The relationship between biodiversity and ecosystem functioning in food webs

Recent theoretical and experimental work provides clear evidence that biodiversity loss can have profound impacts on functioning of natural and managed ecosystems and the ability of ecosystems to deliver ecological services to human societies. Work on simplified ecosystems in which the diversity of a single trophic level is manipulated shows that diversity can enhance ecosystem processes such as primary productivity and nutrient retention. Theory also strongly suggests that biodiversity can act as biological insurance against potential disruptions caused by environmental changes. However, these studies generally concern a single trophic level, primary producers for the most part. Changes in biodiversity also affect ecosystem functioning through trophic interactions. Here we review, through the analysis of a simple ecosystem model, several key aspects inherent in multitrophic systems that may strongly affect the relationship between diversity and ecosystem processes. Our analysis shows that trophic interactions have a strong impact on the relationships between diversity and ecosystem functioning, whether the ecosystem property considered is total biomass or temporal variability of biomass at the various trophic levels. In both cases, food-web structure and trade-offs that affect interaction strength have major effects on these relationships. Multitrophic interactions are expected to make biodiversity–ecosystem functioning relationships more complex and non-linear, in contrast to the monotonic changes predicted for simplified systems with a single trophic level.     

Linking biodiversity indicators,ecosystem functioning,provision of services and human well-being in estuarine systems: Application of a conceptual framework

Assuming that human well-being strongly relies on the services provided by well-functioning ecosystems, changes in the ecological functioning of any system can have direct and indirect effects on human welfare. Intensive land use and tourism have expanded in recent decades along coastal ecosystems, together with increasing demands for water, food and energy; all of these factors intensify the exploitation of natural resources. Many of the interrelations between ecosystem functioning and the provision of ecosystem services (ES) still require quantification in estuarine systems. A conceptual framework to assess such links in a spatially and temporally explicit manner is proposed and applied to the Mondego estuary (Portugal). This framework relies on three consecutive steps and discriminates among biodiversity structural components, ecosystem functioning and stability and the services provided by the ecosystem.Disturbances in abiotic factors were found to have a direct effect on biodiversity, ecosystem functioning and the provision of ES. The observed changes in the species composition of communities had a positive effect on the ecosystem's productivity and stability. Moreover, the observed changes in the estuarine ES provision are likely to arise from changing structural and abiotic factors and in the present case from the loss or decline of locally abundant species. This study also indicates that linear relationships between biodiversity, ecosystem functioning and services provision are unlikely to occur in estuarine systems. Instead, cumulative and complex relations are observed between factors on both temporal and spatial scales. In this context, the results suggest several additional conclusions: (1) biodiversity and ecosystem functioning interaction with human well-being need to be incorporated into decision-making processes aimed at the conservative management of systems; (2) the institutional use of research results must be part of the design and implementation of sustainable management activities; and (3) more integrative tools/studies are required to account for the interactions of estuarine ecosystems with surrounding socio-economic activities. Therefore, when performing integrated assessments of ecosystem dynamics, it becomes essential to consider not only the effects of biodiversity and ecosystem functioning on services provision but also the effects that human well-being and ES provision may have on estuarine biodiversity and ecosystem functioning.The proposed framework implies taking into account both the functional and the commodities points of view upon natural ecosystems and by this representing a line of thought which will deserve further research to explore more in detail the conceptual links between biodiversity–ecosystem functioning–services provided.     

Predicting the responsiveness of soil biodiversity to deforestation: a cross‐biome study

The consequences of deforestation for aboveground biodiversity have been a scientific and political concern for decades. In contrast, despite being a dominant component of biodiversity that is essential to the functioning of ecosystems, the responses of belowground biodiversity to forest removal have received less attention. Single‐site studies suggest that soil microbes can be highly responsive to forest removal, but responses are highly variable, with negligible effects in some regions. Using high throughput sequencing, we characterize the effects of deforestation on microbial communities across multiple biomes and explore what determines the vulnerability of microbial communities to this vegetative change. We reveal consistent directional trends in the microbial community response, yet the magnitude of this vegetation effect varied between sites, and was explained strongly by soil texture. In sandy sites, the difference in vegetation type caused shifts in a suite of edaphic characteristics, driving substantial differences in microbial community composition. In contrast, fine‐textured soil buffered microbes against these effects and there were minimal differences between communities in forest and grassland soil. These microbial community changes were associated with distinct changes in the microbial catabolic profile, placing community changes in an ecosystem functioning context. The universal nature of these patterns allows us to predict where deforestation will have the strongest effects on soil biodiversity, and how these effects could be mitigated.     

