Invertebrate herbivory increases along an experimental gradient of grassland plant diversity

Plant diversity is a key driver of ecosystem functioning best documented for its influence on plant productivity. The strength and direction of plant diversity effects on species interactions across trophic levels are less clear. For example, with respect to the interactions between herbivorous invertebrates and plants, a number of competing hypotheses have been proposed that predict either increasing or decreasing community herbivory with increasing plant species richness. We investigated foliar herbivory rates and consumed leaf biomass along an experimental grassland plant diversity gradient in year eight after establishment. The gradient ranged from one to 60 plant species and manipulated also functional group richness (from one to four functional groups—legumes, grasses, small herbs, and tall herbs) and plant community composition. Measurements in monocultures of each plant species showed that functional groups differed in the quantity and quality of herbivory damage they experienced, with legumes being more damaged than grasses or non-legume herbs. In mixed plant communities, herbivory increased with plant diversity and the presence of two key plant functional groups in mixtures had a positive (legumes) and a negative (grasses) effect on levels of herbivory. Further, plant community biomass had a strong positive impact on consumed leaf biomass, but little effect on herbivory rates. Our results contribute detailed data from a well-established biodiversity experiment to a growing body of evidence suggesting that an increase of herbivory with increasing plant diversity is the rule rather than an exception. Considering documented effects of herbivory on other ecosystem functions and the increase of herbivory with plant diversity, levels of herbivory damage might not only be a result, but also a trigger within the diversity–productivity relationship.     

Community maturity, species saturation and the variant diversity-productivity relationships in grasslands

Detailed knowledge of the relationship between plant diversity and productivity is critical for advancing our understanding of ecosystem functioning and for achieving success in habitat restoration efforts. However, effects and interactions of diversity, succession and biotic invasions on productivity remain elusive. We studied newly established communities in relation to preexisting homogeneous vegetation invaded by exotic plants in the northern Great Plains, USA, at four study sites for 3 years. We observed variant diversity–productivity relationships for the seeded communities (generally positive monotonic at three sites and non-monotonic at the other site) but no relationships for the resident community or the seeded and resident communities combined at all sites and all years. Community richness was enhanced by seeding additional species but productivity was not. The optimal diversity (as indicated by maximum productivity) changed among sites and as the community developed. The findings shed new light on ecosystem functioning of biodiversity under different conditions and have important implications for restoration.     

Biodiversity, productivity and the temporal stability of productivity: patterns and processes

Theory predicts that the temporal stability of productivity, measured as the ratio of the mean to the standard deviation of community biomass, increases with species richness and evenness. We used experimental species mixtures of grassland plants to test this hypothesis and identified the mechanisms involved. Additionally, we tested whether biodiversity, productivity and temporal stability were similarly influenced by particular types of species interactions. We found that productivity was less variable among years in plots planted with more species. Temporal stability did not depend on whether the species were planted equally abundant (high evenness) or not (realistically low evenness). Greater richness increased temporal stability by increasing overyielding, asynchrony of species fluctuations and statistical averaging. Species interactions that favoured unproductive species increased both biodiversity and temporal stability. Species interactions that resulted in niche partitioning or facilitation increased both productivity and temporal stability. Thus, species interactions can promote biodiversity and ecosystem services.     

高寒草甸不同草地类型功能群多样性及组成对植物群落生产力的影响

对不同类型草地功能群多样性和组成与植物群落生产力之间的关系进行了探讨。结果表明：(1)在矮嵩草(Kobresia humlis)草甸和金露梅(Potentilla froticosa)灌丛中,豆科植物的作用比较明显,而其他功能群植物的作用较弱。(2)在藏嵩草(Kobresia tibetica)沼泽化草甸和小嵩草(K．pygmaca)草甸中,虽然杂类草、C3植物和莎草科植物功能群的生产力占群落初级生产力的比例较大,但二者在统计上没有显著性差异,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响,更主要的是受到功能群内物种密度和均匀度的影响,即功能群组成比功能群多样性更能说明对生态系统过程的影响。(3)不同类型草地群落植物功能群盖度与群落初级生产力呈显著的线性相关。(4)不同类型草地群落生产力与功能群内物种数的变化均表现为单峰曲线关系,即功能群内物种数处于中间水平时,群落生产力最高。     

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.     

