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 assembly in plant assemblages and its modulation by productivity and disturbance

Understanding how communities assemble is a key challenge in ecology. Conflicting hypotheses suggest that plant traits within communities should show divergence to reflect strategies to reduce competition or convergence to reflect strong selection for the environmental conditions operating. Further hypotheses suggest that plant traits related to productivity show convergence within communities, but those related to disturbance show divergence. Data on functional diversity (FD_var) of 12 traits from 30 communities ranging from arable fields, mown and grazed grasslands to moorland and woodland were employed to test this using randomisations tests and correlation and regression analysis. No traits showed consistent significant convergence or divergence in functional diversity. When correlated to measures of the environment, the most common pattern was for functional diversity to decline (7 out of 12 traits) and the degree of convergence (7 out of 12 traits) to increase as the levels of productivity (measured as primary productivity, soil nitrogen release and vegetation C:N) and disturbance increased. Convergence or a relationship between functional diversity and the environment was not seen for a number of important traits, such as LDMC and SLA, which are considered as key predictors of ecosystem function. The analysis indicates that taking into account functional diversity within a system may be a necessary part of predicting the relationship between plant traits and ecosystem function, and that this may be of particular importance within less productive and less disturbed systems.     

环境因子对海岛植物茎、叶功能性状的影响

植物功能性状与环境之间的关系是功能性状研究的重点,海岛作为独特的生态系统,其植物功能性状必然和大陆存在差异。为了明确海岛植被的生态适应机制,该文以平潭岛森林群落为研究对象,通过测定茎、叶10个功能性状,以及地形和土壤9个环境因子,探讨了植物功能性状之间的权衡关系,分析了环境因子对海岛植物功能性状的影响。结果表明:(1)比叶面积(SLA)与叶氮含量(LNC)、叶磷含量(LPC)呈正相关,与叶厚度(LT)、叶干物质含量(LDMC)、茎组织密度(STD)、叶碳含量(LCC)呈负相关; LDMC与LNC、茎氮含量(SNC)呈负相关; LT与STD呈正相关,与LNC和LPC呈负相关; LPC与LNC、SNC呈正相关;茎和叶C、N含量均呈正相关。(2)土壤有机质和TN是海岛植物功能性状的主要土壤影响因子。然而,由于土壤中磷含量的缺乏,LNC、茎磷含量(SPC)、SNC均与土壤全磷呈正相关; LDMC与土壤全氮呈正相关;STD与土壤有机质呈正相关; SLA随着土壤pH的增加而增加。(3)坡位和坡度是海岛植物功能性状的主要地形影响因子。SLA、SPC随着海拔上升而下降; STD、LDMC随着海拔和坡度增大而增大; LNC、LPC阴坡大于阳坡。该研究为海岛植被修复和重建提供了参考依据。     

Grazing increases functional richness but not functional divergence in Tibetan alpine meadow plant communities

Plant community diversity and ecosystem function are conditioned by competition among co-occurring species for multiple resources. Previous studies suggest that removal of standing biomass by grazing decreases competition for light, but coincident grazing effects on competition for soil nutrients remain largely unknown in Tibetan rangelands where grazing tends to deplete soil phosphorus availability. We measured five functional traits indicative of plant productivity and stoichiometry leaf carbon concentration (LCC), leaf nitrogen concentration (LNC), leaf phosphorus concentration (LPC), specific leaf area (SLA), leaf dry matter content (LDMC) for component species of plant communities in grazed and ungrazed plots in five Tibetan alpine meadows. We examined the diversity of traits singly Rao index of functional diversity (FDrao) and in aggregate functional richness (FRic), functional divergence (FDiv), and functional evenness (FEve) in response to grazing. We tested whether foliar trait diversity increases with nutrient competition but decreases with light competition when competitive exclusion is reduced by grazing. The FDrao of LPC significantly increased under grazing, but FDrao for LCC, LNC and SLA tended to decrease. The FDrao of LDMC increased at the drier site but decreased at the wettest site. There was a strong negative association between increase in FDrao of LPC and decrease in soil nutrients, especially soil phosphorus availability. The FRic for all five traits together increased with species diversity following grazing, but neither FDiv nor FEve differed significantly between grazed and ungrazed plots at most sites. Grazing in Tibetan alpine meadows tends to increase competition for soil phosphorus while decreasing competition for light, resulting in an increase in the functional richness in grazed plant communities without any significant changes in the overall functional diversity of foliar traits. Our study highlights the potential importance of grazing mediated competition for multiple resources in alpine meadow ecosystems.     