Biodiversity–ecosystem function experiments reveal the mechanisms underlying the consequences of biodiversity change in real world ecosystems

In a recent Forum paper, Wardle (Journal of Vegetation Science, 2016) questions the value of biodiversity–ecosystem function (BEF) experiments with respect to their implications for biodiversity changes in real world communities. The main criticism is that the previous focus of BEF experiments on random species assemblages within each level of diversity has ‘limited the understanding of how natural communities respond to biodiversity loss.’ He concludes that a broader spectrum of approaches considering both non‐random gains and losses of diversity is essential to advance this field of research. Wardle's paper is timely because of recent observations of frequent local and regional biodiversity changes across ecosystems. While we appreciate that new and complementary experimental approaches are required for advancing the field, we question criticisms regarding the validity of BEF experiments. Therefore, we respond by briefly reiterating previous arguments emphasizing the reasoning behind random species composition in BEF experiments. We describe how BEF experiments have identified important mechanisms that play a role in real world ecosystems, advancing our understanding of ecosystem responses to species gains and losses. We discuss recent examples where theory derived from BEF experiments enriched our understanding of the consequences of biodiversity changes in real world ecosystems and where comprehensive analyses and integrative modelling approaches confirmed patterns found in BEF experiments. Finally, we provide some promising directions in BEF research.     

Perspectives on the link between ecosystem services and biodiversity: The assessment of the nursery function

The relationship between biodiversity and each ecosystem service or bundle of ecosystem services (e.g. win−win, win−lose or win−neutral) is an active field of research that requires structured and consistent information. The application of that research for conservation and decision-making can be hampered by the ambiguity found in the definition of the nursery function under the ecosystem service perspective. In this paper, we review how the role of nursery habitats is included in the ecosystem services literature, covering conceptual, biophysical and economic reflections. The role of ecosystems as nurseries is mostly analyzed in coastal environments. The main observation is that there is no consensus on the consideration of the nursery function as a service (e.g. which species or habitats) or on how to assess it (e.g. which indicators or valuation methods). After that review, we analyze three different interpretations given to the nursery function, namely the ecological, conservationist and economic point of view; and we distinguish between different types of assessment that may consider the nursery function.We conclude that the nursery function can be considered an ecosystem service on its own right when it is linked to a concrete human benefit and not when it is represented with indicators of general biodiversity or ecosystem condition. Thus, the analysis of the delivery of ecosystem services should be differentiated from the analysis of ecological integrity. Only with this distinction science may be able to quantify the link between biodiversity and ecosystem services and policy may be effective in halting biodiversity loss. Similar considerations could apply for other biodiversity constituents that may be treated as ecosystem services.     

Effects of plant diversity,community composition and environmental parameters on productivity in montane European grasslands

In the past years, a number of studies have used experimental plant communities to test if biodiversity influences ecosystem functioning such as productivity. It has been argued, however, that the results achieved in experimental studies may have little predictive value for species loss in natural ecosystems. Studies in natural ecosystems have been equivocal, mainly because in natural ecosystems differences in diversity are often confounded with differences in land use history or abiotic parameters. In this study, we investigated the effect of plant diversity on ecosystem functioning in semi-natural grasslands. In an area of 10×20 km, we selected 78 sites and tested the effects of various measures of diversity and plant community composition on productivity. We separated the effects of plant diversity on ecosystem functioning from potentially confounding effects of community composition, management or environmental parameters, using multivariate statistical analyses. In the investigated grasslands, simple measures of biodiversity were insignificant predictors of productivity. However, plant community composition explained productivity very well (R²=0.31) and was a better predictor than environmental variables (soil and site characteristics) or management regime. Thus, complex measures such as community composition and structure are important drivers for ecosystem functions in semi-natural grasslands. Furthermore, our data show that it is difficult to extrapolate results from experimental studies to semi-natural ecosystems, although there is a need to investigate natural ecosystems to fully understand the relationship of biodiversity and ecosystem functioning.     