Focusing on individual plants to understand community scale biodiversity effects: the case of root distribution in grasslands

Spatial resource partitioning between species via differences in rooting depth is one of the main explanations for the positive biodiversity–productivity relationship. However, evidence for the importance of this mechanism is limited. This may be due to the community scale at which these interactions are often investigated. Community measures represent net outcomes of species interactions and may obscure the mechanisms underlying belowground interactions. Here, we assess the performance of ~1700 individual plants and their heterospecific neighbours over three growing seasons in experimental grassland plots containing one, four or sixteen different plant species and tested whether their performance in mixtures compared to monocultures was related to their own rooting depth versus the rooting depth of their heterospecific neighbours. Overall, individuals of deep-rooting species performed better in mixtures and this effect significantly increased when surrounded by more shallow-rooting species. This effect was not apparent for the shallow rooting species. Together, including both deep and shallow rooting species increased mixture performance. Our results show that taking the perspective of the individual rather than the community can elucidate the interactions between species that contribute to positive biodiversity effects, emphasizing the need for studies at different scales to disentangle the myriad interactions that take place in diverse communities.     

Diversity–productivity relationships in two ecologically realistic rarity–extinction scenarios

To develop a better understanding of how biodiversity loss and productivity are related, we need to consider ecologically realistic rarity (i.e. reduced evenness and increased dominance) and extinction (i.e. reduced richness) scenarios. Furthermore, we need to identify and better understand the factors that influence species and community yielding behaviors because the general conditions for overyielding are the same as those for coexistence. We established experimental tallgrass prairie plots in Iowa to determine how two ecologically realistic rarity–extinction scenarios influenced aboveground net primary productivity (ANPP) and disassembly. Equal‐mass seedlings of six tallgrass prairie species were transplanted into field plots to establish realistic declining species evenness (high, medium, low) and richness (4, 1) treatments. Across declining evenness treatments, the relative abundance of the ubiquitous tall species Andropogon gerardii increased, the relative abundance of the tall species Salvia azurea was constant, and the relative abundance of two short (dissimilar height scenario) or two tall species (tall scenario) decreased. Monocultures of Andropogon represented a continuation of this trend until there was complete dominance by Andropogon and extinction of all other species. Our treatments also allowed us to test if variation in plant height contributes to the complementarity effect. Niche partitioning in plant height was not positively related to complementarity. The effects of declining species evenness and richness on the diversity–productivity relationship were different for these two ecologically realistic rarity–extinction scenarios. Specifically, as diversity declined across treatments, ANPP and the selection effects decreased in tall communities, but not in dissimilar communities. Additionally, differences between these two scenarios revealed that decreased species yielding behavior is associated with two tallgrass prairie extinction risk factors, rarity and short height. The differences between these scenarios demonstrate the importance of incorporating the known patterns of diversity declines into future studies.     

A functional trait-based approach to understand community assembly and diversity–productivity relationships over 7 years in experimental grasslands

Several multi-year biodiversity experiments have shown positive species richness–productivity relationships which strengthen over time, but the mechanisms which control productivity are not well understood. We used experimental grasslands (Jena Experiment) with mixtures containing different numbers of species (4, 8, 16 and 60) and plant functional groups (1–4; grasses, legumes, small herbs, tall herbs) to explore patterns of variation in functional trait composition as well as climatic variables as predictors for community biomass production across several years (from 2003 to 2009). Over this time span, high community mean trait values shifted from the dominance of trait values associated with fast growth to trait values suggesting a conservation of growth-related resources and successful reproduction. Increasing between-community convergence in means of several productivity-related traits indicated that environmental filtering and exclusion of competitively weaker species played a role during community assembly. A general trend for increasing functional trait diversity within and convergence among communities suggested niche differentiation through limiting similarity in the longer term and that similar mechanisms operated in communities sown with different diversity. Community biomass production was primarily explained by a few key mean traits (tall growth, large seed mass and leaf nitrogen concentration) and to a smaller extent by functional diversity in nitrogen acquisition strategies, functional richness in multiple traits and functional evenness in light-acquisition traits. Increasing species richness, presence of an exceptionally productive legume species (Onobrychis viciifolia) and climatic variables explained an additional proportion of variation in community biomass. In general, community biomass production decreased through time, but communities with higher functional richness in multiple traits had high productivities over several years. Our results suggest that assembly processes within communities with an artificially maintained species composition maximize functional diversity through niche differentiation and exclusion of weaker competitors, thereby maintaining their potential for high productivity.     

