighbor trait distance;NN）来探讨群落中单个性状和综合性状的分布格局。同时结合地形因子采用回归分析探讨功能性状的分布格局对局域生境变化的响应。基于PW的结果显示：单个性状中除叶面积外,其余性状的分布格局均为聚集分布多于离散分布;基于NN的结果显示：除叶面积和最大树高外,其余性状的分布格局为聚集分布多于离散分布。此外,由6种单个性状组成的综合性状的分布格局同样为聚集分布多于离散分布。基于回归分析的结果显示：森林群落中功能性状的分布格局受到海拔、坡度和坡向等因素的显著影响,而凹凸度的影响则不显著。研究结果表明包括环境过滤和生物相互作用的非随机过程能够影响温带针阔混交林的群落构建过程,中性过程对该区域群落构建过程的影响不显著。     

鼎湖山南亚热带常绿阔叶林叶功能性状沿群落垂直层次的种内变异

探究功能性状沿着环境梯度如何变化一直以来是基于性状的群落生态学的核心问题之一。尽管功能性状存在种内和种间变异, 但种内变异沿环境梯度如何变化仍有待探究。本文以鼎湖山南亚热带常绿阔叶林1.44 ha塔吊样地内16个树种的2,820个个体为研究对象, 探究4种叶功能性状(比叶面积、叶干物质含量、叶厚度和叶面积)沿群落垂直层次的种内变异。首先, 利用随机效应线性模型量化塔吊样地内的种内变异和种间变异; 其次, 利用Kmeans函数将森林的垂直层次划分为灌木层、亚冠层和林冠层, 并通过构建回归模型探究叶功能性状在群落垂直层次中的种内变异格局。最后, 应用混合线性模型和单因素方差分析的方法探究叶功能性状沿垂直层次的种内变异是否具有物种依赖性。结果表明: 在局域群落中, 并非所有叶功能性状的种内变异都低于种间变异; 叶功能性状在不同垂直层次的种内变异格局存在显著差异, 且种内变异与垂直范围呈正相关; 叶功能性状的种内变异具有较强的物种依赖性, 因此树种差异相对于小环境解释了更多的性状变异; 此外, 不同叶功能性状的种内变异沿垂直层次的变化趋势并不一致。本研究发现种内变异对于物种共存具有重要作用。     

Intraspecific functional trait variability across different spatial scales: a case study of two dominant trees in Korean pine broadleaved forest

Intraspecific functional trait variability plays an important role in the response of plants to environmental changes. However, it is still unclear how the variability differs across three nested spatial scales (individual, plot, and site) and which determinants (climatic, soil, and ontogenetic variables) shape the trait variability. Along a latitudinal gradient in Korean pine broadleaved forest of northeast China, we quantified the extent of intraspecific variability of four functional traits in two dominant trees Pinus koraiensis and Fraxinus mandshurica at eight sites, including specific leaf area, leaf dry matter content (morphological traits) and leaf nitrogen content, leaf phosphorus content (physiological traits). Results showed a large trait variation within and between species (coefficient variation: 6.07–23.3%). The leaf physiological traits of F. mandshurica and morphological traits of P. koraiensis were more responsive at site scale, while the morphological traits of F. mandshurica and physiological traits of P. koraiensis were more responsive at individual scale. In addition, abiotic and biotic factors explaining functional trait variation differ markedly between the two tree species, with physiological trait of F. mandshurica being more associated with climate and soil, while traits variability in P. koraiensis was not affected by climate, soil, and ontogeny, except for leaf phosphorus content. Overall, we can predict that the physiological traits of broadleaved species tend to be more sensitive to environmental changes, while pines are more sensitive to competition. It is critical to determine which spatial scale and trait type should be taken into account in predictive models of vegetation dynamics.     

