Plant diversity effects on arthropods and arthropod-dependent ecosystem functions in a biodiversity experiment

Biodiversity-ecosystem function experiments test how species diversity influences fundamental ecosystem processes. Historically, arthropod driven functions, such as herbivory and pest-control, have been thought to be influenced by direct and indirect associations among species. Although a number of studies have evaluated how plant diversity affects arthropod communities and arthropod-mediated ecosystem processes, it remains unclear whether diversity effects on arthropods are sufficiently consistent over time such that observed responses can be adequately predicted by classical hypotheses based on associational effects. By combining existing results from a long-term grassland biodiversity experiment (Jena Experiment) with new analyses, we evaluate the consistency of consumer responses within and across taxonomic, trophic, and trait-based (i.e. vertical stratification) groupings, and we consider which changes in arthropod community composition are associated with changes in consumer-mediated ecosystem functions.Overall, higher plant species richness supported more diverse and complex arthropod communities and this pattern was consistent across multiple years. Vegetation-associated arthropods responded more strongly to changes in plant species richness than ground-dwelling arthropods. Additionally, increases in plant species richness were associated with shifts in the species-abundance distributions for many, but not all taxa. For example, highly specialized consumers showed a decrease in dominance and an increase in the number of rare species with increasing plant species richness. Most ecosystem processes investigated responded to increases in plant species richness in the same way as the trophic group mediating the process, e.g. both herbivory and herbivore diversity increase with increasing plant species richness. In the Jena Experiment and other studies, inconsistencies between predictions based on classic hypotheses of associational effects and observed relationships between plant species richness and arthropod diversity likely reflect the influence of multi-trophic community dynamics and species functional trait distributions. Future research should focus on testing a broader array of mechanisms to unravel the biological processes underlying the biodiversity-ecosystem functioning relationships.     

Moving beyond the guild concept: developing a practical functional trait framework for terrestrial beetles

1. New logical and analytical frameworks for studying functional traits have led to major advances in plant and freshwater ecology at local and global scales. The ecological and taxonomic diversity of terrestrial adult beetles (Coleoptera) means that functional trait approaches should have considerable power to illuminate the function not only of these animals but also of the ecosystems in which they occur. 2. Even though the functional trait concept is not new in ecology, it is still plagued with inconsistencies in methodology and terminology. Plant‐based studies have shown that an integrated and relatively consistent functional trait approach facilitates comparisons between studies, and allows the full utility and predictive capacity of trait‐based approaches to be realised. 3. This review outlines a logical framework for adult beetle functional trait studies using uniform terminology and methodology similar to those used by plant ecologists. Beetle life‐history and ecomorphological trait studies are synthesised and it is shown that a combination of both is analogous to the functional trait approach. A general functional trait list for beetles and potential functional links is outlined, as are potential analysis approaches. A consistent functional trait approach, coupled with advances in molecular techniques, has the capability to provide deeper insights into beetle community assembly and how beetles impact ecosystems and will enable worldwide comparisons and predictions to be made.     

Land‐use effects on the functional distinctness of arthropod communities

Land‐use change is a major driver of the global loss of biodiversity, but it is unclear to what extent this also results in a loss of ecological traits. Therefore, a better understanding of how land‐use change affects ecological traits is crucial for efforts to sustain functional diversity. To this end we tested whether higher species richness or taxonomic distinctness generally leads to increased functional distinctness and whether intensive land use leads to functionally more narrow arthropod communities. We compiled species composition and trait data for 350 species of terrestrial arthropods (Araneae, Carabidae and Heteroptera) in different land‐use types (forests, grasslands and arable fields) of low and high land‐use intensity. We calculated the average functional and taxonomic distinctness and the rarified trait richness for each community. These measures reflect the range of traits, taxonomic relatedness and number of traits that are observed in local communities. Average functional distinctness only increased significantly with species richness in Carabidae communities. Functional distinctness increased significantly with taxonomic distinctness in communities of all analyzed taxa suggesting a high functional redundancy of taxonomically closely related species. Araneae and Heteroptera communities had the expected lower functional distinctness at sites with higher land‐use intensity. More frequently disturbed land‐use types such as managed grasslands or arable fields were characterized by species with smaller body sizes and higher dispersal abilities and communities with lower functional distinctness or trait richness. Simple recommendations about the conservation of functional distinctness of arthropod communities in the face of future land‐use intensification and species loss are not possible. Our study shows that these relationships depend on the studied taxa and land‐use type. However, for some arthropod groups functional distinctness is threatened by intensification and conversion from less to more frequently disturbed land‐uses.     