Similarity of plant functional traits and aggregation pattern in a subtropical forest

The distribution of species and communities in relation to environmental heterogeneity is a central focus in ecology. Co‐occurrence of species with similar functional traits is an indication that communities are determined in part by environmental filters. However, few studies have been designed to test how functional traits are selectively filtered by environmental conditions at local scales. Exploring the relationship between soil characteristics and plant traits is a step toward understanding the filtering hypothesis in determining plant distribution at local scale. Toward this end, we mapped all individual trees (diameter >1 cm) in a one‐ha subtropical forest of China in 2007 and 2015. We measured topographic and detailed soil properties within the field site, as well as plant leaf functional traits and demographic rates of the seven most common tree species. A second one‐ha study plot was established in 2015, to test and validate the general patterns that were drawn from first plot. We found that variation in species distribution at local scale can be explained by soil heterogeneity and plant functional traits. (From first plot). (1) Species dominant in habitats with high soil ammonium nitrogen and total phosphorus tended to have high specific leaf area (SLA) and relative growth rate (RGR). (2) Species dominant in low‐fertility habitats tended to have high leaf dry matter content (LDMC), ratio of chlorophyll a and b (ratioab), and leaf thickness (LT). The hypothesis that functional traits are selected in part by environmental filters and determine plant distribution at local scale was confirmed by the data of the first plot and a second regional site showed similar species distribution patterns.     

胡杨叶功能性状特征及其对地下水埋深的响应

叶片性状反映了植物对环境的适应能力及其自我调控能力。以塔里木干旱荒漠区建群种胡杨（Populus euphratica）为研究对象,通过分析自然生长状况下胡杨叶功能性状对地下水埋深（GWD）的响应及功能性状间的权衡关系,揭示胡杨对极端干旱荒漠环境的生态适应策略。结果表明：胡杨7个叶功能性状种内变异程度不同（9.20%-40.02%）,叶面积（LA）变异程度最大,叶干物质含量（LDMC）与叶片含水量（LWC）变异程度较低,GWD梯度上表现出较大的分化变异特征。叶性状在不同GWD之间差异显著（P<0.05）,与GWD呈极显著相关（P<0.01）。比叶面积（SLA）、LA、LWC与叶干重（LDM）呈极显著正相关（P<0.01）,与叶厚度（LT）、叶组织密度（LTD）、LDMC呈极显著的负相关（P<0.01）;LDMC与LT、LTD,LWC与LA、SLA呈极显著正相关（P<0.01）,反映胡杨通过叶性状间的相互调节与权衡来适应干旱荒漠环境。逐步回归分析表明LA、LT对GWD变化最敏感,可间接借助这2个性状来预测干旱荒漠区地下水埋深变化。随GWD降低,胡杨SLA、LA、LDM、LWC减小,而LT、LTD、LDMC增大,其由高生长速率、资源利用能力的开拓型策略转变为以增强自身养分储存、防御能力的保守型策略,拓宽了生态幅和增强其在干旱逆境的适合度。可见,极端干旱荒漠区胡杨形成了小的LA、SLA、LDM,大的LT、LDMC、LTD等一系列有利于减少水分散失、储存养分和增强耐旱能力的干旱性状组合,这可能是其适应干旱贫瘠环境的生态策略。     

Is leaf dry matter content a better predictor of soil fertility than specific leaf area?

Background and Aims

Specific leaf area (SLA), a key element of the ‘worldwide leaf economics spectrum’, is the preferred ‘soft’ plant trait for assessing soil fertility. SLA is a function of leaf dry matter content (LDMC) and leaf thickness (LT). The first, LDMC, defines leaf construction costs and can be used instead of SLA. However, LT identifies shade at its lowest extreme and succulence at its highest, and is not related to soil fertility. Why then is SLA more frequently used as a predictor of soil fertility than LDMC?

Methods

SLA, LDMC and LT were measured and leaf density (LD) estimated for almost 2000 species, and the capacity of LD to predict LDMC was examined, as was the relative contribution of LDMC and LT to the expression of SLA. Subsequently, the relationships between SLA, LDMC and LT with respect to soil fertility and shade were described.