Convergent elevation trends in canopy chemical traits of tropical forests

The functional biogeography of tropical forests is expressed in foliar chemicals that are key physiologically based predictors of plant adaptation to changing environmental conditions including climate. However, understanding the degree to which environmental filters sort the canopy chemical characteristics of forest canopies remains a challenge. Here, we report on the elevation and soil‐type dependence of forest canopy chemistry among 75 compositionally and environmentally distinct forests in nine regions, with a total of 7819 individual trees representing 3246 species collected, identified and assayed for foliar traits. We assessed whether there are consistent relationships between canopy chemical traits and both elevation and soil type, and evaluated the general role of phylogeny in mediating patterns of canopy traits within and across communities. Chemical trait variation and partitioning suggested a general model based on four interconnected findings. First, geographic variation at the soil‐Order level, expressing broad changes in fertility, underpins major shifts in foliar phosphorus (P) and calcium (Ca). Second, elevation‐dependent shifts in average community leaf dry mass per area (LMA), chlorophyll, and carbon allocation (including nonstructural carbohydrates) are most strongly correlated with changes in foliar Ca. Third, chemical diversity within communities is driven by differences between species rather than by plasticity within species. Finally, elevation‐ and soil‐dependent changes in N, LMA and leaf carbon allocation are mediated by canopy compositional turnover, whereas foliar P and Ca are driven more by changes in site conditions than by phylogeny. Our findings have broad implications for understanding the global ecology of humid tropical forests, and their functional responses to changing climate.     

Human‐aided admixture may fuel ecosystem transformation during biological invasions: theoretical and experimental evidence

Biological invasions can transform our understanding of how the interplay of historical isolation and contemporary (human‐aided) dispersal affects the structure of intraspecific diversity in functional traits, and in turn, how changes in functional traits affect other scales of biological organization such as communities and ecosystems. Because biological invasions frequently involve the admixture of previously isolated lineages as a result of human‐aided dispersal, studies of invasive populations can reveal how admixture results in novel genotypes and shifts in functional trait variation within populations. Further, because invasive species can be ecosystem engineers within invaded ecosystems, admixture‐induced shifts in the functional traits of invaders can affect the composition of native biodiversity and alter the flow of resources through the system. Thus, invasions represent promising yet under‐investigated examples of how the effects of short‐term evolutionary changes can cascade across biological scales of diversity. Here, we propose a conceptual framework that admixture between divergent source populations during biological invasions can reorganize the genetic variation underlying key functional traits, leading to shifts in the mean and variance of functional traits within invasive populations. Changes in the mean or variance of key traits can initiate new ecological feedback mechanisms that result in a critical transition from a native ecosystem to a novel invasive ecosystem. We illustrate the application of this framework with reference to a well‐studied plant model system in invasion biology and show how a combination of quantitative genetic experiments, functional trait studies, whole ecosystem field studies and modeling can be used to explore the dynamics predicted to trigger these critical transitions.     

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.     

Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain,Southwest China

Characterizing trait variation across different ecological scales in plant communities has been viewed as a way to gain insights into the mechanisms driving species coexistence. However, little is known about how changes in intraspecific and interspecific traits across sites influence species richness and community assembly, especially in understory herbaceous communities. Here we partitioned the variance of four functional traits (maximum height, leaf thickness, leaf area and specific leaf area) across four nested biological scales: individual, species, plot, and elevation to quantify the scale-dependent distributions of understory herbaceous trait variance. We also integrated the comparison of the trait variance ratios to null models to investigate the effects of different ecological processes on community assembly and functional diversity along a 1200-m elevational gradient in Yulong Mountain. We found interspecific trait variation was the main trait variation component for leaf traits, although intraspecific trait variation ranged from 10% to 28% of total variation. In particular, maximum height exhibited high plasticity, and intraspecific variation accounted for 44% of the total variation. Despite the fact that species composition varied across elevation and species richness decreased dramatically along the elevational gradient, there was little variance at our largest (elevation) scale in leaf traits and functional diversity remained constant along the elevational gradient, indicating that traits responded to smaller scale influences. External filtering was only observed at high elevations. However, strong internal filtering was detected along the entire elevational gradient in understory herbaceous communities, possibly due to competition. Our results provide evidence that species coexistence in understory herbaceous communities might be structured by differential niche-assembled processes. This approach--integrating different biological scales of trait variation--may provide a better understanding of the mechanisms involved in the structure of communities.     

