Transgressive Overyielding Research Articles

Benefits of sward diversity for agricultural grasslands

A pan-European experiment carried out at 28 sites across Europe showed strong benefits of sward diversity in agricultural grasslands. We systematically varied the relative abundance of four agronomic plant species (sown species evenness), and found that 4-species mixtures yielded more forage than could be expected on the basis of the monoculture yields. Mixtures generally yielded more than even the best performing monoculture (transgressive overyielding). Mixtures strongly reduced the incidence of unsown species in the sward. These diversity effects were consistent over the wide range of environmental conditions and persisted over three harvest years and in highly fertilized conditions. These results indicate a strong potential for agronomic mixtures to contribute to more sustainable agricultural systems. Agronomic diversity can improve forage yield and reduce weed invasion in intensively managed grasslands, and may also enhance the provision of other ecosystem services.

Can complementarity in water use help to explain diversity–productivity relationships in experimental grassland plots?

Positive diversity-productivity relationships have repeatedly been found in experimental grassland plots, but mechanistic explanations are still under debate. We tested whether complementarity for the exploitation of the soil water resource helps to explain these relationships. In the dry summer of 2003, evapotranspiration (ET) was assessed at the Swedish BIODEPTH site using two different approaches: snapshot measurements of canopy surface temperature and simulation of time-accumulated ET by means of a soil water balance model. More diverse plots were characterized by lower surface temperatures and higher accumulated ET. Transgressive overyielding tests revealed that ET in polycultures was higher than in the best-performing monocultures, but this pattern was reversed at high degrees of water stress. Our results indicate that a more complete exploitation of soil water by more diverse grassland systems is on the one hand likely to be a driver for their increased biomass production, but on the other hand causes the more diverse communities to be affected earlier by drought. Nevertheless, the results also suggest that productivity may (at least partially) be maintained under dry conditions due to the higher likelihood of including drought-tolerant species in the more diverse communities.

Ecosystem functioning in stream assemblages from different regions: contrasting responses to variation in detritivore richness, evenness and density

1. The diversity of species traits in a biological assemblage varies not only with species richness, but also with species evenness and organism density, which together influence the concentration of traits within functional guilds. Potential trait diversity at local scales is also constrained by the regional species pool. Implications of such variation for spatio-temporal variability in biodiversity-ecosystem functioning relationships are likely to be complex, but are poorly understood. 2. In microcosm experiments conducted at laboratories in Sweden, Ireland and Romania, we investigated effects of species richness, evenness and density of stream-living detritivores on two related processes: detritivore leaf-processing efficiency (LPE) and growth. Assemblage composition varied among laboratories: one taxonomic order (Plecoptera) was studied in Sweden, whereas two orders, encompassing wider trait variation, were studied in Romania (Trichoptera and Plecoptera) and Ireland (Trichoptera and Isopoda). 3. Relationships between density and both LPE and growth ranged from negative to positive across the study species, highlighting the potential for density-dependent variation in process rates to alter ecosystem functioning, but indicating that such effects depend on species identity. 4. LPE varied with species diversity in the two more heterogeneous assemblages, but whereas LPE in the Romanian study was generally enhanced as richness increased, LPE in the Irish study increased only in less-even polycultures dominated by particular species. Transgressive overyielding was detected in the Irish experiment, indicating complementary resource use and/or facilitation (complementarity). These mechanisms could not be distinguished from the selection effect in the Romanian study. 5. Growth was elevated in Romanian species mixtures, reflecting positive complementarity, but lower than expected growth in some Swedish mixtures was associated with negative complementarity, indicating interspecific interference competition. 6. Our results emphasize the potential importance of detritivore diversity for stream ecosystem functioning, but both the effects of diversity on the studied processes, and the mechanisms underlying those effects, were specific to each assemblage and process. Such variability highlights challenges in generalizing impacts of diversity change for functional integrity in streams and other ecosystems in which the occurrence of important species traits fluctuates over relatively small spatio-temporal scales.

Experimental evidence for the effects of dung beetle functional group richness and composition on ecosystem function in a tropical forest

Much of the literature on the relationship between species richness or functional group richness and measures of ecosystem function focuses on a restricted set of ecosystem function measures and taxonomic groups. Few such studies have been carried out under realistic levels of diversity in the field, particularly in high diversity ecosystems such as tropical forests. We used exclusion experiments to study the effects of dung beetle functional group richness and composition on two interlinked and functionally important ecological processes, dung removal and secondary seed dispersal, in evergreen tropical forest in Sabah, Malaysian Borneo. Overall, both dung and seed removal increased with dung beetle functional group richness. However, levels of ecosystem functioning were idiosyncratic depending on the identity of the functional groups present, indicating an important role for functional group composition. There was no evidence for interference or competition among functional groups. We found strong evidence for overyielding and transgressive overyielding, suggesting complementarity or facilitation among functional groups. Not all mixtures showed transgressive overyielding, so that complementarity was restricted to particular functional group combinations. Beetles in a single functional group (large nocturnal tunnellers) had a disproportionate influence on measures of ecosystem function: in their absence dung removal is reduced by approximately 75%. However, a full complement of functional groups is required to maximize ecosystem functioning. This study highlights the importance of both functional group identity and species composition in determining the ecosystem consequences of extinctions or altered patterns in the relative abundance of species.

