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.     

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.     

Incorporating intraspecific variation in tests of trait-based community assembly

Environmental filtering and niche differentiation are processes proposed to drive community assembly, generating nonrandom patterns in community trait distributions. Despite the substantial intraspecific trait variation present in plant communities, most previous studies of trait-based community assembly have used species mean trait values and therefore not accounted for intraspecific variation. Using a null model approach, I tested for environmental filtering and niche differentiation acting on three key functional traits??vegetative height, specific leaf area (SLA), and leaf dry matter content (LDMC)??in old-field plant communities. I also examined how accounting for intraspecific variation at the among-plot and individual levels affected the detection of nonrandom assembly patterns. Tests using fixed species mean trait values provided evidence of environmental filtering acting on height and SLA and niche differentiation acting on SLA. Including plot-level intraspecific variation increased the strength of these patterns, indicating an important role of intraspecific variation in community assembly. Tests using individual trait data indicated strong environmental filtering acting on all traits, but provided no evidence of niche differentiation, although these signals may have been obscured by the effects of dispersal limitation and spatial aggregation of conspecific individuals. There was also strong evidence of nonrandom assembly of individuals within single species, with the strength of environmental filtering varying among species. This study demonstrates that, while analyses using fixed species mean trait values can provide insights into community assembly processes, accounting for intraspecific variation provides a more complete view of communities and the processes driving their assembly.     

Trait coordination and environmental filters shape functional trait distributions of forest understory herbs

1. Understanding the drivers of trait selection is critical for resolving community assembly processes. Here, we test the importance of environmental filtering and trait covariance for structuring the functional traits of understory herbaceous communities distributed along a natural environmental resource gradient that varied in soil moisture, temperature, and nitrogen availability, produced by different topographic positions in the southern Appalachian Mountains.
2. To uncover potential differences in community‐level trait responses to the resource gradient, we quantified the averages and variances of both abundance‐weighted and unweighted values for six functional traits (vegetative height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, and leaf δ¹³C) using 15 individuals of each of the 108 species of understory herbs found at two sites in the southern Appalachians of western North Carolina, USA.
3. Environmental variables were better predictors of weighted than unweighted community‐level average trait values for all but height and leaf N, indicating strong environmental filtering of plant abundance. Community‐level variance patterns also showed increased convergence of abundance‐weighted traits as resource limitation became more severe.
4. Functional trait covariance patterns based on weighted averages were uniform across the gradient, whereas coordination based on unweighted averages was inconsistent and varied with environmental context. In line with these results, structural equation modeling revealed that unweighted community‐average traits responded directly to local environmental variation, whereas weighted community‐average traits responded indirectly to local environmental variation through trait coordination.
5. Our finding that trait coordination is more important for explaining the distribution of weighted than unweighted average trait values along the gradient indicates that environmental filtering acts on multiple traits simultaneously, with abundant species possessing more favorable combinations of traits for maximizing fitness in a given environment.

     

Community Functional Responses to Soil and Climate at Multiple Spatial Scales: When Does Intraspecific Variation Matter?

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.     

New method for rapid assessment of the functional composition of herbaceous plant communities

Abstract We propose a rapid sampling method to assess the functional composition of herbaceous plant communities without prior knowledge of the floristic composition. To determine the community‐level value of traits (‘aggregated trait values’) for a plant community, a standardized population‐centred method exists, but requires substantial manpower and reliable botanical knowledge. We tested an alternative method, the trait transect, using four subalpine pastures in the Beaufortain region (Northern French Alps) selected along a fertility gradient. We applied both methods to measure five commonly used ‘soft traits’ known to be responsive to soil nutrient availability: plant vegetative and reproductive height, specific leaf area, leaf dry matter and nitrogen contents. We tested whether the variation of these traits along the gradient detected with the population‐centred method was also detected with the trait transect. Both methods detected expected trends in the traits in response to the fertility gradient. The trait transect method was as efficient as the population‐centred method and is recommended as an appropriate tool for monitoring ecosystem changes in response to environmental conditions and management, especially in species‐rich communities.     