Evolutionary dynamics of the leaf phenological cycle in an oak metapopulation along an elevation gradient

It has been predicted that environmental changes will radically alter the selective pressures on phenological traits. Long‐lived species, such as trees, will be particularly affected, as they may need to undergo major adaptive change over only one or a few generations. The traits describing the annual life cycle of trees are generally highly evolvable, but nothing is known about the strength of their genetic correlations. Tight correlations can impose strong evolutionary constraints, potentially hampering the adaptation of multivariate phenological phenotypes. In this study, we investigated the evolutionary, genetic and environmental components of the timing of leaf unfolding and senescence within an oak metapopulation along an elevation gradient. Population divergence, estimated from in situ and common‐garden data, was compared to expectations under neutral evolution, based on microsatellite markers. This approach made it possible (1) to evaluate the influence of genetic correlation on multivariate local adaptation to elevation and (2) to identify traits probably exposed to past selective pressures due to the colder climate at high elevation. The genetic correlation was positive but very weak, indicating that genetic constraints did not shape the local adaptation pattern for leaf phenology. Both spring and fall (leaf unfolding and senescence, respectively) phenology timings were involved in local adaptation, but leaf unfolding was probably the trait most exposed to climate change‐induced selection. Our data indicated that genetic variation makes a much smaller contribution to adaptation than the considerable plastic variation displayed by a tree during its lifetime. The evolutionary potential of leaf phenology is, therefore, probably not the most critical aspect for short‐term population survival in a changing climate.     

Patch isolation and shape predict plant functional diversity in a naturally fragmented forest

Aims It is known that taxonomic diversity can be predicted by the spatial configuration of the habitat, in particular by its area and degree of isolation. However, taxonomic diversity is a poor predictor of ecosystem functioning. While functional diversity is strongly linked to the functionality and stability of ecosystems, little is known about how changes in the spatial configuration of the habitat affect functional diversity. In this study, we evaluated whether the spatial configuration of forest patches predicts the functional diversity of plants in a fragmented forest.Methods Five functional leaf traits (leaf dry matter content, leaf punch force, specific leaf area, leaf size and leaf thickness) were measured for 23 dominant plant species in 20 forest patches in a naturally fragmented forest on the Yucatan Peninsula. Abundance-weighted multivariate and individual trait metrics of functional diversity were calculated and correlated with size, degree of isolation and the shape of forest patches.Important findings Patch shape was negatively correlated with multivariate and individual trait (leaf dry matter content and leaf size) metrics of functional diversity. Patch isolation measures were also negatively correlated with individual trait (leaf dry matter content, leaf punch force and leaf size) metrics of functional diversity. In other words, greater patch shape irregularity and isolation degree impoverish plant functional variability. This is the first report of the negative effects of patch shape irregularity and isolation on the functional diversity of plant communities in a forest that has been fragmented for a long time.     

Elevation,Topography, and Edge Effects Drive Functional Composition of Woody Plant Species in Tropical Montane Forests

Tropical montane forests comprise heterogeneous environments along natural gradients of topography and elevation. Human‐induced edge effects further increase the environmental heterogeneity in these forests. The simultaneous effects of natural and human‐induced gradients on the functional diversity of plant leaf traits are poorly understood. In a tropical montane forest in Bolivia, we studied environmental gradients associated with elevation (from 1900 m to 2500 m asl), topography (ridge and gorge), and edge effects (forest edge vs. forest interior), and their relationship with leaf traits and resource‐use strategies. First, we investigated associations of environmental conditions (soil properties and microclimate) with six leaf traits, measured on 119 woody plant species. Second, we evaluated changes in functional composition with community‐weighted means and functional structure with multidimensional functional diversity indices (FRic, FEve and FDiv). We found significant associations between leaf traits and soil properties in accordance with the trade‐off between acquisition and conservation of resources. Functional composition of leaf traits shifted from the dominance of acquisitive species in habitats at low altitudes, gorges, and forest interior to the dominance of conservative species in habitats at high altitudes, ridges, and forest edges. Functional structure was only weakly associated with the environmental gradients. Natural and human‐induced environmental gradients, especially soil properties, are important for driving leaf traits and resource‐use strategies of woody plants. Nevertheless, weak associations between functional structure and environmental gradients suggest a high redundancy of functional leaf traits in this tropical montane forest.     