Key Results

Although LD is strongly related to LDMC, and LDMC and LT each contribute equally to the expression of SLA, the exact relationships differ between ecological groupings. LDMC predicts leaf nitrogen content and soil fertility but, because LT primarily varies with light intensity, SLA increases in response to both increased shade and increased fertility.

Conclusions

Gradients of soil fertility are frequently also gradients of biomass accumulation with reduced irradiance lower in the canopy. Therefore, SLA, which includes both fertility and shade components, may often discriminate better between communities or treatments than LDMC. However, LDMC should always be the preferred trait for assessing gradients of soil fertility uncoupled from shade. Nevertheless, because leaves multitask, individual leaf traits do not necessarily exhibit exact functional equivalence between species. In consequence, rather than using a single stand-alone predictor, multivariate analyses using several leaf traits is recommended.     

Do Clonal and Bud Bank Traits Vary in Correspondence with Soil Properties and Resource Acquisition Strategies? Patterns in Alpine Communities in the Scandian Mountains

Plant traits associated with resource acquisition strategies (specific leaf area (SLA), leaf dry matter content (LDMC), leaf size and plant height) change along gradients of soil properties, being the most conservative in a resource-poor environment and the most dynamic in a resource-rich environment. Clonal attributes also vary along soil and other environmental conditions. We hypothesized that in alpine communities in the Scandian Mts. (1) the average composition of traits in a plant assemblage in terms of i) the predominance of different clonal growth organ types, ii) the number of buds in the bud bank, iii) the distribution of the bud-bank (above- and below ground), iv) the distance of lateral spread and v) the longevity of plant – offspring connections would change along a gradient of soil properties and (2) that this variation would be in correspondence with that of traits associated with resource acquisition strategies (SLA, LDMC, leaf size and plant height). Analysis of clonal and bud bank traits for species of alpine communities supported our first hypothesis: with decreasing soil quality the most common clonal growth organs were rhizomes, and there was a predominance of perennial bud banks located at the soil surface or below-ground, low rates of lateral spread and long persistence of plant – offspring connections. Our second hypothesis was partly supported. As predicted, at the level of the plant assemblage, these clonal and bud bank traits were positively associated with LDMC, and negatively with leaf size and plant height. These observations reinforce the hypotheses about trade-offs between acquisition and retention strategies in plants. The only result that was in contradiction with our expectations was the lack of correspondence between clonal and bud bank traits and SLA that could be attributed to errors associated to the measurement of the area of narrow and small leaves or to the dependence of the SLA index on species-specific morphological attributes.     

生境异质性对鼎湖山常绿阔叶林群落功能多样性的影响

物种共存机制一直以来是群落生态学的研究热点。为了探讨异质生境条件下鼎湖山常绿阔叶林群落功能多样性变化,找到其变化的主要环境驱动因子,该研究利用位于鼎湖山20 hm~2监测样地第2次群落调查数据并选择代表不同生境(海拔和地形)的27个样方(20 m×20 m),于2013年夏季在样地内所选样方中测定所有胸径≥1 cm树种的叶片功能性状。所测性状包括形态学性状(比叶面积、叶片干物质含量、叶面积以及叶片长宽比)和化学计量学性状(叶片碳、氮、磷的含量),结合地形和土壤数据并通过分析功能多样性随环境梯度的变化,探讨了环境过滤和竞争在鼎湖山群落物种共存中的相对重要性。结果表明:功能分歧度和群落权重平均值与环境因素关系密切,尤其是海拔、凹凸度和土壤养分。环境条件较好区域(微尺度高海拔、高凹凸度和土壤养分含量)的植物采取统一的养分有效保存(低SLA,高LDMC)的适应策略(功能分歧度低),环境过滤所起作用更强;植物在相反的环境条件下,采取快速生长策略(高SLA,低LDMC),能够更好地适应环境的变化,且性状变化是多样的(功能分歧度高),在该条件下竞争作用更为显著。叶面积和叶片氮含量的分歧度在环境条件较好的区域增大,这与其他功能性状不一致,说明不同生态位轴(环境因素)影响不同性状的分歧度变化,并且在局域尺度上植物为了更好地适应环境变化采取了多样的适应策略。     

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.     

