Overyielding among plant functional groups in a long-term experiment

A 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.     

Cascading extinctions, functional complementarity, and selection in two-trophic-level model communities: A trait-based mechanistic approach

The influence of diversity on ecosystem functioning and ecosystem services is now well established. Yet predictive mechanistic models that link species traits and community-level processes remain scarce, particularly for multitrophic systems. Here we revisit MacArthur's classical consumer resource model and develop a trait-based approach to predict the effects of consumer diversity on cascading extinctions and aggregated ecosystem processes in a two-trophic-level system. We show that functionally redundant efficient consumers generate top-down cascading extinctions. This counterintuitive result reveals the limits of the functional redundancy concept to predict the consequences of species deletion. Our model also predicts that the biodiversity-ecosystem functioning relationship is different for different ecosystem processes and depends on the range of variation of consumer traits in the regional species pool, which determines the sign of selection effects. Lastly, competition among resources and consumer generalism both weaken complementarity effects, which suggests that selection effects may prevail at higher trophic levels. Our work emphasizes the potential of trait-based approaches for transforming biodiversity and ecosystem functioning research into a more predictive science.     

Effects of plant diversity on invertebrate herbivory in experimental grassland

The rate at which a plant species is attacked by invertebrate herbivores has been hypothesized to depend on plant species richness, yet empirical evidence is scarce. Current theory predicts higher herbivore damage in monocultures than in species-rich mixtures. We quantified herbivore damage by insects and molluscs to plants in experimental plots established in 2002 from a species pool of 60 species of Central European Arrhenatherum grasslands. Plots differed in plant species richness (1, 2, 4, 8, 16, 60 species), number of functional groups (1, 2, 3, 4), functional group and species composition. We estimated herbivore damage by insects and molluscs at the level of transplanted plant individuals (“phytometer” species Plantago lanceolata, Trifolium pratense, Rumex acetosa) and of the entire plant community during 2003 and 2004. In contrast to previous studies, our design allows specific predictions about the relative contributions of functional diversity, plant functional identity, and species richness in relation to herbivory. Additionally, the phytometer approach is new to biodiversity-herbivory studies, allowing estimates of species-specific herbivory rates within the larger biodiversity-ecosystem functioning context. Herbivory in phytometers and experimental communities tended to increase with plant species richness and the number of plant functional groups, but the effects were rarely significant. Herbivory in phytometers was in some cases positively correlated with community biomass or leaf area index. The most important factor influencing invertebrate herbivory was the presence of particular plant functional groups. Legume (grass) presence strongly increased (decreased) herbivory at the community level. The opposite pattern was found for herbivory in T. pratense phytometers. We conclude that (1) plant species richness is much less important than previously thought and (2) plant functional identity is a much better predictor of invertebrate herbivory in temperate grassland ecosystems.     

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.     

The antioxidation-related functional structure of plant communities: Understanding antioxidation at the plant community level

In the case of ecological stresses, reactive oxygen species (ROS) can be overproduced in plant cells, leading to lipid peroxidation that causes damage or death to cells. To prevent damage, plant tissues contain several antioxidants that scavenge ROS. However, antioxidation at the plant community level still remains unknown and may provide an insight into ecosystem functioning regarding stress resistance. To understand the property, we established the antioxidation-related functional structure based on the concept of the functional structure and the activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), reduced glutathione (GSH), carotenoid (Car) and proline (Pro) and the relative abundance of component species in natural plant communities on the Chinese hilly-gullied Loess Plateau. An information-theoretic (I-T) approach was used to infer the effects of antioxidation-related functional components and stresses that we investigated on lipid peroxidation at the plant community level quantified by the community-weighted mean of malondialdehyde (MDA). We found that the induction of lipid peroxidation was more closely associated with a soil available nitrogen deficiency than it was with an available phosphorus deficiency. However, the inducing effect of soil available nitrogen was finite. The prevention of lipid peroxidation was more closely associated with the community-weighted means of GSH and Pro (CWM GSH and CWM Pro) than it was with other antioxidation-related functional components. However, the efficiency of CWM Pro was quite low; CWM GSH exhibited inefficiency. In addition, antioxidation-related functional components were affected by neither soil available nitrogen nor available phosphorus. Furthermore, by estimating the relative weights of the antioxidants and considering the feasibility of community assemblages, we proposed that Stipa grandis, Leymus secalinus, Stipa bungeana, Phragmites australis, Potentilla tanacetifolia, Artemisia gmelinii, Artemisia scoparia, Heteropappus altaicus and Syringa oblata could be utilized in community assemblages to achieve an antioxidation-functional target. Additionally, appropriate phosphorus application for A. gmelinii and H. altaicus might contribute to maintaining their antioxidation.     