Evolutionary community ecology of plant-associated arthropods in terrestrial ecosystems

In the 21st century, researchers have attempted a synthesis between community ecology and evolutionary biology. This emerging research area, which aims to synthesize community ecology and evolutionary biology, is evolutionary community ecology. Evolutionary community ecology addresses how intraspecific trait variation in community members is essential for predicting community properties and, how community properties are a key component of the selective forces that determine genetic and phenotypic variation in a community member. In this paper, I review recent findings in evolutionary community ecology in plant-associated arthropods in terrestrial ecosystems. I discuss roles of both genetic variation and phenotypic plasticity as a source of trait variation in plants in shaping plant-associated arthropod communities. Also, I discuss effects of genetic variation in herbivores on plant-associated arthropod communities. Furthermore, I highlight community context evolution in which multiple species interactions and community composition affect trait evolution of a community member. Finally, I argue that future studies should investigate a feedback loop between community and evolutionary dynamics beyond unidirectional studies on effects of evolution on a community or vice versa. This approach will provide major insights into mechanistic principles for making predictions of community ecology.     

Functional richness outperforms taxonomic richness in predicting ecosystem functioning in natural phytoplankton communities

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Functional traits explain phytoplankton community structure and seasonal dynamics in a marine ecosystem

A fundamental yet elusive goal of ecology is to predict the structure of communities from the environmental conditions they experience. Trait‐based approaches to terrestrial plant communities have shown that functional traits can help reveal the mechanisms underlying community assembly, but such approaches have not been tested on the microbes that dominate ecosystem processes in the ocean. Here, we test whether functional traits can explain community responses to seasonal environmental fluctuation, using a time series of the phytoplankton of the English Channel. We show that interspecific variation in response to major limiting resources, light and nitrate, can be well‐predicted by lab‐measured traits characterising light utilisation, nitrate utilisation and maximum growth rate. As these relationships were predicted a priori, using independently measured traits, our results show that functional traits provide a strong mechanistic foundation for understanding the structure and dynamics of ecological communities.     

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.     

Does genomic variation in a foundation species predict arthropod community structure in a riparian forest?

Understanding how genetic variation within a foundation species determines the structure of associated communities and ecosystem processes has been an emerging frontier in ecology. Previous studies in common gardens identified close links between intraspecific variation and multispecies community structure, and these findings are now being evaluated directly in the complex natural ecosystem. In this study, we examined to what extent genomic variation in a foundation tree species explains the structure of associated arthropod communities in the field, comparing with spatial, temporal and environmental factors. In a continuous mixed forest, arthropods were surveyed on 85 mature alders (Alnus hirsuta) in 2 years. Moreover, we estimated Nei's genetic distance among the alders based on 1,077 single nucleotide polymorphisms obtained from restricted‐site‐associated DNA sequencing of the alders’ genome. In both years, we detected significant correlations between genetic distance and dissimilarity of arthropod communities. A generalized dissimilarity modelling indicated that the genetic distance of alder populations was the most important predictor to explain the variance of arthropod communities. Among arthropod functional groups, carnivores were consistently correlated with genetic distance of the foundation species in both years. Furthermore, the extent of year‐to‐year changes in arthropod communities was more similar between more genetically closed alder populations. This study demonstrates that the genetic similarity rule would be primarily prominent in community assembly of plant‐associated arthropods under temporally and spatially variable environments in the field.     

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.     