Effects of species and functional group richness on production in two fertility environments: an experiment with communities of perennial plants

The effects of species and functional group richness of sown species and fertilization on productivity and invasion of colonizers (i.e. unsown species) were studied in a three year field experiment. Both species and functional group numbers positively affected the average yield of sown species, but the effect changed among the three years. The biomass of colonizers generally decreased with the biomass of sown species, and so was negatively affected by the diversity characteristics of sown species. All three characteristics of biodiversity (selection, complementarity and overyielding effects) varied greatly during the study period, depending on both species and functional group richness and nutrient amendment. Functional differences among species led to the detection of high values of complementarity (resource use complementarity and/or facilitation), however this effect was not sufficient to cause transgressive overyielding. The compositional difference between fertilized and non-fertilized control plots increased with both time and species richness, whereas the relative difference in productivity decreased. This shows that the changes in species composition toward the one best-suited to the given environment, can lead to optimal resource use among plant species, and, as a consequence, diminish variation in community functioning in a changing environment.

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.

Effect of functional group richness and species richness in manipulated productivity–diversity studies: a glasshouse pot experiment

Species and functional group (grasses, legumes, creeping nonlegume forbs, rosette nonlegume forbs) richness of species assemblages composed of 16 species from four functional plant groups were manipulated to evaluate the productivity-diversity relationships in a greenhouse pot experiment. Pots were filled with sand, and supplied at two levels of nutrients. The plants were grown in monocultures, two, four, eight and 16 species mixtures. Individual two, four, and eight species mixtures differed in the richness of functional groups. Although the two characteristics of biodiversity, i.e. species and functional group richness, were necessarily correlated, it was shown that it is possible to separate their effect statistically, and also test for their common effect without pronounced loss of test power. There was a pronounced increase of average aboveground biomass and a mild increase in belowground biomass with biodiversity. The effect of functional group richness was more pronounced than the effect of the number of species. By using the method of Loreau and Hector (Nature 411 (2001) 72), selection and complementarity effects were statistically separated, and the overyielding index was calculated as a ratio of the productivity of a mixture to the productivity of its most productive component (to demonstrate transgressive overyielding). Positive values of complementarity and transgressive overyielding were both found, particularly in some rich communities and under high nutrient levels. Complementarity significantly increased only with functional group richness and mainly under high nutrients in the belowground biomass. Some species, when grown in monocultures, had decreased productivity under higher nutrients, and thus were more productive in mixtures than in monocultures. It seems that those species suffered from too high nutrient levels when grown in monocultures, but not in the presence of other species, which were able to use the nutrients in high concentrations and effectively decrease the nutrient levels. As a consequence, mixtures of high diversity were always more productive under high nutrients. The difference in species proportions between high and low nutrients, characterized by chord distance, increased with species richness. The relative change in productivity decreased with the number of functional groups. This suggests that species richness might lead to stabilization of aggregate characteristics (like total productivity) under changing environmental conditions by changing the proportions of individual species.

Inorganic soil nitrogen under grassland plant communities of different species composition and diversity

We measured aboveground plant biomass and soil inorganic nitrogen pools in a biodiversity experiment in northern Sweden, with plant species richness ranging from 1 to 12 species. In general, biomass increased and nitrate pools decreased with increasing species richness. Transgressive overyielding of mixed plant communities compared to the most productive of the corresponding monocultures occurred in communities with and without legumes. N2‐fixing legumes had a fertilizing function, while non‐legumes had a N retaining function. Plant communities with only legumes had a positive correlation between biomass and soil nitrate content, whereas in plant communities without legumes they were negatively correlated. Both nitrate and ammonium soil pools in mixed non‐legume communities were approximately equal to the lowest observed in the corresponding monocultures. In mixed legume/non‐legume communities, no correlation was found for soil nitrate with either biomass or legume biomass as percentage of total biomass. The idea of complementarity among species in nitrogen acquisition was supported in both pure non‐legume and mixed non‐legume/legume communities. In the latter, however, facilitation through increased nitrogen availability and retention, was probably dominating. Our results suggest that diversity effects on biomass and soil N pools through resource use complementarity depend on the functional traits of species, especially N2fixation or high productivity.

Overyielding in experimental grassland communities – irrespective of species pool or spatial scale

AbstractIn a large integrated biodiversity project (‘The Jena Experiment’ in Germany) we established two experiments, one with a pool of 60 plant species that ranged broadly from dominant to subordinate competitors on large 20 × 20 m and small 3.5 × 3.5 m plots (= main experiment), and one with a pool of nine potentially dominant species on small 3.5 × 3.5 m plots (= dominance experiment). We found identical positive species richness–aboveground productivity relationships in the main experiment at both scales. This result suggests that scaling up, at least over the short term, is appropriate in interpreting the implications of such experiments for larger‐scale patterns. The species richness–productivity relationship was more pronounced in the experiment with dominant species (46.7 and 82.6% yield increase compared to mean monoculture, respectively). Additionally, transgressive overyielding occurred more frequently in the dominance experiment (67.7% of cases) than in the main experiment (23.4% of cases). Additive partitioning and relative yield total analyses showed that both complementarity and selection effects contributed to the positive net biodiversity effect.

