Revisiting the Holy Grail: using plant functional traits to understand ecological processes

One of ecology's grand challenges is developing general rules to explain and predict highly complex systems. Understanding and predicting ecological processes from species' traits has been considered a ‘Holy Grail’ in ecology. Plant functional traits are increasingly being used to develop mechanistic models that can predict how ecological communities will respond to abiotic and biotic perturbations and how species will affect ecosystem function and services in a rapidly changing world; however, significant challenges remain. In this review, we highlight recent work and outstanding questions in three areas: (i) selecting relevant traits; (ii) describing intraspecific trait variation and incorporating this variation into models; and (iii) scaling trait data to community‐ and ecosystem‐level processes. Over the past decade, there have been significant advances in the characterization of plant strategies based on traits and trait relationships, and the integration of traits into multivariate indices and models of community and ecosystem function. However, the utility of trait‐based approaches in ecology will benefit from efforts that demonstrate how these traits and indices influence organismal, community, and ecosystem processes across vegetation types, which may be achieved through meta‐analysis and enhancement of trait databases. Additionally, intraspecific trait variation and species interactions need to be incorporated into predictive models using tools such as Bayesian hierarchical modelling. Finally, existing models linking traits to community and ecosystem processes need to be empirically tested for their applicability to be realized.     

Land-use intensification reduces functional redundancy and response diversity in plant communities

Ecosystem resilience depends on functional redundancy (the number of species contributing similarly to an ecosystem function) and response diversity (how functionally similar species respond differently to disturbance). Here, we explore how land-use change impacts these attributes in plant communities, using data from 18 land-use intensity gradients that represent five biomes and > 2800 species. We identify functional groups using multivariate analysis of plant traits which influence ecosystem processes. Functional redundancy is calculated as the species richness within each group, and response diversity as the multivariate within-group dispersion in response trait space, using traits that influence responses to disturbances. Meta-analysis across all datasets showed that land-use intensification significantly reduced both functional redundancy and response diversity, although specific relationships varied considerably among the different land-use gradients. These results indicate that intensified management of ecosystems for resource extraction can increase their vulnerability to future disturbances.
Ecology Letters (2010) 13: 76–86     

Scaling environmental change through the community-level: a trait-based response-and-effect framework for plants

Predicting ecosystem responses to global change is a major challenge in ecology. A critical step in that challenge is to understand how changing environmental conditions influence processes across levels of ecological organization. While direct scaling from individual to ecosystem dynamics can lead to robust and mechanistic predictions, new approaches are needed to appropriately translate questions through the community level. Species invasion, loss, and turnover all necessitate this scaling through community processes, but predicting how such changes may influence ecosystem function is notoriously difficult. We suggest that community‐level dynamics can be incorporated into scaling predictions using a trait‐based response–effect framework that differentiates the community response to environmental change (predicted by response traits) and the effect of that change on ecosystem processes (predicted by effect traits). We develop a response‐and‐effect functional framework, concentrating on how the relationships among species' response, effect, and abundance can lead to general predictions concerning the magnitude and direction of the influence of environmental change on function. We then detail several key research directions needed to better scale the effects of environmental change through the community level. These include (1) effect and response trait characterization, (2) linkages between response‐and‐effect traits, (3) the importance of species interactions on trait expression, and (4) incorporation of feedbacks across multiple temporal scales. Increasing rates of extinction and invasion that are modifying communities worldwide make such a research agenda imperative.     

What does species richness tell us about functional trait diversity? Predictions and evidence for responses of species and functional trait diversity to land‐use change

In the conservation literature on land‐use change, it is often assumed that land‐use intensification drives species loss, driving a loss of functional trait diversity and ecosystem function. Modern research, however, does not support this cascade of loss for all natural systems. In this paper we explore the errors in this assumption and present a conceptual model taking a more mechanistic approach to the species–functional trait association in a context of land‐use change. We provide empirical support for our model's predictions demonstrating that the association of species and functional trait diversity follows various trajectories in response to land‐use change. The central premise of our model is that land‐use change impacts upon processes of community assembly, not species per se. From the model, it is clear that community context (i.e. type of disturbance, species pool size) will affect the response trajectory of the relationship between species and functional trait diversity in communities undergoing land‐use change. The maintenance of ecosystem function and of species diversity in the face of increasing land‐use change are complementary goals. The use of a more ecologically realistic model of responses of species and functional traits will improve our ability to make wise management decisions to achieve both aims in specific at‐risk systems.     