退化草地物种种内和种间功能性状变异对放牧的响应

研究草地植物功能性状变异对放牧干扰的响应机制,有助于解析草地生态系统群落构建和功能多样性维持机制及植物对环境的适应及权衡策略。以科尔沁沙地退化草地为对象,研究围封和放牧草地物种多种功能性状(植株高度、根系长度、叶面积、根面积、叶片干物质含量、地上干物质含量、根系干物质含量、比叶面积、比根长和根冠比)变异特征及其对放牧干扰的响应机制。结果表明: 退化草地物种种间性状变异明显高于种内变异,种间性状变异对总体变异的贡献率占比高达70.2%～95.1%,而种内变异仅为4.9%～29.8%,但群落构建中物种的种内变异仍不可忽视。放牧草地物种种间性状变异低于围封草地,且放牧草地种内变异增加而种间变异减小。放牧导致不耐牧的优良禾本科牧草叶面积和叶片干物质含量下降而比根长增加,但耐牧的杂类草则通过增加叶面积和叶片干物质含量并降低比根长来提高在群落中的优势地位。退化草地对放牧响应较为敏感的功能性状有叶面积、叶片干物质含量、比根长和比叶面积。植物叶片性状和根系性状自身及彼此间均呈显著正相关,但放牧会增强根系性状的协同效应而减弱叶片性状的协同效应。说明放牧会驱动植物个体和种群功能性状权衡策略发生改变,进而起到调控植物群落结构和功能的作用。     

Trait‐based community assembly of aquatic macrophytes along a water depth gradient in a freshwater lake

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Plant functional and phylogenetic turnover correlate with climate and land use in the Western Swiss Alps

Aims Understanding the relative importance of historical and environmental processes in the structure and composition of communities is one of the longest quests in ecological research. Increasingly, researchers are relying on the functional and phylogenetic β-diversity of natural communities to provide concise explanations on the mechanistic basis of community assembly and the drivers of trait variation among species. The present study investigated how plant functional and phylogenetic β-diversity change along key environmental and spatial gradients in the Western Swiss Alps.Methods Using the quadratic diversity measure based on six functional traits—specific leaf area, leaf dry matter content, plant height, leaf carbon content, leaf nitrogen content and leaf carbon to nitrogen content alongside a species-resolved phylogenetic tree—we relate variations in climate, spatial geographic, land use and soil gradients to plant functional and phylogenetic turnover in mountain communities of the Western Swiss Alps.Important findings Our study highlights two main points. First, climate and land-use factors play an important role in mountain plant community turnover. Second, the overlap between plant functional and phylogenetic turnover along these gradients correlates with the low phylogenetic signal in traits, suggesting that in mountain landscapes, trait lability is likely an important factor in driving plant community assembly. Overall, we demonstrate the importance of climate and land-use factors in plant functional and phylogenetic community turnover and provide valuable complementary insights into understanding patterns of β-diversity along several ecological gradients.     

Leaf-trait responses to environmental gradients in moorland communities: contribution of intraspecific variation,species replacement and functional group replacement

The values of many important traits of plants in a community change along environmental gradients. Such changes may involve intraspecific variation and replacement by species that have different trait values. We hypothesized that they also involve the variation within and among functional groups (FGs) to the environmental dependence of trait values at the community level. We studied environmental dependence of trait values in 27 moorlands at various scales and analyzed to what extent intraspecific variation, species replacement within FGs and FG replacement contribute to the gradient of community trait values. The community structure in moorlands was influenced mainly by two environmental factors: temperature and water condition. Plants inhabiting sites with low temperature and low-pH generally tended to have lower maximum leaf height, greater leaf mass per area, and smaller leaf size. At the community level, site-mean of maximum leaf height and leaf size generally increased with increasing temperature and water pH. Our analysis demonstrated that the relative contributions of intraspecific variation, species replacement within FGs and FG replacement differed depending on combinations of the traits and environments. The contribution of FG replacement varied considerably among cases (0.6–34.5 %). Species replacement within FGs, which has received little attention in previous studies, was most responsible for the community-level changes (31.6–65.3 %) and intraspecific variation also made a large contribution (22.9–57.9 %). Understanding such various mechanisms involving intraspecific variation and species replacement should help us better predict how the structure and functioning of moorland plant communities will respond to climate change.     