Modelling the joint effects of predator and prey diversity on total prey biomass

Summary Niche differentiation is widely thought to promote both coexistence of many species within a trophic level, and higher total biomass of those species. However, the joint effects of changes in diversity within multiple trophic levels are unknown. The potential for niche differences to promote high diversity and biomass within a trophic level may depend on the diversity and composition of adjacent trophic levels. Using a simple mechanistic model, I examine the joint consequences of changes in diversity on two adjacent trophic levels (‘predators’ and ‘prey’) for the total biomass of prey. Niche differences among prey reflect a trade‐off between ability to compete for resources and ability to avoid/sustain predation. Niche differences among predators reflect differences in ability to consume different prey species. Total prey biomass is a complex, joint function of prey and predator diversity. In particular, transgressive overyielding (when a diverse polyculture of prey outperforms any prey monoculture) can occur for some parameter values when a specialist predator mediates prey coexistence, and for a different set of parameter values when a specialist and a generalist predator together mediate prey coexistence, but is impossible when a generalist predator alone mediates prey coexistence. Increased initial predator diversity tends to reduce total prey biomass even when predators cannot coexist, due to a ‘selection’ effect for the most effective predator. Predators coexist only in prey polycultures. Coexisting predators can, but do not necessarily, reduce total prey biomass to a lower level than could any predator alone. The niche differences that permit coexistence strongly constrain the rank order of total prey biomass across all prey and predator combinations. Knowledge of the key trade‐offs that promote coexistence within food webs can allow powerful qualitative predictions of the effects of species loss on ecosystem‐level properties such as total biomass.

Overyielding among plant functional groups in a long‐term experiment

AbstractA recent debate among ecologists has focused on mechanisms by which species diversity might affect net primary productivity. Communities with more species could use a greater variety of resource capture characteristics, leading to greater use of limiting resources (complementarity) and therefore greater productivity (overyielding). Recent experiments, however, have shown a variety of relationships between diversity and productivity. In an experiment on serpentine grassland communities spanning 8 years, we found that overyielding increased several years after plot establishment. Overyielding varied greatly depending on the functional characteristics of the species involved and the biotic and abiotic environment (particularly water availability). While functional differences among species led to strong complementarity and facilitation, these effects were not sufficient to cause significant transgressive overyielding or consistent increases in productivity with increased plant diversity. These results suggest that greater absolute production with greater diversity may be restricted to particular species combinations or environmental conditions.

The long‐term relationship between plant diversity and total plant biomass depends on the mechanism maintaining diversity

The relationship between plant diversity and total plant biomass is of great current interest in ecology. Niche differences among plants are widely thought to promote both plant coexistence, and higher biomass in more diverse polycultures. Using simple mechanistic models, I demonstrate that not all niche differences among plants are equally likely to promote high total biomass in polyculture. In particular, transgressive overyielding (when a polyculture of plants outperforms any monoculture) occurs for a wide range of parameter values when plants coexist in polyculture due to differences in resource use. Transgressive overyielding occurs only for a limited set of parameter values when specialist herbivores mediate plant coexistence, and is impossible when generalist herbivores mediate coexistence. Niche differences among plants promote high biomass in polyculture only when plants coexist in polyculture at the expense of other trophic levels – that is, by converting into biomass resources that would otherwise be bound in herbivores, or exist in a free state. A major challenge for future work will be to identify the joint consequences of different coexistence mechanisms for plant diversity and ecosystem performance.

Integrating methods that investigate how complementarity influences ecosystem functioning

Separating the mechanisms that influence ecosystem functioning has been a goal of recent high profile experiments. Integrating the various experimental and analytical methods that attempt this goal across terrestrial and aquatic ecosystems, as well as careful definition of ‘complementarity’, produces novel insights and valuable lessons about new directions for research. (1) Experimental designs differ in temporal scale and whether standing stock or another ecosystem process was the response variable. (2) Mathematically identical variables in different designs have contrasting ecological interpretations. For example, different sets of ecological processes can contribute to different variables in different experimental designs. (3) The frequency of transgressive overyielding of standing stock (e.g. total above ground biomass) in polycultures implies little about the prevalence of transgressive overyielding in other ecosystem processes. (4) Measuring the contribution to ecosystem functioning of individual species, rather than just total ecosystem functioning of a polyculture, is not essential for estimating effects of complementarity. (5) Further research will profit from distinguishing standing stock from all other ecosystem functions. (6) None of the analytic methods can distinguish the effects of individual processes or mechanisms such as resource use differentiation, facilitation, or allelopathy, for which additional experimental treatments are required.