Species loss and gain in communities under future climate change: consequences for functional diversity

It is anticipated that anthropogenic climate change will lead to substantial reassembly within communities in coming decades as individual species shift their ranges to track optimal conditions for growth and survival. As species are lost and gained in communities, what are the consequences for functional trait diversity? Functional traits are the characteristics of species that affect individual performance and provide the vital link between biodiversity at the species level and ecosystem function. We investigated how projected changes in species richness in plant communities under climate change scenarios for the decade 2050 will affect the distribution and diversity of five functional traits. We aggregated range change projections made in Maxent for the decade 2050 across all species in the regional pool of littoral rainforest vines in eastern Australia (n = 163 species). The effect of richness changes on trait diversity was assessed in nine rainforest reserves along the east coast of Australia. Although richness was predicted to significantly decline across all communities, functional diversity remained stable, indicating a decoupling in response to climate change at these two different levels of biological organization. A high degree of redundancy in trait composition in communities may buffer against the loss of function in these plant communities. Scaling‐up our understanding of the impact of climate change from the species level to communities is a critical step towards developing conservation strategies aimed at preserving ecosystem function.     

山西五鹿山森林群落木本植物功能多样性

通过选取群落中木本植物种子的扩散方式、传粉方式、植株高度和盖度等13个功能性状,计算出群落的6个功能多样性指数:功能性状距离、功能性状平均距离、功能体积、功能均匀度、功能分散指数和Rao二次熵指数,结合群落物种丰富度指数、Shannon-Wiener指数和物种均匀度指数对山西五鹿山森林群落木本植物功能多样性进行研究。结果表明:(1)功能性状距离、功能性状平均距离、功能体积与物种丰富度、Shannon-Wiener指数显著正相关;功能均匀度与Shannon-Wiener指数、物种均匀度指数显著正相关;功能分散指数、Rao二次熵指数与物种均匀度指数、Shannon-Wiener指数显著正相关;(2)功能多样性的差异很大程度上是由于物种差异所引起的;(3)6个功能多样性指数可分为三类:功能性状距离、功能性状平均距离、功能体积为功能丰富度指数;功能均匀度为功能均匀度指数;功能分散指数和Rao二次熵指数为功能离散度指数。该分类结果符合指数的计算方法和生态学意义,以及相互独立的标准。     

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.     

Trait structure and redundancy determine sensitivity to disturbance in marine fish communities

Trait diversity is believed to influence ecosystem dynamics through links between organismal traits and ecosystem processes. Theory predicts that key traits and high trait redundancy—large species richness and abundance supporting the same traits—can buffer communities against environmental disturbances. While experiments and data from simple ecological systems lend support, large‐scale evidence from diverse, natural systems under major disturbance is lacking. Here, using long‐term data from both temperate (English Channel) and tropical (Seychelles Islands) fishes, we show that sensitivity to disturbance depends on communities’ initial trait structure and initial trait redundancy. In both ecosystems, we found that increasing dominance by climatically vulnerable traits (e.g., small, fast‐growing pelagics/corallivores) rendered fish communities more sensitive to environmental change, while communities with higher trait redundancy were more resistant. To our knowledge, this is the first study demonstrating the influence of trait structure and redundancy on community sensitivity over large temporal and spatial scales in natural systems. Our results exemplify a consistent link between biological structure and community sensitivity that may be transferable across ecosystems and taxa and could help anticipate future disturbance impacts on biodiversity and ecosystem functioning.     

Do temperate tree species diversity and identity influence soil microbial community function and composition?

Studies of biodiversity–ecosystem function in treed ecosystems have generally focused on aboveground functions. This study investigates intertrophic links between tree diversity and soil microbial community function and composition. We examined how microbial communities in surface mineral soil responded to experimental gradients of tree species richness (SR ), functional diversity (FD ), community‐weighted mean trait value (CWM ), and tree identity. The site was a 4‐year‐old common garden experiment near Montreal, Canada, consisting of deciduous and evergreen tree species mixtures. Microbial community composition, community‐level physiological profiles, and respiration were evaluated using phospholipid fatty acid (PLFA ) analysis and the MicroResp^? system, respectively. The relationship between tree species richness and glucose‐induced respiration (GIR ), basal respiration (BR ), metabolic quotient (qCO ₂) followed a positive but saturating shape. Microbial communities associated with species mixtures were more active (basal respiration [BR ]), with higher biomass (glucose‐induced respiration [GIR ]), and used a greater number of carbon sources than monocultures. Communities associated with deciduous tree species used a greater number of carbon sources than those associated with evergreen species, suggesting a greater soil carbon storage capacity. There were no differences in microbial composition (PLFA ) between monocultures and SR mixtures. The FD and the CWM of several functional traits affected both BR and GIR . In general, the CWM of traits had stronger effects than did FD , suggesting that certain traits of dominant species have more effect on ecosystem processes than does FD . Both the functions of GIR and BR were positively related to aboveground tree community productivity. Both tree diversity (SR ) and identity (species and functional identity—leaf habit) affected soil microbial community respiration, biomass, and composition. For the first time, we identified functional traits related to life‐history strategy, as well as root traits that influence another trophic level, soil microbial community function, via effects on BR and GIR .     

