Assessing functional redundancy in chronically trawled benthic communities

Fishing disturbance on ecosystems leads to changes in community structure and composition, which may have drastic implications for ecosystem functional performance. Functional redundancy, defined as species sharing similar functional roles, is a community property that plays an important role in preventing functional changes in ecosystems under pressure. In this study, we suggest that functional redundancy may be achieved through trait abundance (i.e. large amounts of a trait, hereafter “common traits”), or through trait richness (i.e. large numbers of distinct taxa exhibiting the same trait, hereafter “widespread traits”). We assessed the variability of both measures obtained from epifaunal and infaunal communities in soft-bottom trawling grounds. Sampling sites were located in four Mediterranean areas that were subjected to different levels of trawling effort. Common and widespread traits measures were based on the analysis of biological traits linked to key soft-bottoms functions such as nutrient cycling, bentho-pelagic coupling and habitat provision. The role of rare species in both measures was also assessed and we observed that, in our study sites, rare species generally exhibited the same traits as the most abundant species. Common and widespread traits measures provided complementary information on benthic functional redundancy. Thus, we suggest that a combination of the two measures should be used to appropriately assess benthic functional redundancy in trawling grounds. As redundancy is a component of ecosystem resilience, functional redundancy evaluation is important to assess the overall integrity of ecosystems.     

A macroecological perspective of trait patterns in stream communities

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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     

Functional diversity: a tool for answering challenging ecological questions

Functional trait diversity provides a powerful means of addressing ecology's persistent questions, through its dual role as an indicator of mechanisms driving differences in species composition between communities and as a predictor of ecosystem‐level processes. Functional traits provide a means of testing mechanisms behind species turnover between communities because environmental heterogeneity, competition and disturbance influence species fitness via their traits. Functional traits also provide a link between species and multiple ecosystem‐level processes, such as primary productivity, nutrient fluxes and resilience, since species influence these processes via their traits. This special issue demonstrates that functional diversity offers a practical means of investigating ecology's persistent questions.     

Impacts of multiple anthropogenic stressors on stream macroinvertebrate community composition and functional diversity

Ensuring the provision of essential ecosystem services in systems affected by multiple stressors is a key challenge for theoretical and applied ecology. Trait‐based approaches have increasingly been used in multiple‐stressor research in freshwaters because they potentially provide a powerful method to explore the mechanisms underlying changes in populations and communities. Individual benthic macroinvertebrate traits associated with mobility, life history, morphology, and feeding habits are often used to determine how environmental drivers structure stream communities. However, to date multiple‐stressor research on stream invertebrates has focused more on taxonomic than on functional metrics. We conducted a fully crossed, 4‐factor experiment in 64 stream mesocosms fed by a pristine montane stream (21 days of colonization, 21 days of manipulations) and investigated the effects of nutrient enrichment, flow velocity reduction and sedimentation on invertebrate community, taxon, functional diversity and trait variables after 2 and 3 weeks of stressor exposure. 89% of the community structure metrics, 59% of the common taxa, 50% of functional diversity metrics, and 79% of functional traits responded to at least one stressor each. Deposited fine sediment and flow velocity reduction had the strongest impacts, affecting invertebrate abundances and diversity, and their effects translated into a reduction of functional redundancy. Stressor effects often varied between sampling occasions, further complicating the prediction of multiple‐stressor effects on communities. Overall, our study suggests that future research combining community, trait, and functional diversity assessments can improve our understanding of multiple‐stressor effects and their interactions in running waters.     

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.     

Fish communities diverge in species but converge in traits over three decades of warming

Describing the spatial and temporal dynamics of communities is essential for understanding the impacts of global environmental change on biodiversity and ecosystem functioning. Trait‐based approaches can provide better insight than species‐based (i.e. taxonomic) approaches into community assembly and ecosystem functioning, but comparing species and trait dynamics may reveal important patterns for understanding community responses to environmental change. Here, we used a 33‐year database of fish monitoring to compare the spatio‐temporal dynamics of taxonomic and trait structure in North Sea fish communities. We found that the majority of variation in both taxonomic and trait structure was explained by a pronounced spatial gradient, with distinct communities in the southern and northern North Sea related to depth, sea surface temperature, salinity and bed shear stress. Both taxonomic and trait structure changed significantly over time; however taxonomically, communities in the south and north diverged towards different species, becoming more dissimilar over time, yet they converged towards the same traits regardless of species differences. In particular, communities shifted towards smaller, faster growing species with higher thermal preferences and pelagic water column position. Although taxonomic structure changed over time, its spatial distribution remained relatively stable, whereas in trait structure, the southern zone of the North Sea shifted northward and expanded, leading to homogenization. Our findings suggest that global environmental change, notably climate warming, will lead to convergence towards traits more adapted for novel environments regardless of species composition.     

