A conceptual framework for the spatial analysis of functional trait diversity

Functional trait diversity is a popular tool in modern ecology, mainly used to infer assembly processes and ecosystem functioning. Patterns of functional trait diversity are shaped by ecological processes such as environmental filtering, species interactions and dispersal that are inherently spatial, and different processes may operate at different spatial scales. Adding a spatial dimension to the analysis of functional trait diversity may thus increase our ability to infer community assembly processes and to predict change in assembly processes following disturbance or land‐use change. Richness, evenness and divergence of functional traits are commonly used indices of functional trait diversity that are known to respond differently to large‐scale filters related to environmental heterogeneity and dispersal and fine‐scale filters related to species interactions (competition). Recent developments in spatial statistics make it possible to separately quantify large‐scale patterns (variation in local means) and fine‐scale patterns (variation around local means) by decomposing overall spatial autocorrelation quantified by Moran's coefficient into its positive and negative components using Moran eigenvector maps (MEM). We thus propose to identify the spatial signature of multiple ecological processes that are potentially acting at different spatial scales by contrasting positive and negative components of spatial autocorrelation for each of the three indices of functional trait diversity. We illustrate this approach with a case study from riparian plant communities, where we test the effects of disturbance on spatial patterns of functional trait diversity. The fine‐scale pattern of all three indices was increased in the disturbed versus control habitat, suggesting an increase in local scale competition and an overall increase in unexplained variance in the post‐disturbance versus control community. Further research using simulation modeling should focus on establishing the proposed link between community assembly rules and spatial patterns of functional trait diversity to maximize our ability to infer multiple processes from spatial community structure.     

The effects of human disturbance on the species composition,species diversity and functional diversity of a Miombo woodland in northern Malawi

Anthropogenic disturbances have serious impacts on ecosystems across the world. Understanding the effects of disturbance on woodlands, especially in regions where local people depend on these natural resources, is essential for sustainable natural resource management and biodiversity conservation. In this study, we evaluated the effects of anthropogenic disturbance, specifically selective logging of Brachystegia floribunda, on woodlands by comparing species composition, species diversity and functional diversity of woody plants between disturbed and undisturbed woodlands. We combined species data and functional trait data for leaves, fruits and other traits related to resource and disturbance responses to calculate functional indices (functional richness, evenness and divergence) and community‐weighted means of each trait. Shifts in taxonomic species composition were analysed using nonmetric multi‐dimensional scaling. Species composition differed significantly between disturbed and undisturbed woodlands. Tree density was greater in disturbed woodlands, whereas evenness, functional evenness and functional divergence were greater in undisturbed woodlands. In terms of forest cover, selective logging of B. floribunda appeared to have little impact on Miombo woodlands, but some shifts in functional traits, such as the shift from a deciduous to evergreen phenology, may increase the vulnerability of these ecosystems to environmental change, especially drought.     

Climate and small scale factors determine functional diversity shifts of biological soil crusts in Iberian drylands

Understanding functional diversity is critical to manage and preserve biodiversity and ecosystem functioning in the face of global change. However, the efforts to characterize this functional component have been mostly directed to vascular vegetation. We sampled lichen-dominated biological soil crusts (BSCs) in semiarid grasslands along an environmental gradient in the Iberian Peninsula. We characterized five effect functional traits for 31 lichens species, and evaluated the influence of large scale (i.e. precipitation) and small scale factors (i.e. substrate type, shrub presence, Stipa tenacissima presence) on dominant trait values; i.e. community weighted means, and functional divergence; i.e. Rao quadratic entropy in 580 sampling quadrats. Across the gradient, we found multiple trait shifts and a general increase of functional divergence with increasing precipitation. We also observed that substrate type and small scale biotic factors determined shifts in all traits studied, while these factors affected less to functional divergence. Comparing functional diversity with taxonomic diversity, we found contrasting responses to both large and small scale factors. These findings suggest that BSC community trait composition is influenced by multi-scale abiotic and biotic factors with environmental filtering dominating at large spatial scales and limiting similarity at specific small scales. Also, our results emphasize the potential differences between taxonomic and functional diversity in response to environmental factors. We concluded that functional diversity of BSCs not only provides novel and critical knowledge of BSC community structure, but also it should be considered as a critical tool in biodiversity conservation strategies, ecosystem services assessment and ecological modelling.     