Biodiversity loss and the taxonomic bottleneck: emerging biodiversity science

Human domination of the Earth has resulted in dramatic changes to global and local patterns of biodiversity. Biodiversity is critical to human sustainability because it drives the ecosystem services that provide the core of our life-support system. As we, the human species, are the primary factor leading to the decline in biodiversity, we need detailed information about the biodiversity and species composition of specific locations in order to understand how different species contribute to ecosystem services and how humans can sustainably conserve and manage biodiversity. Taxonomy and ecology, two fundamental sciences that generate the knowledge about biodiversity, are associated with a number of limitations that prevent them from providing the information needed to fully understand the relevance of biodiversity in its entirety for human sustainability: (1) biodiversity conservation strategies that tend to be overly focused on research and policy on a global scale with little impact on local biodiversity; (2) the small knowledge base of extant global biodiversity; (3) a lack of much-needed site-specific data on the species composition of communities in human-dominated landscapes, which hinders ecosystem management and biodiversity conservation; (4) biodiversity studies with a lack of taxonomic precision; (5) a lack of taxonomic expertise and trained taxonomists; (6) a taxonomic bottleneck in biodiversity inventory and assessment; and (7) neglect of taxonomic resources and a lack of taxonomic service infrastructure for biodiversity science. These limitations are directly related to contemporary trends in research, conservation strategies, environmental stewardship, environmental education, sustainable development, and local site-specific conservation. Today’s biological knowledge is built on the known global biodiversity, which represents barely 20% of what is currently extant (commonly accepted estimate of 10 million species) on planet Earth. Much remains unexplored and unknown, particularly in hotspots regions of Africa, South Eastern Asia, and South and Central America, including many developing or underdeveloped countries, where localized biodiversity is scarcely studied or described. "Backyard biodiversity", defined as local biodiversity near human habitation, refers to the natural resources and capital for ecosystem services at the grassroots level, which urgently needs to be explored, documented, and conserved as it is the backbone of sustainable economic development in these countries. Beginning with early identification and documentation of local flora and fauna, taxonomy has documented global biodiversity and natural history based on the collection of "backyard biodiversity" specimens worldwide. However, this branch of science suffered a continuous decline in the latter half of the twentieth century, and has now reached a point of potential demise. At present there are very few professional taxonomists and trained local parataxonomists worldwide, while the need for, and demands on, taxonomic services by conservation and resource management communities are rapidly increasing. Systematic collections, the material basis of biodiversity information, have been neglected and abandoned, particularly at institutions of higher learning. Considering the rapid increase in the human population and urbanization, human sustainability requires new conceptual and practical approaches to refocusing and energizing the study of the biodiversity that is the core of natural resources for sustainable development and biotic capital for sustaining our life-support system. In this paper we aim to document and extrapolate the essence of biodiversity, discuss the state and nature of taxonomic demise, the trends of recent biodiversity studies, and suggest reasonable approaches to a biodiversity science to facilitate the expansion of global biodiversity knowledge and to create useful data on backyard biodiversity worldwide towards human sustainability.     

Species and structural diversity affect growth of oak,but not pine,in uneven-aged mature forests

The effects of mixing tree species on tree growth and stand production have been abundantly studied, mostly looking at tree species diversity effects while controlling for stand density and structure. Regarding the shift towards managing forests as complex adaptive systems, we also need insight into the effects of structural diversity. Strict forest reserves, left for spontaneous development, offer unique opportunities for studying the effects of diversity in tree species and stand structure. We used data from repeated inventories in ten forest reserves in the Netherlands and northern Belgium to study the growth of pine and oak. We investigated whether the diversity of a tree's local neighbourhood (i.e., species and structural diversity) is important in explaining its basal area growth. For the subcanopy oak trees, we found a negative effect of the tree species richness of the local neighbours, which – in the studied forests – was closely related to the share of shade-casting tree species in the neighbourhood. The growth of the taller oak trees was positively affected by the height diversity of the neighbour trees. Pine tree growth showed no relation with neighbourhood diversity. Tree growth decreased with neighbourhood density for both species (although no significant relationship was found for the small pines). We found no overall diversity-growth relationship in the studied uneven-aged mature forests; the relationship depended on tree species identity and the aspect of diversity considered (species vs. structural diversity).     

