Diversity-function Relationships Research Articles

The functional role of biodiversity in ecosystems: incorporating trophic complexity

Understanding how biodiversity affects functioning of ecosystems requires integrating diversity within trophic levels (horizontal diversity) and across trophic levels (vertical diversity, including food chain length and omnivory). We review theoretical and experimental progress toward this goal. Generally, experiments show that biomass and resource use increase similarly with horizontal diversity of either producers or consumers. Among prey, higher diversity often increases resistance to predation, due to increased probability of including inedible species and reduced efficiency of specialist predators confronted with diverse prey. Among predators, changing diversity can cascade to affect plant biomass, but the strength and sign of this effect depend on the degree of omnivory and prey behaviour. Horizontal and vertical diversity also interact: adding a trophic level can qualitatively change diversity effects at adjacent levels. Multitrophic interactions produce a richer variety of diversity-functioning relationships than the monotonic changes predicted for single trophic levels. This complexity depends on the degree of consumer dietary generalism, trade-offs between competitive ability and resistance to predation, intraguild predation and openness to migration. Although complementarity and selection effects occur in both animals and plants, few studies have conclusively documented the mechanisms mediating diversity effects. Understanding how biodiversity affects functioning of complex ecosystems will benefit from integrating theory and experiments with simulations and network-based approaches.

Evenness drives consistent diversity effects in intensive grassland systems across 28 European sites

1 Ecological and agronomic research suggests that increased crop diversity in species-poor intensive systems may improve their provision of ecosystem services. Such general predictions can have critical importance for worldwide food production and agricultural practice but are largely untested at higher levels of diversity. 2 We propose new methodology for the design and analysis of experiments to quantify diversity-function relationships. Our methodology can quantify the relative strength of inter-specific interactions that contribute to a functional response, and can disentangle the separate contributions of species richness and relative abundance. 3 Applying our methodology to data from a common experiment at 28 European sites, we show that the above-ground biomass of four-species mixtures (two legumes and two grasses) in intensive grassland systems was consistently greater than that expected from monoculture performance, even at high productivity levels. The magnitude of this effect generally resulted in transgressive overyielding. 4 A combined analysis of first-year results across sites showed that the additional performance of mixtures was driven by the number and strength of pairwise inter-specific interactions and the evenness of the community. In general, all pairwise interactions contributed equally to the additional performance of mixtures; the grass-grass and legume-legume interactions were as strong as those between grasses and legumes. 5 The combined analysis across geographical and temporal scales in our study provides a generality of interpretation of our results that would not have been possible from individual site analyses or experimentation at a single site. 6 Our four-species agricultural grassland communities have proved a simple yet relevant model system for experimentation and development of methodology in diversity-function research. Our study establishes that principles derived from biodiversity research in extensive, semi-natural grassland systems are applicable in intensively managed grasslands with agricultural plant species.

The ecology of restoration: historical links, emerging issues and unexplored realms

AbstractRestoration ecology is a young academic field, but one with enough history to judge it against past and current expectations of the science's potential. The practice of ecological restoration has been identified as providing ideal experimental settings for tests of ecological theory; restoration was to be the ‘acid test’ of our ecological understanding. Over the past decade, restoration science has gained a strong academic foothold, addressing problems faced by restoration practitioners, bringing new focus to existing ecological theory and fostering a handful of novel ecological ideas. In particular, recent advances in plant community ecology have been strongly linked with issues in ecological restoration. Evolving models of succession, assembly and state‐transition are at the heart of both community ecology and ecological restoration. Recent research on seed and recruitment limitation, soil processes, and diversity–function relationships also share strong links to restoration. Further opportunities may lie ahead in the ecology of plant ontogeny, and on the effects of contingency, such as year effects and priority effects. Ecology may inform current restoration practice, but there is considerable room for greater integration between academic scientists and restoration practitioners.