Limitations of trait‐based approaches for stressor assessment: The case of freshwater invertebrates and climate drivers

The appeal of trait‐based approaches for assessing environmental vulnerabilities arises from the potential insight they provide into the mechanisms underlying the changes in populations and community structure. Traits can provide ecologically based explanations for observed responses to environmental changes, along with predictive power gained by developing relationships between traits and environmental variables. Despite these potential benefits, questions remain regarding the utility and limitations of these approaches, which we explore focusing on the following questions: (a) How reliable are predictions of biotic responses to changing conditions based on single trait–environment relationships? (b) What factors constrain detection of single trait–environment relationships, and how can they be addressed? (c) Can we use information on meta‐community processes to reveal conditions when assumptions underlying trait‐based studies are not met? We address these questions by reviewing published literature on aquatic invertebrate communities from stream ecosystems. Our findings help to define factors that influence the successful application of trait‐based approaches in addressing the complex, multifaceted effects of changing climate conditions on hydrologic and thermal regimes in stream ecosystems. Key conclusions are that observed relationships between traits and environmental stressors are often inconsistent with predefined hypotheses derived from current trait‐based thinking, particularly related to single trait–environment relationships. Factors that can influence findings of trait‐based assessments include intercorrelations of among traits and among environmental variables, spatial scale, strength of biotic interactions, intensity of habitat disturbance, degree of abiotic stress, and methods of trait characterization. Several recommendations are made for practice and further study to address these concerns, including using phylogenetic relatedness to address intercorrelation. With proper consideration of these issues, trait‐based assessment of organismal vulnerability to environmental changes can become a useful tool to conserve threatened populations into the future.     

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.     

The functional response and resilience in small waterbodies along land‐use and environmental gradients

There is growing recognition of the essential services provided to humanity by functionally intact ecosystems. Freshwater ecosystems are found throughout agricultural and urban landscapes and provide a wide range of ecosystem services, but globally they are also amongst the most vulnerable. In particular, ponds (lentic waters typically less than 2 ha), provide natural flood management, sequester carbon and hold significant cultural value. However, to inform their management it is important to understand (1) how functional diversity varies in response to disturbance and (2) the link between biodiversity conservation and ecosystem function. In this study, a meta‐analysis of seven separate pond studies from across England and Wales was carried out to explore the effect of urban and agricultural land‐use gradients, shading, emergent vegetation, surface area and pH upon groups of functionally similar members of the macroinvertebrate fauna. Functional effect groups were first identified by carrying out a hierarchical cluster analysis using body size, voltinism and feeding habits (18 categories) that are closely related to biogeochemical processes (e.g. nutrient and carbon recycling). Secondly, the influence of the gradients upon effect group membership (functional redundancy—FR) and the breadth of traits available to aid ecosystem recovery (response diversity) was assessed using species counts and functional dispersion (FDis) using 12 response traits. The effect of land‐use gradients was unpredictable, whilst there was a negative response in both FR and FDis to shading and positive responses to increases in emergent vegetation cover and surface area. An inconsistent association between FDis and FR suggested that arguments for taxonomic biodiversity conservation to augment ecosystem functioning are too simplistic. Thus, a deeper understanding of the response of functional diversity to disturbance could have greater impact with decision‐makers who may relate better to the loss of ecosystem function in response to environmental degradation than species loss alone.     

植物性状研究的机遇与挑战:从器官到群落

植物性状(Plant trait)或植物功能性状(Plant functional trait)通常是指植物对外界环境长期适应与进化后所表现出的可量度、且与生产力优化或环境适应等密切相关的属性。近几十年来,植物性状研究在性状-生产力、性状-养分、性状间相互关系、性状-群落结构维持等方面取得了卓越成就。然而,由于大多数性状调查都是以植物群落内优势种或亚优势种为对象,使其在探讨群落尺度的性状-功能关系、性状数据如何用于改进或优化模型、性状数据如何与遥感连接等问题时,存在空间尺度和量纲不匹配的极大挑战。为了破解上述难题,亟需发展新的、基于单位土地面积的群落性状(Community trait)概念体系、数据源和计算方法等,推动植物性状数据与快速发展的宏观生态学新技术(遥感、模型和通量观测等)相结合,既拓展了植物性状研究范畴,又可推动其更好地服务于区域生态环境问题的解决。所定义的群落性状(如叶片氮含量、磷含量、比叶面积、气孔密度、叶绿素含量等),是在充分考虑群落内所有物种的性状实测数据,再结合比叶面积、生物量异速生长方程和群落结构数据等,推导而成的基于单位土地面积的群落性状。受测试方法的影响,传统的直接算术平均法或相对生物量加权平均法所获得的群落水平的植物性状(如叶片氮含量g/kg或%),虽然可以有效地探讨群落结构维持机制,由于无法实现对群落性状在量纲上向单位土地面积转换,使它很难与模型和遥感数据相匹配。基于单位土地面积的群落性状,可在空间尺度匹配(或量纲匹配)的前提下实现个体水平测定的植物性状数据与生态模型和遥感观测相联系,更好地探讨区域尺度下自然生态系统结构和功能的关系及其对全球变化的响应与适应。同时,它也可更好地建立群落水平的性状-功能的定量关系(非物种水平),为更好地探讨自然群落结构维持机制和生产力优化机制提供了新思路。     