黔中喀斯特山区植物功能性状的变异与关联

认识植物功能性状随演替进展的变化规律和不同性状之间的关系, 有助于从功能生态学的角度来理解群落演替。该文调查了位于贵州省中部的普定县喀斯特山区26个样方的物种组成, 这些样方分别处于灌丛、落叶阔叶林、落叶常绿混交林3个演替阶段; 测量了分布于该区域的82种木本植物的3个功能性状值(叶面积、比叶面积、最大高度); 根据物种在样方中的多度加权计算得到26个样方的性状平均值, 在此基础上分析了随演替进展植物功能性状在群落水平上的变异格局和不同性状之间的相关性; 采用性状梯度分析法分析了各阶段优势物种的功能性状在群落间(β组分)和群落内(α组分)的变异格局及相关性。结果表明: 1)沿灌丛→落叶阔叶林→落叶常绿混交林这一演替顺序, 群落平均叶面积和平均高度逐渐增加, 而群落平均比叶面积则逐渐变小; 2)就群落平均性状值而言, 叶面积与最大高度呈较强的正相关关系, 最大高度与比叶面积、叶面积与比叶面积均呈较强的负相关关系; 3)物种功能性状的α组分之间没有显著的相关关系, 而β组分之间相关性显著。这说明: 随演替的进展, 群落优势物种对环境的适应策略由高速生长转向提高资源利用效率, 而同一群落内共存的物种采取不同的性状组合来适应共同的群落环境。     

Plant communities on infertile soils are less sensitive to climate change

Background and Aims Much evidence suggests that plant communities on infertile soils are relatively insensitive to increased water deficit caused by increasing temperature and/or decreasing precipitation. However, a multi-decadal study of community change in the western USA does not support this conclusion. This paper tests explanations related to macroclimatic differences, overstorey effects on microclimate, variation in soil texture and plant functional traits.Methods A re-analysis was undertaken of the changes in the multi-decadal study, which concerned forest understorey communities on infertile (serpentine) and fertile soils in an aridifying climate (southern Oregan) from 1949–1951 to 2007–2008. Macroclimatic variables, overstorey cover and soil texture were used as new covariates. As an alternative measure of climate-related change, the community mean value of specific leaf area was used, a functional trait measuring drought tolerance. We investigated whether these revised analyses supported the prediction of lesser sensitivity to climate change in understorey communities on infertile serpentine soils.Key Results Overstorey cover, but not macroclimate or soil texture, was a significant covariate of community change over time. It strongly buffered understorey temperatures, was correlated with less change and averaged >50 % lower on serpentine soils, thereby counteracting the lower climate sensitivity of understorey herbs on these soils. Community mean specific leaf area showed the predicted pattern of less change over time in serpentine than non-serpentine communities.Conclusions Based on the current balance of evidence, plant communities on infertile serpentine soils are less sensitive to changes in the climatic water balance than communities on more fertile soils. However, this advantage may in some cases be lessened by their sparser overstorey cover.     

Using plant functional traits to explain community composition across a strong environmental filter in Australian alpine snowpatches

Environmental filters act to limit the local community assemblage from the regional species pool by restricting the viable trait states that can occur there. In alpine snowpatches, the timing of snowmelt is a strong environmental filter. In coming decades, the strength of this filter is likely to relax with global climate change. We used three continuous plant functional traits (leaf area, plant height, seed mass) and their divergence (using the FD_var index) to document current patterns of community assembly and predict plant community responses to future environmental filters in alpine snowpatch vegetation. The community trait-weighted mean for leaf area and height, but not seed mass, was significantly higher in early snowmelt zones relative to mid and late melting zones across all snowpatches. Mean FD_var for height (but not leaf area or seed mass), by contrast, was substantially lower in early snowmelt zones, indicating that species growing in early melt zones are consistently taller than those growing in other zones. These results suggest that if climate change leads to earlier snowmelt and hence, a longer growing season, taller (more competitive) species with larger leaf areas (more productive) may replace short species in snowpatches as these plant communities re-assemble in response to changing environmental filters.     

黄土高原典型小流域草地群落功能性状对土壤水分的响应

植物功能性状可以响应生境的变化并决定生态系统的功能,探究植物功能性状间的关系及其随土壤有效水分梯度的变化规律,对认识不同水分条件下植被在群落水平碳水代谢关系和维持水分平衡的生理生态学机制具有重要意义。以甘肃定西典型半干旱黄土小流域草地群落为研究对象,采用排序分析和回归拟合方法,分析了30个代表性草地样地中7个植物功能性状加权均值对土壤有效水分的响应以及响应性状间的相关关系。结果显示:(1)7个性状中,除叶宽与土壤有效水分无明显相关外,叶长、株高、叶面积、比叶面积、叶厚和叶干物质含量均与土壤有效水分显著性相关,可识别为草地在群落水平对土壤水分的响应性状。(2)草地群落通过降低株高,减小叶长、叶面积和比叶面积,增加叶厚和叶干物质含量以适应土壤有效水分减少;其中叶干物质含量的解释度最大,是土壤水分的最优响应性状。(3)除叶厚与叶长无显著相关外,其余功能性状均存在显著相关,说明草地群落的功能性状在土壤水分梯度上已基本形成了一个相互权衡或协同变化的功能性状组合。     