Is community persistence related to diversity? A test with prairie species in a long-term experiment

Community persistence, or the ability of a community to maintain species composition and diversity through time, is a component of stability that is important to restoration. We ran a biodiversity–ecosystem functioning experiment for three years, and then stopped weeding it for 5–6 years, which allowed us to test whether increased plant species diversity and dissimilarity in height would lead to increased community persistence in the face of high invasion pressure by non-native species. Our approach was unique in that the experiment varied richness (one or four species) and evenness (three levels plus monocultures of the dominant species) using two separate dissimilarity types (having all tall species or having tall and short species combined) in six spatiotemporal blocks. Persistence was quantified as to how well positive productivity–diversity relationships, proportion of planted native species, and species richness remained unchanged over time. Thus, high persistence values indicate low levels of invasion and local extinction. We found that the positive relationship between diversity measures and productivity persisted after cessation of weeding. The proportion of planted species was 32% higher in mixture than in monoculture plots, indicating that monocultures were more heavily invaded by non-native species. Reduced evenness did not affect persistence measures in plots with dissimilar heights, but measures declined linearly with decreased evenness in plots with all tall species. Our results suggest that (1) persistence–diversity relationships are likely to vary with the traits of species becoming rare and going extinct, and (2) it is important to restore higher species diversity in restoration projects to favor the long-term persistence of planted species.     

Mycorrhizal suppression and phosphorus addition influence the stability of plant community composition and function in a temperate steppe

Nutrient enrichment can reduce ecosystem stability, typically measured as temporal stability of a single function, e.g. plant productivity. Moreover, nutrient enrichment can alter plant–soil interactions (e.g. mycorrhizal symbiosis) that determine plant community composition and productivity. Thus, it is likely that nutrient enrichment and interactions between plants and their soil communities co-determine the stability in plant community composition and productivity. Yet our understanding as to how nutrient enrichment affects multiple facets of ecosystem stability, such as functional and compositional stability, and the role of above–belowground interactions are still lacking. We tested how mycorrhizal suppression and phosphorus (P) addition influenced multiple facets of ecosystem stability in a three-year field study in a temperate steppe. Here we focused on the functional and compositional stability of plant community; functional stability is the temporal community variance in primary productivity; compositional stability is represented by compositional resistance, turnover, species extinction and invasion. Community variance was partitioned into population variance defined as community productivity weighted average of the species temporal variance in performance, and species synchrony defined as the degree of temporal positive covariation among species. Compared to treatments with mycorrhizal suppression, the intact AM fungal communities reduced community variance in primary productivity by reducing species synchrony at high levels of P addition. Species synchrony and population variance were linearly associated with community variance with the intact AM fungal communities, while these relationships were decoupled or weakened by mycorrhizal suppression. The intact AM fungal communities promoted the compositional resistance of plant communities by reducing compositional turnover, but this effect was suppressed by P addition. P addition increased the number of species extinctions and thus promoted compositional turnover. Our study shows P addition and AM fungal communities can jointly and independently modify the various components of ecosystem stability in terms of plant community productivity and composition.     

Increasing antagonistic interactions cause bacterial communities to collapse at high diversity

Biodiversity is a major determinant of ecosystem functioning. Species-rich communities often use resources more efficiently thereby improving community performance. However, high competition within diverse communities may also reduce community functioning. We manipulated the genotypic diversity of Pseudomonas fluorescens communities, a plant mutualistic species inhibiting pathogens. We measured antagonistic interactions in vitro, and related these interactions to bacterial community productivity (root colonisation) and ecosystem service (host plant protection). Antagonistic interactions increased disproportionally with species richness. Mutual poisoning between competitors lead to a 'negative complementarity effect', causing a decrease in bacterial density by up to 98% in diverse communities and a complete loss of plant protection. The results emphasize that antagonistic interactions may determine community functioning and cause negative biodiversity-ecosystem functioning relationships. Interference competition may thus be an additional key for predicting the dynamics and performance of natural assemblages and needs to be implemented in future biodiversity models.     

Biodiversity influences ecosystem functioning in aquatic angiosperm communities

Knowledge of the connection between aquatic plant diversity and ecosystem processes is still limited. To examine how plant species diversity affects primary productivity, plant nutrient use, functional diversity of secondary producers and population/community stability, we manipulated submerged angiosperm species diversity in a field experiment lasting 15 weeks. Plant richness increased the shoot density for three of four species. Polyculture biomass production was enhanced by increasing richness, with positive complementarity and selection effects causing positive biodiversity effects. Species richness enhanced the community stability for biomass production and shoot density. Sediment ammonium availability decreased with plant diversity, suggesting improved nutrient usage with increasing plant richness. Interestingly, positive multitrophic effects of plant species richness on structural and functional diversity of macrobenthic secondary producers were recorded. The results suggest that mixed seagrass meadows play an important role for ecosystem functioning and thus contribute to the provision of goods and services in coastal areas.     