Biodiversity, ecosystem function and plant traits in mature and immature plant communities

Several studies of synthesized herbaceous plant communities have reported a positive relationship between vascular plant diversity and biomass production (Naeem et al. 1994; Tilman et al. 2001), and the European BIODEPTH experiment has been one of the most influential in this regard (Hector et al. 1999). Much, though not all, of the evidence for this relationship has come from experimental communities established from seed and harvested after only a year or two. It is not clear that such immature plant communities would be expected to provide reliable evidence of the relationship between diversity and ecosystem properties. Here we compare the relationship between diversity and biomass in BIODEPTH with that at the long-term monitoring site at Bibury in Gloucestershire, UK (Dunnett et al. 1998), a natural plant community that has much in common with BIODEPTH. The Bibury site offers a rare opportunity to address questions similar to those tackled by BIODEPTH, while avoiding some of the limitations of short-term experiments. If a positive relationship between vascular plant diversity and biomass production is an important phenomenon, one could argue that it should be detectable in natural vegetation. Previous attempts to explore the relationship between biodiversity and productivity in natural plant communities have been criticized on the grounds that both variables are frequently affected by additional (usually abiotic) variables (Lawton et al. 1998). For example, over a wide range of moderate-to-high biomass values, there is a well known negative relationship between biomass and species richness, driven by increasing competitive exclusion at higher productivity (Grime 1973; Grace 1999; Stevens et al. 2004). In experiments such as BIODEPTH, one aim is to exclude such natural sources of variation and create synthesized plant assemblages that vary only in artificially imposed levels of species richness. Of course, random environmental heterogeneity cannot be completely excluded, and Huston & McBride (2002) have shown how such heterogeneity can give rise to spurious diversity-function relationships. However, synthesized communities may also exhibit systematic variation in productivity equalling or exceeding that occurring in natural habitats, arising from the higher nitrogen availability in communities sown with N-fixing legumes. In BIODEPTH, addition of TriJolium pratense (nomenclature follows Stace 1997) increased biomass, on average, by 360 g m (Hector et al. 1999). As this increase was much larger than the effect of any other species, it is reasonable to attribute this effect to a direct result of the extra N fixed by Trifolium. The increase was also much larger than the effect of a change in diversity itself; doubling species richness increased biomass, on average, by only 80 g m-. The Bibury road verge lies on the north side of Akeman Street, originally a Roman road, in Gloucestershire, UK (National Grid reference SP 119048). The 700 m length of experimental verge was originally chosen

Soil animals alter plant litter diversity effects on decomposition

Most of the terrestrial net primary production enters the decomposer system as dead organic matter, and the subsequent recycling of C and nutrients are key processes for the functioning of ecosystems and the delivery of ecosystem goods and services. Although climatic and substrate quality controls are reasonably well understood, the functional role of biodiversity for biogeochemical cycles remains elusive. Here we ask how altering litter species diversity affects species-specific decomposition rates and whether large litter-feeding soil animals control the litter diversity-function relationship in a temperate forest ecosystem. We found that decomposition of a given litter species changed greatly in the presence of litters from other cooccurring species despite unaltered climatic conditions and litter chemistry. Most importantly, soil fauna determined the magnitude and direction of litter diversity effects. Our data show that litter species richness and soil fauna interactively determine rates of decomposition in a temperate forest, suggesting a combination of bottom-up and top-down controls of litter diversity effects on ecosystem C and nutrient cycling. These results provide evidence that, in ecosystems supporting a well developed soil macrofauna community, animal activity plays a fundamental role for altered decomposition in response to changing litter diversity, which in turn has important implications for biogeochemical cycles and the long-term functioning of ecosystems with ongoing biodiversity loss.

Biodiversity and ecosystem services in agricultural landscapes—are we asking the right questions?

The assumed relationship between biodiversity or local richness and the persistence of ‘ecosystem services’ (that can sustain productivity on-site as well as off-site, e.g. through regulation of water flow and storage) in agricultural landscapes has generated considerable interest and a range of experimental approaches. The abstraction level aimed for, however, may be too high to yield meaningful results. Many of the experiments on which evidence in favour or otherwise are based are artificial and do not support the bold generalisations to other spatial and temporal scales that are often made. Future investigations should utilise co-evolved communities, be structured to investigate the distinct roles of clearly defined functional groups, separate the effects of between- and within-group diversity and be conducted over a range of stress and disturbance situations. An integral part of agricultural intensification at the plot level is the deliberate reduction of diversity. This does not necessarily result in impairment of ecosystem services of direct relevance to the land user unless the hypothesised diversity–function threshold is breached by elimination of a key functional group or species. Key functions may also be substituted with petro-chemical energy in order to achieve perceived efficiencies in the production of specific goods. This can result in the maintenance of ecosystem services of importance to agricultural production at levels of biodiversity below the assumed ‘functional threshold’. However, it can also result in impairment of other services and under some conditions the de-linking of the diversity–function relationship. Avoidance of these effects or attempts to restore non-essential ecosystem services are only likely to be made by land users at the plot scale if direct economic benefit can be thereby achieved. At the plot and farm scales biodiversity is unlikely to be maintained for purposes other than those of direct use or ‘utilitarian’ benefits and often at levels lower than those necessary for maintenance of many ecosystem services. The exceptions may be traditional systems where intrinsic values (social customs) continue to provide reasons for diversity maintenance. High levels of biodiversity in managed landscapes are more likely to be maintained for reasons of intrinsic, serependic (‘option’ or ‘bequest’) values or utilitarian (‘direct use’) than for functional or ecosystem service values. The major opportunity for both maintaining ecosystem services and biodiversity outside conservation areas lies in promoting diversity of land-use at the landscape and farm rather than field scale. This requires, however, an economic and policy climate that favours diversification in land uses and diversity among land users. © 2004 Published by Elsevier B.V.