CWM and Rao’s quadratic diversity: a unified framework for functional ecology

Assessing the effects of environmental constraints on community structure often relies on methods that consider changes in species functional traits in response to environmental processes. Various indices have been proposed to measure relevant aspects of community trait composition from different viewpoints and perspectives. Among these, the ‘community-weighted mean trait value’ (CWM) and the Rao coefficient have been widely used in ecological research for summarizing different facets of functional composition and diversity. Analyzing changes in functional diversity of bee communities along a post-fire successional gradient in southern Switzerland we show that these two measures may be used to describe two complementary aspects of community structure, such as the mean and the dispersion of functional traits within a given species assemblage. While CWM can be adequately used to summarize shifts in mean trait values within communities due to environmental selection for certain functional traits, the Rao coefficient can be effectively applied to analyze patterns of trait convergence or divergence compared to a random expectation.     

刈割与施肥干扰下高寒草甸植物功能性状的构建模式

针对目前关于植物群落功能性状构建模式及其驱动因素存在的争议, 研究了33个物种10个功能性状的构建模式。研究结果显示: (1)在刈割-施肥复合梯度上, 这些功能性状主要表现为随机构建模式, 发生随机性和确定性构建的样方比例分别为82.7%和17.3%; (2)在10个功能性状中, 生长型、生活周期、单株地上干质量、叶面积和叶干质量5个功能性状为随机构建模式, 不受试验处理和群落特征(地上净初级生产力、刈割生物量损失、群落高度)变化的影响。植物倾斜度、繁殖方式、固氮性、株高和比叶面积5个功能性状的构建模式与试验处理或群落特征变化有关, 表现出趋同或趋异构建响应。其中, 植物倾斜度和比叶面积的构建模式仅受群落特征影响, 而固氮性、繁殖方式和株高3个性状的构建模式同时还受刈割或施肥处理的影响, 其构建模式因功能性状而异; (3)群落的生物量损失和地上净初级生产力是解释植物功能性状构建模式变化较理想的群落特征; (4)刈割和施肥处理对株高的构建模式具有相反效应, 而刈割和施肥的交互作用对其无显著影响。上述结果说明该研究群落植物功能性状的构建存在不同模式, 以随机构建模式为主, 确定性构建模式居次要地位。确定性构建模式与试验处理和群落特征变化有关, 而且是性状依赖的。相反选择力对趋同和趋异构建模式的平衡效应能引起功能性状发生随机构建。     

生态学的多样性指数:功能与系统发育

生物多样性是生态学的核心问题。传统的多样性指数仅包含物种数和相对多度的信息,这类基于分类学的多样性指数并不能很好地帮助理解群落构建和生态系统功能。不同物种对群落构建和生态系统功能所起到的作用类型和贡献也不完全相同,且物种在生态过程中的作用和贡献往往与性状密切相关,因此功能多样性已经成为反映物种群落构建、干扰以及环境因素对群落影响的重要指标。同时,由于亲缘关系相近的物种往往具有相似的性状,系统发育多样性也可以作为功能多样性的一个替代。功能多样性和系统发育多样性各自具有优缺点,但二者均比分类多样性更能揭示群落和生态系统的构建、维持与功能。     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.     

More species,but all do the same: contrasting effects of flood disturbance on ground beetle functional and species diversity

The role of habitat disturbance on biodiversity is central as it promotes changes in ecological systems. That said, still little is known about the functional consequences of such changes. Functional diversity can be used to revealing more mechanistically the disturbance effects on communities by considering the richness and the distribution of traits among the species. Here we analyzed the response of functional and species diversity of ground beetles to flood disturbance to better understand the functioning of alluvial invertebrate communities. Ground beetles were sampled in periodically flooded grasslands along the Elbe River in Germany. We used generalized linear mixed effects models to unveil the relationships between flood disturbance, species and functional diversity, respectively. We measured different components of functional diversity (functional richness, evenness, dispersion, and divergence) and analyzed species diversity by means of rarefied species richness, abundances, evenness and Simpson's diversity. We found contrasting relationships in that most species diversity measures peaked at highest disturbance levels, while most functional diversity measures decreased with increasing disturbance intensities. Inversed relationships between species and functional diversity are rarely observed, as most studies report on positive correlations. We explain increasing species diversity with a higher amount of resources available in highly disturbed sites. Decreasing functional diversity is best explained through the convergence of species traits by flood disturbance and uneven resource exploitation in highly disturbed plots (low functional evenness), suggesting strong impacts from functionally different generalist species in floodchannels. We show that the amount of resources available, and how these resources are exploited, play major roles in the functioning of floodplain ground beetle communities.     