Questioning the ecosystem services argument for biodiversity conservation

One of the central justifications for the conservation of biodiversity is the notion that species diversity is essential for the maintenance of ecosystem services. However, an important observation overlooked by proponents of this argument is that most ecosystem services are provided not by whole ecosystems, but by any group of species that fulfils certain basic functional criteria. Distinguishing between services that are resilient in response to species decline, and those that are not, is a far less challenging task than identifying the precise influence on ecosystem functioning of rare species. Conservationists have been almost unanimous in their failure to acknowledge this distinction between resilient and sensitive ecosystem services. Not only does this threaten the credibility of conservation science, but also increases the likelihood that natural area management becomes hijacked by the demand that ecosystem service provision be made the dominant management criteria.     

The more the merrier: Multi-species experiments in ecology

A major objective in ecology is to find general patterns, and to establish the rules and underlying mechanisms that generate those patterns. Nevertheless, most of our current insights in ecology are based on case studies of a single or few species, whereas multi-species experimental studies remain rare. We underline the power of the multi-species experimental approach for addressing general ecological questions, e.g. on species environmental responses or on patterns of among- and within-species variation. We present simulations that show that the accuracy of estimates of between-group differences is increased by maximizing the number of species rather than the number of populations or individuals per species. Thus, the more species a multi-species experiment includes, the more powerful it is. In addition, we discuss some inevitable methodological challenges of multi-species experiments. While we acknowledge the value of single- or few-species experiments, we strongly advocate the use of multi-species experiments for addressing ecological questions at a more general level.     

Bio-economic resource management under threats of environmental catastrophes

We combine ecological and economic dynamics to study the management of a natural resource that supports both ecosystem and human needs. Shrinking the resource base introduces a threat of occurrence of catastrophic ecological events, such as sudden ecosystem collapse. The occurrence conditions involve uncertainty of various types, and the distinction among these types is important for optimal resource management. When uncertainty is due to our ignorance of some aspects of the underlying ecology, the isolated equilibrium states characterizing optimal exploitation for many renewable resource problems become equilibrium intervals. Genuinely stochastic events shift the optimal equilibrium states, but maintain the structure of isolated equilibria.     

Resistance to rust fungi in Lolium perenne depends on within-species variation and performance of the host species in grasslands of different plant diversity

The hypothesis that plant species diversity and genetic variation of the host species decrease the severity of plant diseases is supported by studies of agricultural systems, but experimental evidence from more complex systems is scarce. In an experiment with grassland communities of varying species richness (1, 2, 4, 8, 16, and 60 species) and functional group richness (1, 2, 3, and 4 functional groups), we used different cultivars of Lolium perenne (perennial ryegrass) to study effects of biodiversity and cultivar identity on the occurrence and severity of foliar fungal diseases caused by Puccinia coronata (crown rust) and P. graminis (stem rust). Cultivar monocultures of perennial ryegrass revealed strong differences in pathogen susceptibility among these cultivars. Disease intensity caused by both rust fungi decreased significantly with growing species richness of species mixtures. The response to the diversity gradient was related to the decreased density and size of the host individuals with increasing species richness. The occurrence of other grass species known to be possible hosts of the pathogens in the experimental mixtures did not promote disease intensity in L. perenne, indicating that there was a high host specificity of pathogen strains. Differences in pathogen susceptibility among perennial ryegrass cultivars persisted independent of diversity treatment, host density and host individual size, but resulted in a cultivar-specific pattern of changes in pathogen infestation across the species-richness gradient. Our study provided evidence that within-species variation in pathogen susceptibility and competitive interactions of the host species with the environment, as caused by species diversity treatments, are key determinants of the occurrence and severity of fungal diseases. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Information pyramids for informed biodiversity conservation

We discuss a paradigm for informed ecosystem management that provides a quantitative and rigorous foundation for informing conservation decisions and sustainable ecosystem management. Information pyramids incorporate conceptual and technological advances in ecosystem depiction and provide a framework for the integration and generalization of raw data into forms that are spatially extensive and at the appropriate level of generalization for a particular use. The basic tenets of the pyramid are: (1) Higher levels of the pyramid are entirely derived from a foundation of underlying data. (2) The process of generalization and integration upward should be objective and explicit. (3) Pyramids for different purposes often overlap, with common data and common methods for integration. (4) All levels of the pyramid should be developed together, including base data, methods and kinds of integration, and algorithms for using the information for planning and decision-making. Information pyramids are a powerful approach to organizing research science, and provide a mechanism by which research, data collection, storage and generalization can be focused on conservation outcomes. Common data and methods lead to increased efficiency, while also allowing for separate disciplines and programs. A case study of an integrated pyramid from New Zealand is discussed, which illustrates the characteristics of information pyramids. Components of this pyramid are discussed that provide examples of integration and generalization at various levels of the pyramid, from base data, to derived data, to spatial predictions and classifications, to a method of integrating this information into conservation decisions.     