Species diversity and functional assessment of macroinvertebrate communities in Austrian rivers

We applied an extensive data set from 211 locations along Austrian rivers to assess community structure and the ratios of functional feeding groups of benthic macroinvertebrates. A total of 569 taxa have been identified. At the catchment scale, the Enns, Salzach, and Traun Rivers exhibited the highest taxa richness whereas the Inn River showed the lowest richness. Beta-diversity was highest along the impounded and fragmented Enns and Drau Rivers. Consequently, high corridor diversity corresponded to a low degree of nestedness. Overall, scrapers and gathering-collectors dominated the benthic community. Further, the relationship between habitat conditions and metrics based on functional feeding groups were statistically analyzed to validate the potential of these metrics as indicators of ecosystem attributes. We examined four major ecosystem attributes: species diversity, material cycling, longitudinal material transport, and lateral material input. Multiple regression analyses for midorder rivers demonstrated that metrics were significantly related to habitat conditions. For example, the metric set indicating primary production was positively correlated with periphyton cover, dissolved oxygen, dominant sediment size, and average annual discharge. Overall, most metrics exhibited unique responses to habitat conditions, implying that they are useful proxies of ecosystem attributes. Thus, a function-based approach based on macroinvertebrates has the potential to become an effective tool for the assessment of river ecosystems.     

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.     

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.     

Trait convergence and trait divergence in herbaceous plant communities: Mechanisms and consequences

In landscapes subject to intensive agriculture, both soil fertility and vegetation disturbance are capable of impacting strongly, evenly and simultaneously on the herbaceous plant cover and each tends to impose uniformity on the traits of constituent species. In more natural and ancient grasslands greater spatial and temporal variation in both productivity and disturbance occurs and both factors have been implicated in the maintenance of species‐richness in herbaceous communities. However, empirical data suggest that disturbance is the more potent driver of trait differentiation and species co‐existence at a local scale. This may arise from the great diversity in opportunities for establishment, growth or reproduction that arise when the intensity of competition is reduced by damage to the vegetation. In contrast to the diversifying effects of local disturbances, productivity‐related plant traits (growth rate, leaf longevity, leaf chemistry, leaf toughness, decomposition rate) appear to be less variable on a local scale. This difference in the effects of the productivity and disturbance filters arises from the relative constancy of productivity within the community and the diversity in agency and in spatial and temporal scales exhibited by disturbance events. Also, evolutionary responses to disturbances involve minor adaptive shifts in phenological and regenerative traits and are more likely to occur as micro‐evolutionary steps than the shifts in linked traits in the core physiology associated with the capacity to exploit productive and unproductive habitats. During the assembly of a community and over its subsequent lifespan filters with diversifying and convergent effects may operate simultaneously on recruitment from the local species pool and impose contrasted effects on the similarity of the trait values exhibited by co‐existing species. Moreover, as a consequence of the frequent association of productivity with the convergence filter, an additional difference is predicted in terms of the effects of the two filters on ecosystem functioning. Convergence in traits selected by the productivity filter will exert effects on both the plant community and the ecosystem while divergent effects of the disturbance filter will be restricted to the plant community.     