Partitioning of functional diversity reveals the scale and extent of trait convergence and divergence

Questions: Trait differentiation among species occurs at different spatial scales within a region. How does the partitioning of functional diversity help to identify different community assembly mechanisms? Location: Northeastern Spain. Methods: Functional diversity can be partitioned into within‐community (α) and among‐communities (β) components, in analogy to Whittaker's classical α and β species diversity concept. In light of ecological null models, we test and discuss two algorithms as a framework to measure α and β functional diversity (the Rao quadratic entropy index and the variance of trait values). Species and trait (specific leaf area) data from pastures under different climatic conditions in NE Spain are used as a case study. Results: The proposed indices show different mathematical properties but similarly account for the spatial components of functional diversity. For all vegetation types along the climatic gradient, the observed α functional diversity was lower than expected at random, an observation consistent with the hypothesis of trait convergence resulting from habitat filtering. On the other hand, our data exhibited a remarkably higher functional diversity within communities compared to among communities (α?β). In contrast to the high species turnover, there was a limited functional diversity turnover among communities, and a large part of the trait divergence occurred among coexisting species. Conclusions: Partitioning functional diversity within and among communities revealed that both trait convergence and divergence occur in the formation of assemblages from the local species pool. A considerable trait convergence exists at the regional scale in spite of changes in species composition, suggesting the existence of ecological redundancy among communities.     

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.     

Weak functional response to agricultural landscape homogenisation among plants,butterflies and birds

Measures of functional diversity are expected to predict community responses to land use and environmental change because, in contrast to taxonomic diversity, it is based on species traits rather than their identity. Here, we investigated the impact of landscape homogenisation on plants, butterflies and birds in terms of the proportion of arable field cover in southern Finland at local (0.25 km²) and regional (> 10 000 km²) scales using four functional diversity indices: functional richness, functional evenness, functional divergence and functional dispersion. No uniform response in functional diversity across taxa or scales was found. However, in all cases where we found a relationship between increasing arable field cover and any index of functional diversity, this relationship was negative. Butterfly functional richness decreased with increasing arable field cover, as did butterfly and bird functional evenness. For butterfly functional evenness, this was only evident in the most homogeneous regions. Butterfly and bird functional dispersion decreased in homogeneous regions regardless of the proportion of arable field cover locally. No effect of landscape heterogeneity on plant functional diversity was found at any spatial scale, but plant species richness decreased locally with increasing arable field cover. Overall, species richness responded more consistently to landscape homogenisation than did the functional diversity indices, with both positive and negative effects across species groups. Functional diversity indices are in theory valuable instruments for assessing effects of land use scenarios on ecosystem functioning. However, the applicability of empirical data requires deeper understanding of which traits reliably capture species’ vulnerability to environmental factors and of the ecological interpretation of the functional diversity indices. Our study provides novel insights into how the functional diversity of communities changes in response to agriculturally derived landscape homogenisation; however, the low explanatory power of the functional diversity indices hampers the ability to reliably anticipate impacts on ecosystem functioning.     