Biodiversity and ecosystem functioning in naturally assembled communities

Approximately 25 years ago, ecologists became increasingly interested in the question of whether ongoing biodiversity loss matters for the functioning of ecosystems. As such, a new ecological subfield on Biodiversity and Ecosystem Functioning (BEF) was born. This subfield was initially dominated by theoretical studies and by experiments in which biodiversity was manipulated, and responses of ecosystem functions such as biomass production, decomposition rates, carbon sequestration, trophic interactions and pollination were assessed. More recently, an increasing number of studies have investigated BEF relationships in non‐manipulated ecosystems, but reviews synthesizing our knowledge on the importance of real‐world biodiversity are still largely missing. I performed a systematic review in order to assess how biodiversity drives ecosystem functioning in both terrestrial and aquatic, naturally assembled communities, and on how important biodiversity is compared to other factors, including other aspects of community composition and abiotic conditions. The outcomes of 258 published studies, which reported 726 BEF relationships, revealed that in many cases, biodiversity promotes average biomass production and its temporal stability, and pollination success. For decomposition rates and ecosystem multifunctionality, positive effects of biodiversity outnumbered negative effects, but neutral relationships were even more common. Similarly, negative effects of prey biodiversity on pathogen and herbivore damage outnumbered positive effects, but were less common than neutral relationships. Finally, there was no evidence that biodiversity is related to soil carbon storage. Most BEF studies focused on the effects of taxonomic diversity, however, metrics of functional diversity were generally stronger predictors of ecosystem functioning. Furthermore, in most studies, abiotic factors and functional composition (e.g. the presence of a certain functional group) were stronger drivers of ecosystem functioning than biodiversity per se. While experiments suggest that positive biodiversity effects become stronger at larger spatial scales, in naturally assembled communities this idea is too poorly studied to draw general conclusions. In summary, a high biodiversity in naturally assembled communities positively drives various ecosystem functions. At the same time, the strength and direction of these effects vary highly among studies, and factors other than biodiversity can be even more important in driving ecosystem functioning. Thus, to promote those ecosystem functions that underpin human well‐being, conservation should not only promote biodiversity per se, but also the abiotic conditions favouring species with suitable trait combinations.     

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.     

Do landscape health indices reflect arthropod biodiversity status in the eucalypt woodlands of eastern Australia?

Ecosystem or landscape health indices are important tools for land managers. While strong predictable relationships between these indices and biotic diversity are often generalized, they are seldom validated. Here we use data from a semi‐arid eastern Australian woodland to examine the relationships between arthropod community structure and two sets of landscape health indicators: landscape function analysis (LFA), and a terrestrial index of ecological integrity based on common vegetation metrics (structure, composition and function; SCF). Hierarchical partitioning revealed that the ability of LFA or SCF to account for variation in arthropod richness was low, with the variable of importance taxon‐dependent. Similarly, multivariate analyses indicated relatively weak and inconsistent relationships between LFA and SCF indices and arthropod assemblage structure. Results obtained for additional habitat attributes commonly used in terrestrial vegetation monitoring were similar. Our study indicates that strong predictable relationships are rarely apparent, particularly for arthropods. This indicates that these indices have limited use as surrogates of arthropod biodiversity. These results are contrary to the past literature, highlighting the need for additional research and the development of a conceptual and empirical framework linking health indices and arthropod biodiversity. This is necessary to further the theoretical and practical application of these measurements in environmental management.     

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.     

Human‐aided admixture may fuel ecosystem transformation during biological invasions: theoretical and experimental evidence

Biological invasions can transform our understanding of how the interplay of historical isolation and contemporary (human‐aided) dispersal affects the structure of intraspecific diversity in functional traits, and in turn, how changes in functional traits affect other scales of biological organization such as communities and ecosystems. Because biological invasions frequently involve the admixture of previously isolated lineages as a result of human‐aided dispersal, studies of invasive populations can reveal how admixture results in novel genotypes and shifts in functional trait variation within populations. Further, because invasive species can be ecosystem engineers within invaded ecosystems, admixture‐induced shifts in the functional traits of invaders can affect the composition of native biodiversity and alter the flow of resources through the system. Thus, invasions represent promising yet under‐investigated examples of how the effects of short‐term evolutionary changes can cascade across biological scales of diversity. Here, we propose a conceptual framework that admixture between divergent source populations during biological invasions can reorganize the genetic variation underlying key functional traits, leading to shifts in the mean and variance of functional traits within invasive populations. Changes in the mean or variance of key traits can initiate new ecological feedback mechanisms that result in a critical transition from a native ecosystem to a novel invasive ecosystem. We illustrate the application of this framework with reference to a well‐studied plant model system in invasion biology and show how a combination of quantitative genetic experiments, functional trait studies, whole ecosystem field studies and modeling can be used to explore the dynamics predicted to trigger these critical transitions.     

Fungal functional ecology: bringing a trait‐based approach to plant‐associated fungi

Fungi play many essential roles in ecosystems. They facilitate plant access to nutrients and water, serve as decay agents that cycle carbon and nutrients through the soil, water and atmosphere, and are major regulators of macro‐organismal populations. Although technological advances are improving the detection and identification of fungi, there still exist key gaps in our ecological knowledge of this kingdom, especially related to function . Trait‐based approaches have been instrumental in strengthening our understanding of plant functional ecology and, as such, provide excellent models for deepening our understanding of fungal functional ecology in ways that complement insights gained from traditional and ‐omics‐based techniques. In this review, we synthesize current knowledge of fungal functional ecology, taxonomy and systematics and introduce a novel database of fungal functional traits (Fun^Fun). Fun^Fun is built to interface with other databases to explore and predict how fungal functional diversity varies by taxonomy, guild, and other evolutionary or ecological grouping variables. To highlight how a quantitative trait‐based approach can provide new insights, we describe multiple targeted examples and end by suggesting next steps in the rapidly growing field of fungal functional ecology.     

