Positive Biodiversity Productivity Relationships Research Articles

Eutrophication weakens the positive biodiversity–productivity relationship of benthic diatoms in plateau lakes

Context Freshwater primary productivity is threatened by the decline in biodiversity associated with nutrient enrichment, but there is still uncertainty about how the biodiversity and productivity relationship (BPR) varies with the trophic states. Aims We aimed to examine the variation of benthic diatom BPRs in three plateau lakes with different trophic states and their underlying driving mechanisms. Methods We examined the relationship between diatom taxonomic and functional diversity, niche width, and niche overlap with productivity. Key results The taxonomic and functional diversity, niche width and productivity of benthic diatoms were highest in the mesotrophic lake. The benthic diatom BPRs were linear and positive, with the slope of BPRs being the lowest in eutrophic lake. Motile, non-attached and small-sized diatoms were dominant in eutrophic lake. Nutrient concentrations indirectly affected primary productivity by influencing algal community structure, niche width and biodiversity change. Conclusions Diatom productivity and diversity showed a positive relationship, but nutrient enrichment weakened this relationship. By combining taxonomic and functional diversity indices, supplemented by niche analysis, we can further understand the variation of diatom productivity. Implications The results provide a basis for predicting the changes in BPRs of benthic diatoms in the littoral zone with different trophic states.

Cross-domain diversity effects: linking diatom species richness, intraspecific richness, and biomass production to host-associated bacterial diversity.

Interactions between bacteria and microalgae are important for the functioning of aquatic ecosystems, yet interactions based on the biodiversity of these two taxonomic domains have been scarcely studied. Specifically, it is unclear whether a positive biodiversity-productivity relationship in phytoplankton is largely facilitated by niche partitioning among the phytoplankton organisms themselves or whether associated bacterial communities play an additional role in modifying these diversity effects. Moreover, the effects of intraspecific diversity in phytoplankton communities on bacterial community diversity have not been tested. To address these points, we factorially manipulated both species and intraspecific richness of three diatoms to test the effects of diatom species/strain diversity on biomass production and bacterial diversity in algae-bacteria communities. The results show that diatom intraspecific diversity has significant positive effects on culture biomass and the diversity of the associated free-living bacterial community (0.2-3μm size fraction), which are comparable in magnitude to species diversity effects. However, there were little to no effects of diatom diversity on host-associated bacterial diversity (>3μm size fraction), or of bacterial diversity on biomass production. These results suggest a decoupling of bacterial diversity from the diatom diversity-productivity relationship and provide early insights regarding the relations between diversity across domains in aquatic ecosystems.

Tree diversity increases productivity through enhancing structural complexity across mycorrhizal types.

Tree species diversity and mycorrhizal associations play a central role for forest productivity, but factors driving positive biodiversity-productivity relationships remain poorly understood. In a biodiversity experiment manipulating tree diversity and mycorrhizal associations, we examined the roles of above- and belowground processes in modulating wood productivity in young temperate tree communities and potential underlying mechanisms. We found that tree species richness, but not mycorrhizal associations, increased forest productivity by enhancing aboveground structural complexity within communities. Structurally complex communities were almost twice as productive as structurally simple stands, particularly when light interception was high. We further demonstrate that overyielding was largely explained by positive net biodiversity effects on structural complexity with functional variation in shade tolerance and taxonomic diversity being key drivers of structural complexity in mixtures. Consideration of stand structural complexity appears to be a crucial element in predicting carbon sequestration in the early successional stages of mixed-species forests.

A critical assessment of the biodiversity–productivity relationship in forests and implications for conservation

The question of whether biodiversity conservation and carbon conservation can be synergistic hinges on the form of the biodiversity-productivity relationship (BPR), a fundamental ecological pattern. The stakes are particularly high when it comes to forests, which at a global level comprises a large fraction of both biodiversity and carbon. And yet, in forests, the BPR is relatively poorly understood. In this review, we critically evaluate research on forest BPRs, focussing on the experimental and observational studies of the last 2 decades. We find general support for a positive forest BPR, suggesting that biodiversity and carbon conservation are synergistic to a degree. However, we identify several major caveats: (i) although, on average, productivity may increase with biodiversity, the highest-yielding forests are often monocultures of very productive species; (ii) productivity typically saturates at fewer than ten species; (iii) positive BPRs can be driven by some third variable, in particular stem density, instead of a causal arrow from biodiversity to productivity; (iv) the BPR's sign and magnitude varies across spatial grains and extents, and it may be weak at scales relevant to conservation; and (v) most productivity estimates in forests are associated with large errors. We conclude by explaining the importance of these caveats for both conservation programmes focussed on protection of existing forests and conservation programmes focussed on restoring or replanting forests.