坡向因子对黄土高原草地群落功能多样性的影响

研究群落水平上的植物功能性状特征及功能多样性随坡向的变化规律,对认识不同坡向上的植物群落形成过程具有重要意义。以黄土高原不同坡向上的自然草地群落为研究对象,比较研究了植物功能性状(株高和比叶面积)和功能多样性测度指标(功能丰富度、功能均匀度和功能离散度)随坡向的变化规律。研究结果显示:(1)阴坡的株高和比叶面积显著高于其他坡向;(2)一元性状的功能丰富度在不同坡向间均无显著差异;阴坡和半阴坡的多元性状功能丰富度显著高于阳坡;(3)阴坡的株高功能均匀度显著高于半阳坡,而比叶面积功能均匀度在各坡向的差异并不显著;多元性状功能均匀度在不同坡向差异显著,阴坡最高,半阳坡最低;(4)阴坡和半阴坡的株高功能离散度显著高于阳坡和半阳坡,而半阴坡的比叶面积功能离散度显著高于阳坡;半阴坡的多元性状功能离散度Rao指数显著高于阳坡。研究结果暗示了,在阴坡和半阴坡上,植物对群落内的生态位空间和资源利用更充分,种间竞争强度较低,不同物种之间生态位高度分化;而在阳坡和半阳坡上,由于水分等条件的限制植物可占据的生态位空间有限,导致其对占据的生态位空间使用不足,物种间资源竞争较强烈。在群落水平上,研究地区的植物功能性状及功能多样性随坡向的规律性变化,反映了黄土高原植被群落构建过程中坡向因子对功能性状的筛选效应。该研究结果对该区的植被恢复重建的物种选择及植被布局规划具有重要实践意义。     

Fine‐scale coexistence patterns along a productivity gradient in wet meadows: shifts from trait convergence to divergence

Semi‐natural meadows host a great number of species coexisting at fine spatial scales. Different assembly mechanisms, related to differences in functional traits between species, can influence such coexistence. Coexisting species could be either functionally dissimilar to occupy different niches (‘divergence’) or functionally similar due to exclusion of species with traits less adapted to the prevailing abiotic and biotic conditions (‘convergence’). Various theories differently predict how trait convergence and divergence should differ due to disturbance, along productivity gradients, and across different functional traits. We tested such theories in 21 wet meadows of different productivity in central Europe. In each meadow, four 1 × 1 m plots were established in which disturbance by mowing was combined with fertilization. Species presence was recorded in 100 quadrats 10 × 10 cm in size within the plots over five years. Convergence and divergence were assessed at very fine spatial scales (10 × 10 cm) to focus on the processes driven by the interactions for similar resources. Convergence emerged as the dominant pattern for all traits and across all years. It was particularly strong in the least productive conditions while divergence emerged in some of the most productive meadows. Mowing increased convergence in meadows with low productivity, but increased divergence in productive meadows. Fertilization generally increased convergence, with this increase being more pronounced in mown plots. Convergence in unproductive conditions could be caused by either higher fitness of stress‐tolerant species (more abundant in the species pool of these sites) or by functionally similar species sharing similar patches within fine‐scale heterogeneous plots. This outcome also suggests abiotic filters can have an important role at fine scales, where plant‐ecological theory usually predicts the prevalence of biotic processes.     

Taxonomical vs. functional responses of bee communities to fire in two contrasting climatic regions

1. Valuable insights into mechanisms of community responses to environmental change can be gained by analysing in tandem the variation in functional and taxonomic composition along environmental gradients. 2. We assess the changes in species and functional trait composition (i.e. dominant traits and functional diversity) of diverse bee communities in contrasting fire-driven systems in two climatic regions: Mediterranean (scrub habitats in Israel) and temperate (chestnut forests in southern Switzerland). 3. In both climatic regions, there were shifts in species diversity and composition related to post-fire age. In the temperate region, functional composition responded markedly to fire; however, in the Mediterranean, the taxonomic response to fire was not matched by functional replacement. 4. These results suggest that greater functional stability to fire in the Mediterranean is achieved by replacement of functionally similar species (i.e. functional redundancy) which dominate under different environmental conditions in the heterogeneous landscapes of the region. In contrast, the greater functional response in the temperate region was attributed to a more rapid post-fire vegetation recovery and shorter time-window when favourable habitat was available relative to the Mediterranean. 5. Bee traits can be used to predict the functional responses of bee communities to environmental changes in habitats of conservation importance in different regions with distinct disturbance regimes. However, predictions cannot be generalized from one climatic region to another where distinct habitat configurations occur.     