Predicting productivity in tropical reservoirs: The roles of phytoplankton taxonomic and functional diversity

Primary productivity is intimately linked with biodiversity and ecosystem functioning. Much of what is known today about such relationship has been based on the manipulation of species richness. Other facets of biodiversity, such as functional diversity, have been neglected within this framework, particularly in freshwater systems. We assess the adequacy of different diversity measures, from species richness and evenness, to functional groups richness and functional diversity indices, to predict primary productivity in 19 tropical reservoirs of central Brazil, built to generate hydroelectric energy. We applied linear mixed models (and model selection based on the Akaike’s information criterion) to achieve our goal, using chlorophyll-a concentration as a surrogate for primary productivity. A total of 412 species were collected in this study. Overall we found a positive relation between productivity and diversity, with functional evenness representing the only exception. The most parsimonious models never included functional group classifications, with at least one continuous measure of functional diversity being present in many models. The best model included only species richness and explained 24.1% of variability in productivity. We therefore advise the use of species richness as an indicator of productivity in tropical freshwater environments. However, since the productivity–diversity relationship is known to be scale dependent, we recommend the use of continuous measures of functional diversity in future biodiversity and ecosystem functioning studies, in order to be certain that all functional differences between communities are being accounted for.     

A review and a framework for the integration of biodiversity monitoring at the habitat level

The monitoring of biodiversity at the level of habitats is becoming widespread in Europe and elsewhere as countries establish national habitat monitoring systems and various organisations initiate regional and local schemes. Parallel to this growth, it is increasingly important to address biodiversity changes on large spatial (e.g. continental) and temporal (e.g. decade-long) scales, which requires the integration of currently ongoing monitoring efforts. Here we review habitat monitoring and develop a framework for integrating data or activities across habitat monitoring schemes. We first identify three basic properties of monitoring activities: spatial aspect (explicitly spatial vs. non-spatial), documentation of spatial variation (field mapping vs. remote sensing) and coverage of habitats (all habitats or specific habitats in an area), and six classes of monitoring schemes based on these properties. Then we explore tasks essential for integrating schemes both within and across the major classes. Finally, we evaluate the need and potential for integration of currently existing schemes by drawing on data collected on European habitat monitoring in the EuMon project. Our results suggest a dire need for integration if we are to measure biodiversity changes across large spatial and temporal scales regarding the 2010 target and beyond. We also make recommendations for an integrated pan-European habitat monitoring scheme. Such a scheme should be based on remote sensing to record changes in land cover and habitat types over large scales, with complementary field mapping using unified methodology to provide ground truthing and to monitor small-scale changes, at least in habitat types of conservation importance.     

Exploring spatial indicators for biodiversity accounting

In the context of the System for Environmental-Economic Accounting, biodiversity accounting is being developed as a tool to monitor and increase the understanding of human impacts on biodiversity. Biodiversity accounting aims to structurally measure, monitor and map changes in multiple biodiversity components, as an integral part of a larger system of ecosystem accounts. Both indicators relevant for ecosystem functioning and indicators that reflect the non-use values of biodiversity can be included in biodiversity accounting. In this paper we focus on the latter, and we test the potential applicability of a set of species indicators for developing a biodiversity account in Limburg province, the Netherlands. In particular, we map and analyse a range of indicators reflecting species richness, the presence of rare and threatened species and species diversity. We test spatial correlation to identify the minimum set of indicators that would need to be included in the account. We also evaluate individual indicators using eight different criteria. We show that, in Limburg province, a set of indicators covering at least five species groups is required, and that it would be most meaningful to have indicators reflecting the occurrence of threatened species. However, data availability as well as the most suitable set of indicators are likely to differ between areas, and case studies in other countries are required to support the selection of indicators for biodiversity accounting in an international framework.     