Rapid transgenerational adaptation in response to intercropping reduces competition.

By capitalising on positive biodiversity-productivity relationships, intercropping provides opportunities to improve agricultural sustainability. Intercropping is generally implemented using commercial seeds that were bred for maximal productivity in monocultures, thereby ignoring the ability of plants to adapt over generations to the surrounding neighbourhood, notably through increased complementarity, that is reduced competition or increased facilitation. This is why using monoculture-adapted seeds for intercropping might limit the benefits of crop diversity on yield. However, the adaptation potential of crops and the corresponding changes in complementarity have not been explored in annual crop systems. Here we show that plant-plant interactions among annual crops shifted towards reduced competition and/or increased facilitation when the plants were growing in the same community type as their parents did in the previous two generations. Total yield did not respond to this common coexistence history, but in fertilized conditions, we observed increased overyielding in mixtures with a common coexistence history. Surprisingly, we observed character convergence between species sharing the same coexistence history for two generations, in monocultures but also in mixtures: the six crop species tested converged towards taller phenotypes with lower leaf dry matter content. This study provides the first empirical evidence for the potential of parental diversity affecting plant-plant interactions, species complementarity and therefore potentially ecosystem functioning of the following generations in annual cropping systems. Although further studies are required to assess the context-dependence of these results, our findings may still have important implications for diversified agriculture as they illustrate the potential of targeted cultivars to increase complementarity of species in intercropping, which could be achieved through specific breeding for mixtures.

Using spatially-explicit plant competition models to optimise crop productivity in intercropped systems

Intercropping, by capitalizing on positive biodiversity–productivity relationships, represents a promising option to increase agricultural sustainability. However, the complexity and context-dependency of plant–plant interactions can make it challenging for farmers to find suitable crop combinations. Furthermore, intercropping is usually implemented with standard inter-row spacing and plant densities based on monoculture practices, which might not be the ideal configuration to maximize yield.Here we present a spatially-explicit yield analysis method based on plant ecological interaction models that allowed to optimize crop species combinations and spatial configurations for maximal yield in intercropped systems. We tested this method with three crop species, namely oat, lupine, and camelina. In a first step, field experiments in which crop density and adjacent crop type were varied provided us with indications on which species would compete more with each other. The results showed us that oat and camelina strongly competed with each other. In addition, the distance experiments allowed us to understand how the changes in yield associated with the presence of neighbors vary with distance. This allowed us to find the sets of parameters (identity of neighbors, sowing density, distances between individuals) that optimise intercrop yield (measured as Land Equivalent Ratio [LER]) for the three considered species. Specifically, we show that alternating rows of species led to higher LERs than a homogeneous species mixing, and that 3-species combinations are not necessarily more performant than the best 2-species combinations. In addition, we show that increasing the density of oat is generally beneficial for LER, while increasing the density of lupine is not.By modelling crop yield from simple and reproducible density and distance experiments, our results allow to optimize crop mixtures in terms of species combinations and spatial configurations, for maximal crop yield.

The Complex Biodiversity-Ecosystem Function Relationships for the Qinghai-Tibetan Grassland Community.