Heterogeneous flows foster heterogeneous assemblages: relationships between functional diversity and hydrological heterogeneity in riparian plant communities

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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.     

Hierarchical effects of environmental filters on the functional structure of plant communities: a case study in the French Alps

Understanding the influence of the environment on the functional structure of ecological communities is essential to predict the response of biodiversity to global change drivers. Ecological theory suggests that multiple environmental factors shape local species assemblages by progressively filtering species from the regional species pool to local communities. These successive filters should influence the various components of community functional structure in different ways. In this paper, we tested the relative influence of multiple environmental filters on various metrics of plant functional trait structure (i.e. ‘community weighted mean trait’ and components of functional trait diversity, i.e. functional richness, evenness and divergence) in 82 vegetation plots in the Guisane Valley, French Alps. For the 211 sampled species we measured traits known to capture key aspects of ecological strategies amongst vascular plant species, i.e. leaf traits, plant height and seed mass (LHS). A comprehensive information theory framework, together with null model based resampling techniques, was used to test the various environmental effects. Particular community components of functional structure responded differently to various environmental gradients, especially concerning the spatial scale at which the environmental factors seem to operate. Environmental factors acting at a large spatial scale (e.g. temperature) were found to predominantly shape community weighted mean trait values, while fine‐scale factors (topography and soil characteristics) mostly influenced functional diversity and the distribution of trait values among the dominant species. Our results emphasize the hierarchical nature of ecological forces shaping local species assemblage: large‐scale environmental filters having a primary effect, i.e. selecting the pool of species adapted to a site, and then filters at finer scales determining species abundances and local species coexistence. This suggests that different components of functional community structure will respond differently to environmental change, so that predicting plant community responses will require a hierarchical multi‐facet approach.     

Multi‐scale habitat modification by coexisting ecosystem engineers drives spatial separation of macrobenthic functional groups

By changing habitat conditions, ecosystem engineers increase niche diversity and have profound effects on the distribution and abundances of other organisms. Although many ecosystems contain several engineering species, it is still unclear how the coexistence of multiple engineers affects the physical habitat and the structure of the community on a landscape scale. Here, we investigated through a large‐scale field manipulation how three coexisting engineers on intertidal flats (cockles Cerastoderma edule; lugworms Arenicola marina; blue mussels Mytilus edulis) influence the functional composition of the local macrobenthic community and what the consequences are at the landscape level. By using biological trait analysis (BTA), we show that on the local scale biogenic changes in sediment accumulation and organic matter content translated into specific shifts in the distribution of functional traits within the community. At a landscape scale, the co‐occurrence of multiple ecosystem engineers resulted in the spatial separation of different functional groups, i.e. different functional groups dominated unique complementary habitats. Our results emphasize the role of co‐occurring multiple engineers in shaping natural communities, thus contributing to a better knowledge of community assembly rules. This understanding can profitably be used to improve ecosystem‐based management and conservation actions.     

Fire‐induced taxonomic and functional changes in saproxylic beetle communities in fire sensitive regions

It is often suggested that fire acts as an environmental filter that selects species and functional traits, and reduces trait variability within communities, affecting ecosystem function and underlying services. This may be particularly important in fire‐sensitive ecosystems, such as the central European Alps, where fires are scarce. According to climate and land use change scenarios in Europe, fire risk will increase during the next decades, raising important questions about the maintenance of ecological and functional resilience in these regions. We used two families of saproxylic beetles (i.e. Cerambycidae and Buprestidae) as model group to test the combined effect of fire and altitude on species and trait composition in the central Alps of Switzerland. Trait response was based on weighted means and variation of 15 traits over the communities. Our results showed an overall positive effect of fire on taxonomic and functional diversity, while indicator species and community analyses revealed that the response to fire was also modulated by altitude. The positive effect of fire and the presence of large populations of pyrophilous species suggest co‐evolution with fire and adaptation to disturbance in the Alps. Biodiversity in the central Alps might thus be more resilient to fire than expected. In the light of climatic and land use changes, forest management and species conservation in the central Alps have to consider fire one of the major disruptive factors that have shaped and will shape species composition and ecosystem services.     

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.