The loss of biodiversity conservation in EU research programmes: Thematic shifts in biodiversity wording in the environment themes of EU research programmes FP7 and Horizon 2020

Society has been seeking ways to express biodiversity's value to stimulate its protection. Economic valuation of ecosystem services has had limited success to motivate biodiversity protection and reaching the EU 2020 biodiversity strategy targets is in danger of failure. The expression of biodiversity's value in policy documents thus becomes a topic of discussion, because it greatly influences the ways policy makers think about environmental problems. We present an analysis of the word use related to biodiversity conservation versus ecosystem services in the environment themes of the FP7 and Horizon 2020 research work programs of the European Commission in the period of 2007–2014, and the projects accepted under these themes. We conclude first that biodiversity was lost as a topic in the transition from FP7 to Horizon 2020, accompanied by a three-quarters loss of biodiversity topics in the projects accepted under these research work programs. Moreover, the use of ‘ecosystem services’ was 1.5 times higher at the end of that period compared to the beginning in the research work programs, to the detriment of the use of ‘sustainability’ and ‘conservation’ which halved during that same period. In the light of international commitments to biodiversity conservation, the focus toward ecosystem services and away from conservation is of great concern.     

Improved water retention links high species richness with increased productivity in arctic tundra moss communities

A positive relationship between plant species richness and ecosystem functioning has been found in a number of experimental studies. Positive species interactions at high species numbers have been suggested as a cause, but mechanisms driving positive interactions have not often been tested. In this experiment we asked three questions: (1) What is the relationship between species richness and productivity in experimentally constructed moss communities? (2) Is this relationship affected by plant density? and (3) Can changes in moisture absorption and retention explain observed relationships? To answer these questions we exposed arctic tundra moss communities of different species richness levels (1–11 species) and two different densities in the greenhouse to two levels of drought (short and long). Biomass (by the community and individual species), height and community moisture absorption and retention were measured as response variables. High species diversity increased productivity (more so in low-density plots than in high-density plots), but only when plots were watered regularly. Plot moisture retention was improved at high species richness as well, and plant height and variation in height was increased compared to plants in monoculture. Under high-density and short-drought conditions 10 out of 12 species grew better in mixture than in monoculture, but under the long drought treatment only six species did. A positive feedback loop between biomass and improved humidity under high diversity was supported by path analysis. We conclude that in this community the relationship between species richness and productivity depends on moisture availability and density, with improved water absorption and retention likely to be the mechanism for increased plant growth when drought periods are short. Furthermore, since this is the opposite of what has been found for temperate moss communities, conclusions from one system cannot automatically be extrapolated to other systems.     

Think ratio! A stoichiometric view on biodiversity–ecosystem functioning research

Ecological stoichiometry (ES) has become one of the most pervasive theoretical frameworks in environmental sciences and biology in the last two decades. ES allows predicting processes on all organizational levels from subcellular structures to ecosystems by relating the elemental composition and demand of organisms to the relative availability of resources. However, ES has been rarely used to understand and predict the relationship between biodiversity and ecosystem functioning (BEF), although ES would be ideally suited as it makes predictions on both population processes underlying biodiversity as well as on matter transformations underlying ecosystem processes. Here, we propose to link the two fields of research on ES and BEF relationships and highlight a number of potential avenues for further research. First, we cast a stoichiometric view on drivers of biodiversity change. Second, we address the stoichiometric underpinning of biodiversity–productivity relationships. Third, we discuss potential interactions between stoichiometry and diversity in a food web context.     

Do experiments exploring plant diversity–ecosystem functioning relationships inform how biodiversity loss impacts natural ecosystems?

An enormous recent research effort focused on how plant biodiversity (notably species richness) influences ecosystem functioning, usually through experiments in which diversity is varied through random draws of species from a species pool. Such experiments are increasingly used to predict how species losses influence ecosystem functioning in ‘real’ ecosystems. However, this assumes that comparisons of experimental communities with low vs high species richness are analogous to comparisons of natural communities from which species either have or have not been lost. I explore the validity of this assumption, and highlight difficulties in using such experiments to draw conclusions about the ecosystem consequences of biodiversity loss in natural systems. Notably, these experiments do not mimic what happens in real ecosystems either when local extinctions occur or when species losses are offset by gains of new species. Despite limitations, this single experimental approach for studying how biodiversity loss affects ecosystems has often been advocated and implemented at the expense of other approaches; this limits understanding of how natural ecosystems respond to biodiversity loss. I conclude that a broader spectrum of approaches, and more explicit consideration of how species losses and gains operate in concert to influence ecosystems, will help progress this field.