氮添加对宁夏荒漠草原植物初级生产力的影响机制

许多研究探索了与全球变化相关的生态系统功能的变化,但对生态系统功能变化的机制与途径了解较少。初级生产力是生态系统功能的重要组分,但关于氮(N)添加下荒漠草原植物群落初级生产力如何变化以及变化机制尚未明确,N是否通过影响生物多样性来影响荒漠草原初级生产力?为此,本研究在荒漠草原开展了为期4年的N添加控制实验(2018—2021年),试验处理包括对照和4个N添加水平(5、10、20和40 g m^-2 a^-1),研究了N添加对荒漠草原物种多样性、功能多样性、初级生产力及其关系的影响。结果表明：(1)N添加处理(2018—2021年)改变了植物物种多样性及功能多样性,但年际间变化趋势不同。N添加处理第四年(2021年)荒漠草原植物功能多样性(Rao指数)、群落加权平均值-株高、功能均匀度和功能离散度均显著增加,而荒漠草原植物物种丰富度和Shannon-Wiener指数均显著降低。(2)N添加可以通过影响物种丰富度和功能多样性进而间接地促进荒漠草原初级生产力,但群落加权性状值-株高对初级生产力的影响是正效应,而物种丰富度和功能离散度对初级生产力的影响是...     

Species evenness and invasion resistance of experimental grassland communities

Concern for biodiversity loss coupled with the accelerated rate of biological invasions has provoked much interest in assessing how native plant species diversity affects invasibility. Although experimental studies extensively document the effects of species richness on invader performance, the role of species evenness in such studies is rarely examined. Species evenness warrants more attention because the relative abundances of species can account for substantially more of the variance in plant community diversity and tend to change more rapidly and more frequently in response to disturbances than the absolute numbers of species. In this study, we experimentally manipulated species evenness within native prairie grassland mesocosms. We assessed how evenness affected primary productivity, light availability and the resistance of native communities to invasion. The primary productivity of native communities increased significantly with species evenness, and this increase in productivity was accompanied by significant decreases in light availability. However, evenness had no effect on native community resistance to invasion by three common exotic invasive species. In this study, niche complementarity provides a potential mechanism for the effects of evenness on productivity and light availability, but these effects apparently were not strong enough to alter the invasibility of the experimental communities. Our results suggest that species evenness enhances community productivity but provides no benefit to invasion resistance in otherwise functionally diverse communities.     

Across species‐pool aggregation alters grassland productivity and diversity

Plant performance is determined by the balance of intra‐ and interspecific neighbors within an individual's zone of influence. If individuals interact over smaller scales than the scales at which communities are measured, then altering neighborhood interactions may fundamentally affect community responses. These interactions can be altered by changing the number (species richness), abundances (species evenness), and positions (species pattern) of the resident plant species, and we aimed to test whether aggregating species at planting would alter effects of species richness and evenness on biomass production at a common scale of observation in grasslands. We varied plant species richness (2, 4, or 8 species and monocultures), evenness (0.64, 0.8, or 1.0), and pattern (planted randomly or aggregated in groups of four individuals) within 1 × 1 m plots established with transplants from a pool of 16 tallgrass prairie species and assessed plot‐scale biomass production and diversity over the first three growing seasons. As expected, more species‐rich plots produced more biomass by the end of the third growing season, an effect associated with a shift from selection to complementarity effects over time. Aggregating conspecifics at a 0.25‐m scale marginally reduced biomass production across all treatments and increased diversity in the most even plots, but did not alter biodiversity effects or richness–productivity relationships. Results support the hypothesis that fine‐scale species aggregation affects diversity by promoting species coexistence in this system. However, results indicate that inherent changes in species neighborhood relationships along grassland diversity gradients may only minimally affect community (meter) – scale responses among similarly designed biodiversity–ecosystem function studies. Given that species varied in their responses to local aggregation, it may be possible to use such species‐specific results to spatially design larger‐scale grassland communities to achieve desired diversity and productivity responses.     