Plant Diversity Impacts Decomposition and Herbivory via Changes in Aboveground Arthropods

Loss of plant diversity influences essential ecosystem processes as aboveground productivity, and can have cascading effects on the arthropod communities in adjacent trophic levels. However, few studies have examined how those changes in arthropod communities can have additional impacts on ecosystem processes caused by them (e.g. pollination, bioturbation, predation, decomposition, herbivory). Therefore, including arthropod effects in predictions of the impact of plant diversity loss on such ecosystem processes is an important but little studied piece of information. In a grassland biodiversity experiment, we addressed this gap by assessing aboveground decomposer and herbivore communities and linking their abundance and diversity to rates of decomposition and herbivory. Path analyses showed that increasing plant diversity led to higher abundance and diversity of decomposing arthropods through higher plant biomass. Higher species richness of decomposers, in turn, enhanced decomposition. Similarly, species-rich plant communities hosted a higher abundance and diversity of herbivores through elevated plant biomass and C:N ratio, leading to higher herbivory rates. Integrating trophic interactions into the study of biodiversity effects is required to understand the multiple pathways by which biodiversity affects ecosystem functioning.     

Applying trait‐based models to achieve functional targets for theory‐driven ecological restoration

Manipulating community assemblages to achieve functional targets is a key component of restoring degraded ecosystems. The response‐and‐effect trait framework provides a conceptual foundation for translating restoration goals into functional trait targets, but a quantitative framework has been lacking for translating trait targets into assemblages of species that practitioners can actually manipulate. This study describes new trait‐based models that can be used to generate ranges of species abundances to test theories about which traits, which trait values and which species assemblages are most effective for achieving functional outcomes. These models are generalisable, flexible tools that can be widely applied across many terrestrial ecosystems. Examples illustrate how the framework generates assemblages of indigenous species to (1) achieve desired community responses by applying the theories of environmental filtering, limiting similarity and competitive hierarchies, or (2) achieve desired effects on ecosystem functions by applying the theories of mass ratios and niche complementarity. Experimental applications of this framework will advance our understanding of how to set functional trait targets to achieve the desired restoration goals. A trait‐based framework provides restoration ecology with a robust scaffold on which to apply fundamental ecological theory to maintain resilient and functioning ecosystems in a rapidly changing world.     

Agroenergy Crops Influence the Diversity, Biomass, and Guild Structure of Terrestrial Arthropod Communities

Expanded production of contemporary bioenergy crops (e.g., corn) is considered a threat to the conservation of biodiversity, yet next-generation perennially based crops (switchgrass, mixed-grass?Cforb prairie) may represent an opportunity for enhancing biodiversity in agricultural landscapes. We employed a multi-scaled approach to investigate the relative importance of feedstock selection, forb content, patch size, and landscape-scale habitat structure and composition as factors shaping the diversity and abundance of terrestrial arthropod communities and the biomass of functional groups of arthropods associated with the provisioning of ecosystem services. Compared to intensively managed annual corn fields, switchgrass and mixed-grass?Cforb prairie plantings were associated with a 230% and 324% increase in arthropod family diversity and a 750% and 2,700% increase in arthropod biomass, respectively. Biomass of arthropod pollinators, herbivores, predators, and parasites were similarly the highest in mixed-grass?Cforb prairie, intermediate in switchgrass plantings, and the lowest in cornfields. Community-wide biomass and that of several functional arthropod groups were positively linked to increasing forest cover and land cover diversity surrounding biomass plantings, while pollinator and detritivore biomass was lower in smaller fields. Results not only suggest that the choice of biomass feedstock will play an important role in shaping within-field arthropod diversity but also indicate an important role for the composition of this surrounding landscape. Collectively, our results suggest that selection of perennially based biomass feedstocks along with careful attention to crop placement have important potential to enhance biodiversity conservation and the provisioning of ecologically and economically important arthropod-mediated ecosystem services in future agricultural landscapes.