Intraspecific plant–soil feedback and intraspecific overyielding in Arabidopsis thaliana

Understanding the mechanisms of community coexistence and ecosystem functioning may help to counteract the current biodiversity loss and its potentially harmful consequences. In recent years, plant–soil feedback that can, for example, be caused by below‐ground microorganisms has been suggested to play a role in maintaining plant coexistence and to be a potential driver of the positive relationship between plant diversity and ecosystem functioning. Most of the studies addressing these topics have focused on the species level. However, in addition to interspecific interactions, intraspecific interactions might be important for the structure of natural communities. Here, we examine intraspecific coexistence and intraspecific diversity effects using 10 natural accessions of the model species Arabidopsis thaliana (L.) Heynh. We assessed morphological intraspecific diversity by measuring several above‐ and below‐ground traits. We performed a plant–soil feedback experiment that was based on these trait differences between the accessions in order to determine whether A. thaliana experiences feedback at intraspecific level as a result of trait differences. We also experimentally tested the diversity–productivity relationship at intraspecific level. We found strong differences in above‐ and below‐ground traits between the A. thaliana accessions. Overall, plant–soil feedback occurred at intraspecific level. However, accessions differed in the direction and strength of this feedback: Some accessions grew better on their own soils, some on soils from other accessions. Furthermore, we found positive diversity effects within A. thaliana: Accession mixtures produced a higher total above‐ground biomass than accession monocultures. Differences between accessions in their feedback response could not be explained by morphological traits. Therefore, we suggest that they might have been caused by accession‐specific accumulated soil communities, by root exudates, or by accession‐specific resource use based on genetic differences that are not expressed in morphological traits. Synthesis. Our results provide some of the first evidence for intraspecific plant–soil feedback and intraspecific overyielding. These findings may have wider implications for the maintenance of variation within species and the importance of this variation for ecosystem functioning. Our results highlight the need for an increased focus on intraspecific processes in plant diversity research to fully understand the mechanisms of coexistence and ecosystem functioning.     

Does functional homogenization accompany taxonomic homogenization of British birds and how do biotic factors and climate affect these processes?

Environmental change has reshuffled communities often causing taxonomic homogenization rather than differentiation. Some studies suggest that this increasing similarity of species composition between communities is accompanied by an increase in similarity of trait composition—functional homogenization—although different methodologies have failed to come to any consistent conclusions. Functional homogenization could have a large effect on ecosystem functioning and stability. Here, we use the general definition of homogenization as “reduced spatial turnover over time” to compare changes in Simpson's beta diversity (taxonomic turnover) with changes in Rao's quadratic entropy beta diversity (functional turnover) in British breeding birds at three spatial scales. Using biotic and climatic variables, we identify which factors may predispose a site to homogenization. The change in turnover measures between two time periods, 20 years apart, was calculated. A null model approach was taken to identify occurrences of functional homogenization and differentiation independent of changes in taxonomic turnover. We used conditional autoregressive models fitted using integrated nested Laplace approximations to determine how environmental drivers and factors relating to species distributions affect changes in spatial turnover of species and functional diversity. The measurement of functional homogenization affects the chance of rejection of the null models, with many sites showing taxonomic homogenization unaccompanied by functional homogenization, although occurrence varies with spatial scale. At the smallest scale, while temperature‐related variables drive changes in taxonomic turnover, changes in functional turnover are associated with variation in growing degree days; however, changes in functional turnover become more difficult to predict at larger spatial scales. Our results highlight the multifactorial processes underlying taxonomic and functional homogenization and that redundancy in species traits may allow ecosystem functioning to be maintained in some areas despite changes in species composition.     