A role for the sampling effect in invaded ecosystems

Species loss and invasion of exotic species are two components of global biodiversity change that are expected to influence ecosystem functioning. Yet how they interact in natural settings remains unclear. Experiments have revealed two major mechanisms for the observed increase in primary productivity with plant species richness. Plant productivity may rise with species richness due to the increased amount of resources used by more diverse communities (niche complementarity) or through the increased probability of including a highly productive, dominant species in the community (sampling effect). Current evidence suggests that niche complementarity is the most relevant mechanism, whereas the sampling effect would only play a minor and transient role in natural systems. In turn, exotic species can invade by using untapped resources or because they possess a fitness advantage over resident species allowing them to dominate the community. We argue that the sampling effect can be a significant biodiversity mechanism in ecosystems invaded by dominant exotic species, and that the effect can be persistent even after decades of succession. We illustrate this idea by analyzing tree species richness–productivity relationships in a subtropical montane forest (NW Argentina) heavily invaded by Ligustrum lucidum, an evergreen tree from Asia. We found that the forest biomass increased along a natural gradient of tree species richness whether invaded by L. lucidum or not. Consistent with the sampling effect, L. lucidum invasion tripled total tree biomass irrespective of species richness, and monocultures of L. lucidum were more productive than any of the most species‐rich, uninvaded communities. Hence, the sampling effect may not be restricted to randomly assembled, synthetic communities. We emphasize that studying invaded ecosystems may provide novel insights on the mechanisms underlying the effect of biodiversity on ecosystem function.     

Determining the mechanism by which fish diversity influences production

Understanding the ability of biodiversity to govern ecosystem function is essential with current pressures on natural communities from species invasions and extirpations. Changes in fish communities can be a major determinant of food web dynamics, and even small shifts in species composition or richness can translate into large effects on ecosystems. In addition, there is a large information gap in extrapolating results of small-scale biodiversity–ecosystem function experiments to natural systems with realistic environmental complexity. Thus, we tested the key mechanisms (resource complementarity and selection effect) for biodiversity to influence fish production in mesocosms and ponds. Fish diversity treatments were created by replicating species richness and species composition within each richness level. In mesocosms, increasing richness had a positive effect on fish biomass with an overyielding pattern indicating species mixtures were more productive than any individual species. Additive partitioning confirmed a positive net effect of biodiversity driven by a complementarity effect. Productivity was less affected by species diversity when species were more similar. Thus, the primary mechanism driving fish production in the mesocosms was resource complementarity. In the ponds, the mechanism driving fish production changed through time. The key mechanism was initially resource complementarity until production was influenced by the selection effect. Varying strength of intraspecific interactions resulting from differences in resource levels and heterogeneity likely caused differences in mechanisms between the mesocosm and pond experiments, as well as changes through time in the ponds. Understanding the mechanisms by which fish diversity governs ecosystem function and how environmental complexity and resource levels alter these relationships can be used to improve predictions for natural systems.     

Genotypic richness and dissimilarity opposingly affect ecosystem functioning

Biodiversity is an essential determinant of ecosystem functioning. Numerous studies described positive effects of diversity on the functioning of communities arising from complementary resource use and facilitation. However, high biodiversity may also increase competitive interactions, fostering antagonism and negatively affecting community performance. Using experimental bacterial communities we differentiated diversity effects based on genotypic richness and dissimilarity. We show that these diversity characteristics have opposite effects on ecosystem functioning. Genotypic dissimilarity governed complementary resource use, improving ecosystem functioning in complex resource environments. Contrastingly, genotypic richness drove allelopathic interactions, mostly reducing ecosystem functioning. The net biodiversity effect on community performance resulted from the interplay between the genetic structure of the community and resource complexity. These results demonstrate that increasing richness, without concomitantly increasing dissimilarity, can decrease ecosystem functioning in simple environments due to antagonistic interactions, an effect insufficiently considered so far in mechanistic models of the biodiversity-ecosystem functioning relationship.     

Compensation: an alternative method for analyzing diversity-productivity experiments

Although recent experimental results demonstrate a positive effect of diversity on primary productivity, the interpretation of these experiments has been controversial, creating a need for new methods of analysis. The methods developed in response to this need all use the production of individual species grown in monocultures to calculate the expected production of each species mixture, then analyze departures from these expectations as a function of species richness. We propose an alternative method that treats the same assembly experiments as species removals, and calculates the expected production of each mixture based on the production of individual species when grown together in the full community (the experimental mixture containing all species in the pool). Using the observed production of the full community, and the observed and expected productions of less diverse mixtures, we calculate an index of compensation that measures the degree of functional recovery following species loss. To explore whether losses of dominant versus subordinate species have different ecosystem effects, we suggest a multiple regression approach that tests the influence of both species richness and expected production on compensation. If compensation varies with species richness or expected production consistently in many experimental systems, then we may be able to predict the ecosystem effect of different types of extinctions.
While existing monoculture approaches more directly test hypotheses about complementary resource use, the compensation approach offers two advantages: 1) it is more appropriate for testing how extinctions will affect ecosystem function, and 2) it may provide an important link between assembly experiments in artificial communities and removal experiments in natural systems.