Unifying functional trait approaches to understand the assemblage of ecological communities: synthesizing taxonomic divides

Functional traits have long been considered the ‘holy grail’ in community ecology due to their potential to link phenotypic variation with ecological processes. Advancements across taxonomic disciplines continue to support functional ecology's objective to approach generality in community assembly. However, a divergence of definitions, aims and methods across taxa has created discord, limiting the field's predictive capacity. Here, we provide a guide to support functional ecological comparisons across taxa. We describe advances in cross‐taxa functional research, identify gaps in approaches, synthesize definitions and unify methodological considerations. When deciding which traits to compare, particularly response traits, we advocate selecting functionally analogous traits that relate to community assembly processes. Finally, we describe at what scale and for which questions functional comparisons across taxa are useful and when other approaches may be more constructive. Our approach promotes standardized methods for integrative research across taxa to identify broad trends in community assembly.     

cati: an R package using functional traits to detect and quantify multi‐level community assembly processes

Community ecologists are active in describing species by their functional traits, quantifying the functional structure of plant and animal assemblages and inferring community assembly processes with null‐model analyses of trait distribution and functional diversity indices. Intraspecific variation in traits and effects of spatial scale are potentially important in these analyses. Here, we introduce the R package cati (Community Assembly by Traits: Individuals and beyond) available on CRAN, for the analysis of community assembly with functional traits. cati builds on a recent approach to community assembly that explicitly incorporates individual differences in community assembly analyses and decomposes phenotypic variations across scales and organizational levels, based on three phenotypic variance ratios, termed the T‐statistics. More generally, the cati package 1) calculates a variety of single‐trait and multi‐trait indices from interspecific and intraspecific trait measures; 2) it partitions functional trait variation among spatial and taxonomic levels; 3) it implements a palette of flexible null models for detecting non‐random patterns of functional traits. These patterns can be used to draw inferences about hypotheses of community assembly such as environmental filtering and species interactions. The basic input for cati is a data frame in which columns are traits, rows are species or individuals, and entries are the measured trait values. The cati package can also incorporate a square distance matrix into analyses, which could include phylogenetic or genetic distances among individuals or species. Users select from a variety of functional trait metrics and analyze these relative to a null model that specifies trait distributions in a regional source pool.     

Low functional diversity and no redundancy in British avian assemblages

1. Spatial and temporal patterns in functional diversity can reveal the patterns and processes behind community assembly and whether ecological redundancy exists. Here, we analyse functional diversity in British avian assemblages over a period of about 20 years. 2. Functional diversity is generally lower than expected by chance, indicating that assemblages contain species with relatively similar functional traits. One potential explanation is filtering for traits suitable to particular habitats, though other explanations exist. 3. There was no evidence of ecological redundancy over the 20 years. In fact, changes in functional diversity were almost exactly proportional to changes in species richness. 4. The absence of functional redundancy results from little redundancy intrinsic to the species' functional relationships and also because compositional change was nonrandom. Observed extinction and colonization events caused greater changes in functional diversity than if these events were random. 5. Our findings suggest that community assembly is influenced by the traits of species and that observed changes in functional diversity provide no reason to believe that the functioning of natural systems is buffered against change by ecological redundancy.     

Plant diversity induces shifts in the functional structure and diversity across trophic levels

Changes to primary producer diversity can cascade up to consumers and affect ecosystem processes. Although the effect of producer diversity on higher trophic groups have been studied, these studies often quantify taxonomy‐based measures of biodiversity, like species richness, which do not necessarily reflect the functioning of these communities. In this study, we assess how plant species richness affects the functional composition and diversity of higher trophic levels and discuss how this might affect ecosystem processes, such as herbivory, predation and decomposition. Based on six different consumer traits, we examined the functional composition of arthropod communities sampled in experimental plots that differed in plant species richness. The two components we focused on were functional variation in the consumer community structure (functional structure) and functional diversity, expressed as functional richness, evenness and divergence. We found a consistent positive effect of plant species richness on the functional richness of herbivores, carnivores, and omnivores, but not decomposers, and contrasting patterns for functional evenness and divergence. Increasing plant species richness shifted the omnivore community to more predatory and less mobile species, and the herbivore community to more specialized and smaller species. This was accompanied by a shift towards more species occurring in the vegetation than in the ground layer. Our study shows that plant species richness strongly affects the functional structure and diversity of aboveground arthropod communities. The observed shifts in body size (herbivores), specialization (herbivores), and feeding mode (omnivores) together with changes in the functional diversity may underlie previously observed increases in herbivory and predation in plant communities of higher diversity.