Biotic Regulation of CO<Subscript>2</Subscript> Uptake–Climate Responses: Links to Vegetation Properties

Identifying the plant traits and patterns of trait distribution in communities that are responsible for biotic regulation of CO₂ uptake–climate responses remains a priority for modeling terrestrial C dynamics. We used remotely sensed estimates of gross primary productivity (GPP) from plots planted to different combinations of perennial grassland species in order to determine links between traits and GPP–climate relationships. Climatic variables explained about 50% of the variance in temporal trends in GPP despite large variation in CO₂ uptake among seasons, years, and plots of differing composition. GPP was highly correlated with contemporary changes in net radiation (Rn) and precipitation deficit (potential evapotranspiration minus precipitation) but was negatively correlated with precipitation summed over 210 days prior to flux measurements. Plots differed in GPP–Rn and GPP–water (deficit, precipitation) relationships. Accounting for differences in GPP–climate relationships explained an additional 11% of variance in GPP. Plot differences in GPP–Rn and GPP–precipitation slopes were linked to differences in community-level light-use efficiency (GEE*). Plot differences in GPP–deficit slopes were linked to differences in a species abundance-weighted index of specific leaf area (SLA). GEE* and weighted SLA represent vegetation properties that may regulate how CO₂ uptake responds to climatic variation in grasslands.     

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.

     

A Multidimensional Functional Trait Approach Reveals the Imprint of Environmental Stress in Mediterranean Woody Communities

Water availability is one of the most important factors determining species distribution, plant community structure and ecosystem functioning. We explore how the functional structure of Mediterranean woody plant communities varies along a regional gradient of aridity in the Andalusian region (south Spain). We question whether communities located in more arid sites show more similarity in their functional structure when compared with communities located in wetter sites or whether, instead, there is divergence in their functional spaces. We selected five aridity zones (three sampling sites per zone) and measured 13 traits of different functional dimensions (including leaf, stem and root traits) in 74 woody plant species. We quantified functional space differences using the n-dimensional niche space approach (hypervolume). We found a larger functional space for the wetter communities compared with the more arid communities, which showed greater overlap of the trait space occupation. Our results indicate that aridity acts as a key abiotic filter affecting various metrics of the community trait structure, in accordance with the plant economics spectrum. We have also documented consistent variation in the functional space, supporting lower functional diversity under more harsh climatic conditions. The trend of functional space variation along the aridity gradient was different when considering traits from only one plant organ. Thus, the filtering process driving the functional structure of the communities studied here largely depends on the trait axis considered; for example, the root dimension showed considerable variation in wet environments, whereas the leaf dimension exhibited a larger functional space in the drier habitats.     

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

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

Plant functional traits and community assembly along interacting gradients of productivity and fragmentation

Quantifying the association of plant functional traits to environmental gradients is a promising approach for understanding and projecting community responses to land use and climatic changes. Although habitat fragmentation and climate are expected to affect plant communities interactively, there is a lack of empirical studies addressing trait associations to fragmentation in different climatic regimes.In this study, we analyse data on the key functional traits: specific leaf area (SLA), plant height, seed mass and seed number. First, we assess the evidence for the community assembly mechanisms habitat filtering and competition at different spatial scales, using several null-models and a comprehensive set of community-level trait convergence and divergence indices. Second, we analyse the association of community-mean traits with patch area and connectivity along a south–north productivity gradient.We found clear evidence for trait convergence due to habitat filtering. In contrast, the evidence for trait divergence due to competition fundamentally depended on the null-model used. When the null-model controlled for habitat filtering, there was only evidence for trait divergence at the smallest sampling scale (0.25 m × 0.25 m). All traits varied significantly along the S–N productivity gradient. While plant height and SLA were consistently associated with fragmentation, the association of seed mass and seed number with fragmentation changed along the S–N gradient.Our findings indicate trait convergence due to drought stress in the arid sites and due to higher productivity in the mesic sites. The association of plant traits to fragmentation is likely driven by increased colonization ability in small and/or isolated patches (plant height, seed number) or increased persistence ability in isolated patches (seed mass).Our study provides the first empirical test of trait associations with fragmentation along a productivity gradient. We conclude that it is crucial to study the interactive effects of different ecological drivers on plant functional traits.