Diversity and productivity of plant communities across the Inland Northwest,USA

No definitive explanation for the form of the relationship between species diversity and ecosystem productivity exists nor is there agreement on the mechanisms linking diversity and productivity across scales. Here, we examine changes in the form of the diversity–productivity relationship within and across the plant communities at three observational scales: plots, alliances, and physiognomic vegetation types (PVTs). Vascular plant richness data are from 4,760 20 m² vegetation field plots. Productivity estimates in grams carbon per square meter are from annual net primary productivity (ANPP) models. Analyses with generalized linear models confirm scale dependence in the species diversity–productivity relationship. At the plot focus, the observed diversity–productivity relationship was weak. When plot data were aggregated to a focus of vegetation alliances, a hump-shaped relationship was observed. Species turnover among plots cannot explain the observed hump-shaped relationship at the alliance focus because we used mean plot richness across plots as our index of species richness for alliances and PVTs. The sorting of alliances along the productivity gradient appears to follow regional patterns of moisture availability, with alliances that occupy dry environments occurring within the increasing phase of the hump-shaped pattern, alliances that occupy mesic to hydric environments occurring near the top or in the decreasing phase of the curve, and alliances that occupy the wettest environments having the fewest species and the highest ANPP. This pattern is consistent with the intermediate productivity theory but appears to be inconsistent with the predictions of water–energy theory.     

Plant functional diversity mediates the effects of vegetation and soil properties on community-level plant nitrogen use in the restoration of semiarid sandy grassland

Species-rich plant communities use nitrogen (N) more efficiently in grassland ecosystems; however, the role of plant functional diversity in affecting community level plant N-use has received little attention. We examined plant N content, stock and N-use efficiency at community-level along a restoration gradient of sandy grassland (mobile dune, semi-fixed dune, fixed dune and grassland) in Horqin Sand Land, northern China. We used the functional trait-based approach to examine how plant functional diversity, reflected by the most abundant species’ traits (community-weighted mean, CWM) and the dispersion of functional trait values (FDis), affected N-use efficiency in sandy grassland restoration. We further used the structure equation model (SEM) to evaluate the direct or indirect effects of plant species richness, biomass, functional diversity and soil properties on community-level plant N-use efficiency. We found that plant biomass and its N stock increased following sandy grassland restoration, and there were lower plant N content and higher N-use efficiency in semi-fixed dune, fixed dune and grassland as compared with mobile dune. N-use efficiency was positively associated with plant species richness, biomass, CWM plant height, CWM leaf C:N, FDis and soil gradient, but SEM results showed that species richness, CWM leaf C:N, plant biomass and FDis controlled by soil properties were the main factors exerting direct effects. CWM plant height also had a positive effect on N-use efficiency through its indirect effect on plant biomass. Soil gradient increased N-use efficiency through an indirect effect on vegetation rather than a direct effect. Final SEM models based on different plant functional diversity explained over 74% of variances in N-use efficiency. Effects of plant functional diversity on N-use efficiency supported both the mass ratio hypothesis and the complementarity hypothesis. Our results clearly highlight the important role of plant functional diversity in mediating the effects of vegetation and soil properties on community level plant N-use in sandy grassland ecosystems.     

Natural enemy diversity and biological control: Making sense of the context-dependency

Numerous studies have demonstrated that diverse predator assemblages can be more effective at controlling prey populations. Yet, other studies have shown no effect of predator diversity on prey mortality, or even negative effects (for example due to intraguild predation or interference). Much research emphasis has been placed on the traits of predators that maximise functional complementarity. However, comparatively less attention has been paid to the traits of the prey or habitat that may maximise predator diversity effects, even though there must be a variety of prey niches available to be partitioned in order for niche complementarity to occur. Following this logic, we review six hypotheses for when diverse enemy assemblages should be most effective: when 1) prey communities are diverse; 2) prey have complex life cycles; 3) prey are patchily distributed in space or time; 4) studies are conducted at larger spatial and temporal scales; 5) plant structures are complex; 6) prey are abundant. Many of these hypotheses lack direct tests, particularly in agricultural systems, but we find little or no direct or indirect support for hypotheses 1, 4, 5 and 6. However, previous work does provide some support for hypotheses 2 and 3. We discuss methods to test these hypotheses directly, and suggest that natural enemy diversity may only benefit the biological control of arthropods in heterogeneous systems.     