三江平原农田地表和地下土壤螨类丰富度与环境因子的空间关联性

地表和地下土壤动物群落空间格局及其与环境因子的空间作用关系,是揭示地表-地下生态系统格局与过程及生物多样性维持机制的重要基础。于2011年在三江平原农田生态系统,在50m×50m的空间尺度内,基于地统计空间分析方法,揭示地表和地下土壤螨群落及不同螨类物种丰富度的空间格局,并分析这种空间格局与土壤含水量、土壤p H值及大豆株高空间格局的空间关联性。半方差函数和普通克里格插值表明,8月份地表、地下和10月份地下螨群落及这些群落内大部分螨类物种在特定空间尺度内形成集群,表现为空间异质性特征,且这种空间分异多由结构性因素或结构性因素和随机性因素共同调控。交叉方差函数表明,土壤螨群落和不同螨类物种的空间格局与环境因子的空间格局在多种尺度上表现出复杂的空间关联性(正的或负的)。但简单Mantel检验仅发现8月份地表中气门亚目未定种1(Mesostigmata unidentified sp.1)和大豆株高存在明显的正的空间关联性。研究结果表明地下螨群落和生长季节的地表螨群落具有明显的空间异质性结构,地表和地下螨群落及大多数螨类物种丰富度与土壤含水量、土壤p H值及大豆株高的空间关联性并不显著。促进地表-地下生态系统土壤动物群落空间格局研究,为地表-地下格局与过程研究奠定基础。     

桂林岩溶石山青冈群落植物功能性状的种间和种内变异研究

研究植物功能性状的种间和种内变异,有助于揭示植物对环境的适应策略及其群落构建机制研究。以桂林岩溶石山青冈群落主要木本植物为研究对象,选取叶干物质含量（LDMC）、叶厚度（LT）、比叶面积（SLA）和木材密度（WD）4个功能性状,采用混合线性模型结合方差分解方法对其在种内和种间尺度的变异程度和相对贡献进行分析,同时运用零模型方法探讨驱动岩溶石山青冈群落构建的潜在机制。结果表明：（1） LDMC,LT,SLA和WD这4个性状在种内和种间水平上均存在不同程度的变异幅度,种间变异大于种内变异,但种内变异亦不容忽略。（2）零模型检验表明,将种内变异纳入考虑的环境过滤检测得到改善,基于个体性状值的尺度考虑群落构建机制研究是必要的。（3）对于桂林岩溶石山青冈群落,环境过滤作用确实是群落构建的重要驱动机制,与相似性限制作用和随机作用共同塑造了以青冈为优势物种的群落构建。     

松嫩草地66种草本植物叶片性状特征

植物叶片功能性状及其相互关系越来越受到关注.以松嫩草地66种草本植物为研究对象,测量叶片干物质含量、比叶面积、叶片厚度、叶片氮含量、叶片磷含量、叶绿素含量和类胡萝卜素含量,检验性状间的相互关系,比较不同功能群(多年生根茎禾草,多年生丛生禾草,多年生杂类草,1年生或2年生草本)间性状的差异性.结果表明,叶片厚度变异系数最大,比叶面积、叶片氮含量、叶片磷含量、叶绿素含量和类胡萝卜素含量之间存在显著的正相关关系;叶片于物质含量与叶片磷含量没有显著的相关关系,与其它叶片性状呈显著的负相关关系;叶片厚度只与叶片干物质含量和比叶面积呈显著的负相关关系,与其它叶片性状不相关.叶片干物质含量、比叶面积、叶片厚度、叶片氮、磷含量在4个功能群间差异显著,叶绿素含量和类胡萝卜素含量在各个功能群间差异不显著;多年生根茎禾草和多年生丛生禾草叶片的7个性状差异不显著;多年生根茎禾草和多年生丛生禾草的叶片干物质含量高于多年生杂类草和1年生或2年生草本,其它性状小于这两个功能群.     

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.     