Species richness–decomposition relationships depend on species dominance

AbstractHuman disturbances both decrease the number of species in ecosystems and change their relative abundances. Here we present field evidence demonstrating that shifts in species abundances can have effects on ecosystem functioning that are as great as those from shifts in species richness. We investigated spatial and temporal variability of leaf decomposition rates and community metrics of leaf‐eating invertebrates (shredders) in streams. The shredder community composition dramatically influenced the diversity–function relationship; decomposition was much higher for a given species richness at sites with high species dominance than at sites where dominance was low. Decomposition rates also markedly depended on the identity of the dominant species. Further, dominance effects on decomposition varied seasonally and the number of species required for maintaining decomposition increased with increasing evenness. These findings reveal important but less obvious aspects of the biodiversity–ecosystem functioning relationship.

Sensitivity of Primary Production to Changes in the Architecture of Belowground Food Webs

The proposed mechanisms for the species diversity-function relationship in plant communities stress the recognition of functional properties of species, and interactions between plants and soil processes. As resource availability to plants is influenced by the architecture of decomposer food webs, it has been hypothesised that the diversity of decomposers can also control ecosystem processes, including primary production. We manipulated the complexity of soil animal communities in a miniecosysten experiment in which a boreal forest floor with birch seedlings infected with mycorrhizal fungi was created. The soil animal diversity ranged from zero to typical species richness of soil fauna (approximately 50 taxa) in coniferous forests. Between these extremes was a nested factorial design with manually assembled communities consisting of two sets of one-species, and two sets of five-species animal communities within fungivorous and microbi-detritivorous trophic groups. To investigate the role of predators in system functioning, the miniecosystems with fungivorous and microbidetritivorous fauna were established either with or without mesostigmatid mites as top predators. The miniecosystems were incubated in a climate chamber with varying illumination and temperature regimes for 40 weeks. Our experiment provides evidence that primary productivity is generally insensitive to variation at the species level or even at the level of trophic groups. Although top predators generally reduced prey population size, no effect was found on systen functioning. However, the removal of microbe- or detritus-feeding fauna, especially the microbi-detritivore Cognettia sphagnetorum reduced plant N uptake and accumulation of plant biomass. The functional importance of soil fauna was inversely related to the trophic position of the group. Our results suggest that ecosystem functioning is robust against species extinctions in belowground food webs, and that primary production is predominantly controlled by organisms at low trophic positions in the decomposer food web.

Which species? What kind of diversity? Which ecosystem function? Some problems in studies of relations between biodiversity and ecosystem function

I examine a number of problems that need to be identified and accounted for when examining the relationships between diversity and ecosystem function. Among these are measures of diversity and complexity in ecosystems: species richness, diversity indices, functional groups, keystone species, connectance, etc, all of which may be difficult to relate to ecosystem function. Several important distinctions, when testing diversity–function relationships empirically, are discussed: Diversity of functional groups, diversity within functional groups vs. total diversity; manipulating variables such as body-size distributions vs. manipulating diversity per se; effects of diversity vs. effects of biomass; and diversity–function relations under stable vs. changing conditions or perturbations. It is argued that for the management and development of sustainable ecosystems, it is probably more important to understand the linkages between key species or functional groups and ecosystem function, rather than focusing on species diversity. This is because there are possible mechanistic relations between what species do in ecosystems and ecosystem function. Diversity, being an abstract and aggregated property of the species in the context of communities and ecosystems, lacks such direct relations to ecosystem functions.