Density may alter diversity–productivity relationships in experimental plant communities

Contemporary biodiversity experiments, in which plant species richness is manipulated and aboveground productivity of the system measured, generally demonstrate that lowering plant species richness reduces productivity. However, we propose that community density may in part compensate for this reduction of productivity at low diversity. We conducted a factorial experiment in which plant functional group richness was held constant at three, while plant species richness increased from three to six to 12 species and community density from 440 to 1050 to 2525 seedlings m⁻². Response variables included density, evenness and above- and belowground biomass at harvest. The density gradient converged slightly during the course of the experiment due to about 10% mortality at the highest sowing density. Evenness measured in terms of aboveground biomass at harvest significantly declined with density, but the effect was weak. Overall, aboveground, belowground and total biomass increased significantly with species richness and community density. However, a significant interaction between species richness and community density occurred for both total and aboveground biomass, indicating that the diversity–productivity relationship was flatter at higher than at lower density. Thus, high species richness enabled low-density communities to reach productivity levels otherwise seen only at high density. The relative contributions of the three functional groups C₃, C₄ and nitrogen-fixers to aboveground biomass were less influenced by community density at high than at low species richness. We interpret the interaction effects between community density and species richness on community biomass by expanding findings about constant yield and size variation from monocultures to plant mixtures.     

辽河河口湿地自然保护区植物群落生态的研究

通过野外调查和定量分析,对辽河河口湿地自然保护区主要植物群落的结构、功能及其演替规律进行了研究。结果表明,土壤水盐含量与植物群落分布密切相关,用组平均法将４５个芦苇群落样方分为芦苇草甸和芦苇沼泽两大类,用ＰＣＡ排序对环境进行分类,确定出影响植物群落空间配置的主导因子是土壤水分和土壤盐分。     

Seed banks of a southern Australian wetland: the influence of water regime on the final floristic composition

The sediments of a southern Australian wetland complex were investigated to determine the germinable seed bank. The number of seeds ranged from 22,000 to 78,000 m^–2. Non-metric multi dimensional scaling (NMS) ordination based upon species composition separated the four basins but each contained the representatives of the major functional groups: terrestrial, amphibious and submerged. To test the influence of water regime on the final floristic composition derived from these seed banks, sediments were positioned at three elevations (0, 30 and 80 cm) and subjected to three hydrologic regimes (static water level, and draw down rates of 2.5 and 7 cm week^–1). Species compositions were analysed after 98 days via ordination. Despite significant differences in the initial seed bank composition the final floristic compositions were correlated with the water regime and independent of the initial seed bank composition. Species groups were segregated on a basis of whether sediments were continuously exposed to the atmosphere, the rate of draw down and the water depth. Moist sediments were dominated by species belonging to all the main functional groups. Sediments subject to rapid drying were dominated by terrestrial species whereas sediments that were flooded for the majority of the time were dominated by submerged species.     

Deep placement: A method of nitrogen fertilizer application compatible with algal nitrogen fixation in wetland rice soils

Summary The effect of different methods of nitrogen fertilizer application on the algal flora and biological nitrogen fixation (Acetylene-reducing activity) in a wetland rice soil was studied in pot and field experiments. Broadcast application of urea inhibited nitrogen fixation and favored the growth of green algae. In contrast, deep placement of urea supergranules (1–2 g urea granules) did not suppress the growth of N₂-fixing blue-green algae and permitted acetylene-reducing activity on the soil surface to continue virtually uninhibited.     

Soil and plant tests for available sulfur in wetland rice soils

Summary Soil tests, plant performance, and plant tissue analyses were used to study the availability of sulfur to wetland rice in 30 Philippine soils. The critical concentrations of available sulfur by the calcium phosphate, lithium chloride, ammonium acetate, and hydrochloric acid extractions were 9, 25, 30, and 5 mg/kg, respectively. The critical total sulfur limits were 0.11% in the shoot at maximum tillering 0.055% in the straw at maturity, and 0.065% in the grain. The critical N:S ratio was 15 in the shoot at maximum tillering, 14 in the straw at maturity, and 26 in the grain. The critical sulfate-sulfur limit was 150 mg/kg in the shoot at maximum tillering and 100 mg/kg in the straw at maturity. The critical sulfate-sulfur/total sulfur percentage ratio was 15% in the shoot at maximum tillering and the straw at maturity. Plant performance, judged by appearance and yield of dry matter, straw, and grain, was generally poorer in the sulfur deficient soils than in the other soils. Although the calcium phosphate and ammonium acetate methods gave a better correlation between plant performance and available sulfur than the others, all four methods separated sulfur-deficient soils from non-deficient ones. The hydrochloric acid method merits further study because it is simple and versatile.