基于功能性状的生态系统服务研究框架

功能性状通过影响生态系统的属性和过程及其维持来影响生态系统服务。功能多样性-生态系统功能关系的研究有助于深入探讨生态系统服务形成机制, 也为生态系统服务研究提供了一个切入点。该文对目前的功能性状和生态系统服务研究框架进行了介绍, 回顾了功能多样性-生态系统功能关系的研究现状, 总结了目前功能性状在生态系统服务研究中的应用, 提出了基于功能性状的生态系统服务研究框架。在这个研究框架中, 首先选取对生态系统功能影响显著的非生物因子和功能多样性指数, 然后量化非生物因子和功能多样性与生态系统功能, 以及生态系统功能-生态系统服务之间的关系, 进而构建功能多样性与生态系统服务的数量关系。与此同时, 利用群落构建理论和物种共存机制分析功能多样性-生态系统功能变化的机制联系, 以研究生态系统服务形成和变化机制, 为生态系统服务管理决策提供科学依据。     

Functional traits explain waterbirds' host status,subtype richness,and community-level infection risk for avian influenza

Species functional traits can influence pathogen transmission processes, and consequently affect species' host status, pathogen diversity, and community-level infection risk. We here investigated, for 143 European waterbird species, effects of functional traits on host status and pathogen diversity (subtype richness) for avian influenza virus at species level. We then explored the association between functional diversity and HPAI H5Nx occurrence at the community level for 2016/17 and 2021/22 epidemics in Europe. We found that both host status and subtype richness were shaped by several traits, such as diet guild and dispersal ability, and that the community-weighted means of these traits were also correlated with community-level risk of H5Nx occurrence. Moreover, functional divergence was negatively associated with H5Nx occurrence, indicating that functional diversity can reduce infection risk. Our findings highlight the value of integrating trait-based ecology into the framework of diversity–disease relationship, and provide new insights for HPAI prediction and prevention.     

群落/生态系统功能多样性研究方法及展望

功能多样性是指影响群落/生态系统功能的物种性状值和范围,是解释和预测生态系统结构和功能的有效手段之一,可将植物个体尺度与群落尺度和生态系统尺度的相关生态学问题联系起来。虽已发展出多种功能多样性定量化研究方法,但不同方法结果差异较大,难以进行多研究间的比较研究。比较探讨各功能多样性研究方法的优缺点有利于拓展功能多样性内涵,也有助于功能生态学的应用与发展。回顾了当前10种功能多样性的定量化研究方法,并指出选取合适功能多样性方法的关键在于,应考虑选取群落/生态系统中的哪些物种、哪些功能性状、选取的功能性状数目、以及如何对功能性状权重等。对比发现,功能分散性指数和Rao二次熵系数的研究方法在众多方法中优势明显,具有较高的应用潜力;标准化功能多样性的研究方法在未来仍需进一步完善。利用功能多样性指数预测群落/生态系统过程和功能当前仍多侧重于理论研究,野外实证研究较为缺乏,是功能生态学未来研究的重点和难点之一。     

Functional diversity is positively associated with biomass for lake diatoms

1. Recent work has emphasised the benefit of using functional measures when relating biodiversity to ecosystem functioning. In this study, we investigated the extent to which functional and taxonomic diversity might be related to summed biovolume in community assemblages of 212 species of diatoms collected from 65 temperate lakes in western and central Quebec, Canada. 2. We quantified functional diversity as both the total path‐length of a functional dendrogram (FD) and the variance in species traits (TV) for a given community. Selected traits included both size and responses to a set of environmental variables known to be influential for diatom communities. 3. Species richness, as well as both FD and TV, was positively associated with total diatom biovolume at the level of the entire diatom community, suggesting that diversity in response types (particularly to total phosphorus and pH) is important for diatom community production. 4. Although functional measures of diversity did not provide enhanced explanatory power over species richness, we argue that an exploration of functional traits potentially allows greater insight into the mechanisms underlying biodiversity–ecosystem functioning relations, indicating which traits might be most influential in driving community biomass production.     

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.     