格氏栲天然林林窗植物群落功能性状的变异

林窗是森林更新演替的重要环节, 揭示林窗环境下功能性状变异来源及其相对贡献, 有助于阐明植物对林窗环境的响应。该研究以中亚热带格氏栲(Castanopsis kawakamii)天然林为对象, 设置9个不同大小的林窗样地, 运用方差分解探讨林窗、物种和个体对叶性状变异的相对贡献, 采用线性回归分析不同大小林窗下群落性状变化及种间和种内性状变异的重要性。研究发现: (1)格氏栲天然林林窗植物比叶面积、叶干物质含量、叶厚和叶绿素含量由种间性状变异主导, 叶氮含量由种内性状变异主导, 叶磷含量受林窗大小影响最大。(2)群落叶磷含量与林窗大小具有显著正相关关系, 土壤温度和水解氮含量对群落叶磷含量具有显著正效应, 土壤有效磷含量具有显著负效应。(3)沿林冠开放度的群落叶磷含量变化主要由种内性状变异引起, 优势种扮演着重要角色。结果表明, 格氏栲天然林林窗环境下植物功能性状仍以种间性状变异为主(平均41%), 但沿林窗环境梯度的群落性状变化主要源自种内性状变异, 通过植物表型可塑性响应环境改变, 优势种作用明显。     

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

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

Intraspecific trait variation across elevation predicts a widespread tree species' climate niche and range limits

Global change is widely altering environmental conditions which makes accurately predicting species range limits across natural landscapes critical for conservation and management decisions. If climate pressures along elevation gradients influence the distribution of phenotypic and genetic variation of plant functional traits, then such trait variation may be informative of the selective mechanisms and adaptations that help define climatic niche limits. Using extensive field surveys along 16 elevation transects and a large common garden experiment, we tested whether functional trait variation could predict the climatic niche of a widespread tree species (Populus angustifolia) with a double quantile regression approach. We show that intraspecific variation in plant size, growth, and leaf morphology corresponds with the species' total climate range and certain climatic limits related to temperature and moisture extremes. Moreover, we find evidence of genetic clines and phenotypic plasticity at environmental boundaries, which we use to create geographic predictions of trait variation and maximum values due to climatic constraints across the western US. Overall, our findings show the utility of double quantile regressions for connecting species distributions and climate gradients through trait‐based mechanisms. We highlight how new approaches like ours that incorporate genetic variation in functional traits and their response to climate gradients will lead to a better understanding of plant distributions as well as identifying populations anticipated to be maladapted to future environments.     

Functional trait variation of forest understorey plant communities across Europe

Global environmental changes are expected to alter the functional characteristics of understorey herb-layer communities, potentially affecting forest ecosystem functioning. However, little is known about what drives the variability of functional traits in forest understories. Here, we assessed the role of different environmental drivers in shaping the functional trait distribution of understorey herbs in fragmented forests across three spatial scales. We focused on 708 small, deciduous forest patches located in 16 agricultural landscape windows, spanning a 2500-km macroclimatic gradient across the temperate forest biome in Europe. We estimated the relative effect of patch-scale, landscape-scale and macroclimatic variables on the community mean and variation of plant height, specific leaf area and seed mass. Macroclimatic variables (monthly temperature and precipitation extremes) explained the largest proportion of variation in community trait means (on average 77% of the explained variation). In contrast, patch-scale factors dominated in explaining community trait variation (on average 68% of the explained variation). Notably, patch age, size and internal heterogeneity had a positive effect on the community-level variability. Landscape-scale variables explained only a minor part of the variation in both trait distribution properties. The variation explained by shared combinations of the variable groups was generally negligible. These findings highlight the importance of considering multiple spatial scales in predictions of environmental-change effects on the functionality of forest understories. We propose that forest management sustainability could benefit from conserving larger, historically continuous and internally heterogeneous forest patches to maximise ecosystem service diversity in rural landscapes.