柯-青冈常绿阔叶林优势树种叶片性状变异及适应策略

厘清叶片功能性状的变异及性状间的权衡关系，对揭示植物对环境变化的响应及适应策略具有重要意义。以中亚热带柯(Lithocarpus glaber)-青冈(Cyclobalanopsis glauca)常绿阔叶林为对象，测量了1 hm²固定监测样地内6个优势树种的叶面积(LA)、比叶面积(SLA)、干物质含量(LDMC)、叶片厚度(LT)、叶片碳(LC)、叶片氮(LN)、叶片磷(LP)含量和叶片碳氮比(LC ∶ LN)8个功能性状，采用多重比较、主成分分析(PCA)法分析了群落叶片功能性状的生活型、种内和种间变异及性状间关系。结果表明：(1)群落内叶片功能性状种内、种间差异显著，变异中等(CV： 0.02-0.59)，其结构性状的可塑性较化学性状保守，变异格局符合"性状空间变异分割假说"；针叶树种的LA、SLA显著低于阔叶树种，常绿树种的LC和LDMC最小，而落叶树种的SLA、LN和LP最大以及LT和LC ∶ LN最小。(2)群落叶片功能性状变异主要来源于生活型和种间变异，种内变异亦有显著贡献；生活型对多数性状的贡献率较大，其对LDMC、LC ∶ LN的贡献分别高达93.11%和91.76%；种间变异(LDMC除外)对结构性状的贡献率均高于化学性状；种内变异对LP的贡献率(23.66%)较种间变异高。(3)叶片性状之间多表现出显著相关关系，针叶树和阔叶树在PCA排序图中聚集于不同区域，叶经济型谱在柯-青冈群落中具有适用性。不同树种通过叶片结构、化学性状之间的权衡策略来适应环境变化，从而实现群落物种共存。结果可为理解森林群落物种的环境适应策略、预测群落动态变化和制定植被恢复措施提供科学依据。     

Divergent response and adaptation of specific leaf area to environmental change at different spatio-temporal scales jointly improve plant survival

Specific leaf area (SLA) is one of the most important plant functional traits. It integrates multiple functions and reflects strategies of plants to obtain resources. How plants employ different strategies (e.g., through SLA) to respond to dynamic environmental conditions remains poorly understood. This study aimed to explore the spatial variation in SLA and its divergent adaptation through the lens of biogeographic patterns, evolutionary history, and short-term responses. SLA data for 5424 plant species from 76 natural communities in China were systematically measured and integrated with meta-analysis of field experiments (i.e., global warming, drought, and nitrogen addition). The mean value of SLA across all species was 21.8 m² kg⁻¹, ranging from 0.9 to 110.2 m² kg⁻¹. SLA differed among different ecosystems, temperature zones, vegetation types, and functional groups. Phylogeny had a weak effect on SLA, but plant species evolved toward higher SLA. Furthermore, SLA responded nonlinearly to environmental change. Unexpectedly, radiation was one of the main factors determining the spatial variation in SLA on a large scale. Conversely, short-term manipulative experiments showed that SLA increased with increased resource availability and tended to stabilize with treatment duration. However, different species exhibited varying response patterns. Overall, variation in long-term adaptation of SLA to environmental gradients and its short-term response to resource pulses jointly improve plant adaptability to a changing environment. Overall SLA-environment relationships should be emphasized as a multidimensional strategy for elucidating environmental change in future research.     

Congruency analysis of species ranking based on leaf traits: which traits are the more reliable?

Nine leaf traits (area, fresh weight, dry weight, volume, density, thickness, specific leaf area (SLA), dry matter content (LDMC), leaf nitrogen content (LNC)) from ten plant species at eight sites in southern mediterranean France were investigated in order to assess their variability along a climatic gradient and their ranking congruency power. After examination of trait correlation patterns, we reduced the nine initial leaf traits to four traits, representative of three correlation groups: allometric traits (dry weight), functional traits (SLA and dry matter percentage) and Leaf Thickness. We analysed the variability of these four leaf traits at species and site level. We observed that between species variation (between 64.5 for SLA and 91% for LDMC) is higher than within species variation. Allowing a good congruency of species ranking assessed by spearman rank correlation () and a good reallocation of individuals to species by discriminant analysis. A site level variability (between 0.7% for Dry weight and 6.9% for SLA) was identified and environmental parameters (altitude, temperature, precipitation, nitrogen, pH) were considered as probable control factors. We found significant correlation between SLA, LDMC and the average minimum temperature (respectively r=0.87 and r=-0,9) and no correlation for the other traits or environmental parameters. Furthermore, we conclude that two leaf traits appear to be central in describing species: specific leaf area (SLA), percentage of dry matter (LDMC. While, SLA and LDMC are strongly correlated, LDMC appears to be less variable than SLA. According to our results the Dry Matter Content (or its reversal Leaf Water Content) appears the best leaf trait to be quantified for plant functional screening. Leaf thickness appeared to be rather uncorrelated with other leaf traits and show no environmental contingency; its variability could not have been explained in this study. Further studies should focus on this trait. This revised version was published online in June 2006 with corrections to the Cover Date.     