基于功能性状的中亚热带岩溶常绿落叶阔叶混交林物种多样性维持研究

理解植物功能性状和功能实体在森林群落的分布,有助于探讨物种丧失对森林生态系统功能、冗余和恢复力的影响。为了解脆弱的岩溶石山森林在应对生物多样性丧失的生态系统反馈,对桂林岩溶石山两块1hm²的常绿落叶阔叶混交林的木本植物数据进行了分析。包括基于功能性状计算功能多样性、构建功能实体计算功能冗余以及采用Pearson相关分析和Mantel检验评估物种多样性指标在生态系统的维持机制。结果显示:(1)青冈+大叶榉树群落的功能多样性指标均低于鱼骨木+青冈+圆叶乌桕群落,且两个群落间功能均匀度不相关(P ＞ 0.05),功能丰富度、功能离散度和Rao''s二次熵呈现极显著相关性(P ＜ 0.001),功能分散度呈现显著相关性(P ＜ 0.05)。(2)两个群落的物种丰富度与功能冗余指标表现出相似的线性关系,即物种丰富度与功能实体等级、功能冗余、功能超冗余呈正相关,与功能脆弱性呈负相关关系。(3)不同植物功能性状间、不同功能多样性指标间和不同功能冗余指标间的相关性较强,功能多样性指标和功能冗余指标间无显著相关性,但功能性状与功能多样性指标、功能性状与功能冗余指标均存在不同程度的相关性。而在功能实体与物种多样性指标的相关性方面,呈现出同物种丰富度与物种多样性指标相似的显著度。另外,物种多度与物种丰富度、功能分散度、功能离散度、Rao''s二次熵及功能脆弱性均显著相关。总之,在岩溶石山常绿落叶阔叶混交林中,高功能多样性的群落存在高功能冗余的现象,但功能多样性和功能冗余是相互独立的因素;物种丰富度高的群落所提供的保险效应无法抵消其生态系统的脆弱性。因此,不能仅通过保护物种丰富度来维持生态系统的特有功能,还应充分考虑多度对生态系统功能的贡献,以更有效地实现对岩溶石山森林生态系统的保护。     

Drivers of productivity and its temporal stability in a tropical tree diversity experiment

There is increasing evidence that mixed‐species forests can provide multiple ecosystem services at a higher level than their monospecific counterparts. However, most studies concerning tree diversity and ecosystem functioning relationships use data from forest inventories (under noncontrolled conditions) or from very young plantation experiments. Here, we investigated temporal dynamics of diversity–productivity relationships and diversity–stability relationships in the oldest tropical tree diversity experiment. Sardinilla was established in Panama in 2001, with 22 plots that form a gradient in native tree species richness of one‐, two‐, three‐ and five‐species communities. Using annual data describing tree diameters and heights, we calculated basal area increment as the proxy of tree productivity. We combined tree neighbourhood‐ and community‐level analyses and tested the effects of both species diversity and structural diversity on productivity and its temporal stability. General patterns were consistent across both scales indicating that tree–tree interactions in neighbourhoods drive observed diversity effects. From 2006 to 2016, mean overyielding (higher productivity in mixtures than in monocultures) was 25%–30% in two‐ and three‐species mixtures and 50% in five‐species stands. Tree neighbourhood diversity enhanced community productivity but the effect of species diversity was stronger and increased over time, whereas the effect of structural diversity declined. Temporal stability of community productivity increased with species diversity via two principle mechanisms: asynchronous responses of species to environmental variability and overyielding. Overyielding in mixtures was highest during a strong El Niño‐related drought. Overall, positive diversity–productivity and diversity–stability relationships predominated, with the highest productivity and stability at the highest levels of diversity. These results provide new insights into mixing effects in diverse, tropical plantations and highlight the importance of analyses of temporal dynamics for our understanding of the complex relationships between diversity, productivity and stability. Under climate change, mixed‐species forests may provide both high levels and high stability of production.     

Connections between species diversity and genetic diversity

Species diversity and genetic diversity remain the nearly exclusive domains of community ecology and population genetics, respectively, despite repeated recognition in the literature over the past 30 years of close parallels between these two levels of diversity. Species diversity within communities and genetic diversity within populations are hypothesized to co‐vary in space or time because of locality characteristics that influence the two levels of diversity via parallel processes, or because of direct effects of one level of diversity on the other via several different mechanisms. Here, we draw on a wide range of studies in ecology and evolution to examine the theoretical underpinnings of these hypotheses, review relevant empirical literature, and outline an agenda for future research. The plausibility of species diversity–genetic diversity relationships is supported by a variety of theoretical and empirical studies, and several recent studies provide direct, though preliminary support. Focusing on potential connections between species diversity and genetic diversity complements other approaches to synthesis at the ecology–evolution interface, and should contribute to conceptual unification of biodiversity research at the levels of genes and species.