Using phylogeny and functional traits for assessing community assembly along environmental gradients: A deterministic process driven by elevation

Community assembly processes is the primary focus of community ecology. Using phylogenetic‐based and functional trait‐based methods jointly to explore these processes along environmental gradients are useful ways to explain the change of assembly mechanisms under changing world. Our study combined these methods to test assembly processes in wide range gradients of elevation and other habitat environmental factors. We collected our data at 40 plots in Taibai Mountain, China, with more than 2,300 m altitude difference in study area and then measured traits and environmental factors. Variance partitioning was used to distinguish the main environment factors leading to phylogeny and traits change among 40 plots. Principal component analysis (PCA) was applied to colligate other environment factors. Community assembly patterns along environmental gradients based on phylogenetic and functional methods were studied for exploring assembly mechanisms. Phylogenetic signal was calculated for each community along environmental gradients in order to detect the variation of trait performance on phylogeny. Elevation showed a better explanatory power than other environment factors for phylogenetic and most traits’ variance. Phylogenetic and several functional structure clustered at high elevation while some conserved traits overdispersed. Convergent tendency which might be caused by filtering or competition along elevation was detected based on functional traits. Leaf dry matter content (LDMC) and leaf nitrogen content along PCA 1 axis showed conflicting patterns comparing to patterns showed on elevation. LDMC exhibited the strongest phylogenetic signal. Only the phylogenetic signal of maximum plant height showed explicable change along environmental gradients. Synthesis. Elevation is the best environment factors for predicting phylogeny and traits change. Plant's phylogenetic and some functional structures show environmental filtering in alpine region while it shows different assembly processes in middle‐ and low‐altitude region by different trait/phylogeny. The results highlight deterministic processes dominate community assembly in large‐scale environmental gradients. Performance of phylogeny and traits along gradients may be independent with each other. The novel method for calculating functional structure which we used in this study and the focus of phylogenetic signal change along gradients may provide more useful ways to detect community assembly mechanisms.     

Relationships between leaf morphological traits, nutrient concentrations and isotopic signatures for Mediterranean woody plant species and communities

Background and aims

Soil factors are driving forces that influence spatial distribution and functional traits of plant species. We test whether two anchor morphological traits—leaf mass per area (LMA) and leaf dry matter content (LDMC)—are significantly related to a broad range of leaf nutrient concentrations in Mediterranean woody plant species. We also explore the main environmental filters (light availability, soil moisture and soil nutrients) that determine the patterns of these functional traits in a forest stand.

Methods

Four morphological and 19 chemical leaf traits (macronutrients and trace elements and δ¹³C and δ¹⁵N signatures) were analysed in 17 woody plant species. Community-weighted leaf traits were calculated for 57 plots within the forest. Links between LMA, LDMC and other leaf traits were analysed at the species and the community level using standardised major axis (SMA) regressions

Results

LMA and LDMC were significantly related to many leaf nutrient concentrations, but only when using abundance-weighted values at community level. Among-traits links were much weaker for the cross-species analysis. Nitrogen isotopic signatures were useful to understand different resource-use strategies. Community-weighted LMA and LDMC were negatively related to light availability, contrary to what was expected.

Conclusion

Community leaf traits have parallel shifts along the environmental factors that determine the community assembly, even though they are weakly related across individual taxa. Light availability is the main environmental factor determining this convergence of the community leaf traits.     

Partitioning the impact of abiotic factors and spatial patterns on species richness and community structure of ground ant assemblages in four Bornean rainforests

Contrasting theories have been proposed to explain the structure of ecological communities. Here, we studied the impact of environmental factors and spatial patterns on ground‐foraging ant communities in four different forest types of Gunung Mulu National Park in Sarawak, Borneo, Malaysia. Forest types differed in their environmental parameters and were inhabited by distinct ant communities, with various indicator species characteristic for single forest types. Three environmental parameters, soil volume, number of trees and amount of leaf litter, had the most influence on ant communities. Spatial patterns were correlated with environmental parameters and also influenced ant communities. Environmental parameters influenced community composition only moderately (r²=0.14), but had a high impact on species richness (r²=0.44). Spatial patterns explained only a small fraction of the total variance in species patterns, while much of the residual space in the ordination space of ant community patterns remained unexplained. We conclude that environmental parameters shape the number of niches within a tropical soil habitat, but identities of species that occupy those niches are accounted for by other factors like competition, traits and neutral processes that may further reduce unexplained variance in species ordination.