Despite the long history of the study of the biodiversity-ecosystem function relationship, uncertainty remains about the relationship of natural grassland ecosystems under stressful conditions. Recently, trait- and phylogenetic-based tests provide a powerful way to detect the relationship in different spaces but have seldom been applied to stressful zones on a large spatial scale. We selected Qinghai-Tibetan as the study area and collected a grassland community database involving 581 communities. We calculated biomass and species’, functional, and phylogenetic diversity of each community and examined their relationships by using linear and non-linear regression models. Results showed an overall positive biodiversity-productivity relationship in species’, functional and phylogenetic space. The relationship, however, was non-linear, in which biodiversity explained better the variation in community biomass when species diversity was more than a threshold, showing a weak effect of biodiversity on ecosystem function in low species diversity communities. We also found a filled triangle for the limit of the relationship between species and functional diversity, implying that functional diversity differs significantly among communities when their species diversity is low but finally converges to be a constant with increasing communities’ species diversity. Our study suggests that multiple niche processes may structure the grassland communities, and their forces tend to balance in high-biodiversity communities.

Using plant traits to understand the contribution of biodiversity effects to annual crop community productivity.

Increasing biodiversity generally enhances productivity through selection and complementarity effects not only in natural, but also in agricultural, systems. However, the quest to explain why diverse cropping systems are more productive than monocultures remains a central goal in agricultural science. In a mesocosm experiment, we constructed monocultures, two‐ and four‐species mixtures from eight crop species with or without fertilizer and both in temperate Switzerland and dry, Mediterranean Spain. We measured physical factors and plant traits and related these in structural equation models to selection and complementarity effects to explain seed yield differences between monocultures and mixtures. Increased crop diversity increased seed yield in Switzerland. This positive biodiversity effect was driven to almost the same extent by selection and complementarity effects, which increased with plant height and specific leaf area (SLA), respectively. Also, ecological processes driving seed yield increases from monocultures to mixtures differed from those responsible for seed yield increases through the diversification of mixtures from two to four species. Whereas selection effects were mainly driven by one species, complementarity effects were linked to larger leaf area per unit leaf weight. Seed yield increases due to mixture diversification were driven only by complementarity effects and were not mediated through the measured traits, suggesting that ecological processes beyond those measured in this study were responsible for positive diversity effects on yield beyond two‐species mixtures. By understanding the drivers of positive biodiversity–productivity relationships, we can improve our ability to predict species combinations that enhance ecosystem functioning and can promote sustainable agricultural production.

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.

Effects of local neighbourhood diversity on crown structure and productivity of individual trees in mature mixed-species forests

BackgroundSpecies-specific genotypic features, local neighbourhood interactions and resource supply strongly influence the tree stature and growth rate. In mixed-species forests, diversity-mediated biomass allocation has been suggested to be a fundamental mechanism underlying the positive biodiversity-productivity relationships. Empirical evidence, however, is rare about the impact of local neighbourhood diversity on tree characteristics analysed at a very high level of detail. To address this issue we analysed these effects on the individual-tree crown architecture and tree productivity in a mature mixed forest in northern Germany.MethodsOur analysis considers multiple target tree species across a local neighbourhood species richness gradient ranging from 1 to 4. We applied terrestrial laser scanning to quantify a large number of individual mature trees (N = 920) at very high accuracy. We evaluated two different neighbour inclusion approaches by analysing both a fixed radius selection procedure and a selection based on overlapping crowns.Results and conclusionsWe show that local neighbourhood species diversity significantly increases crown dimension and wood volume of target trees. Moreover, we found a size-dependency of diversity effects on tree productivity (basal area and wood volume increment) with positive effects for large-sized trees (diameter at breast height (DBH) > 40 cm) and negative effects for small-sized (DBH < 40 cm) trees. In our analysis, the neighbour inclusion approach has a significant impact on the outcome. For scientific studies and the validation of growth models we recommend a neighbour selection by overlapping crowns, because this seems to be the relevant scale at which local neighbourhood interactions occur. Because local neighbourhood diversity promotes individual-tree productivity in mature European mixed-species forests, we conclude that a small-scale species mixture should be considered in management plans.

Plant water uptake along a diversity gradient provides evidence for complementarity in hydrological niches

Biodiversity enhances a variety of ecosystem processes, and yet the underlying mechanisms through which these relationships occur remain a critical knowledge gap. Here, we used the natural abundance of stable isotopes to measure depth of water uptake in five common grassland species (Asclepias tuberosa, Lespedeza capitata, Liatris aspera, Schizachyrium scoparium and Sorghastrum nutans) growing across an experimental grassland diversity gradient. Using this approach, we addressed the following questions: 1) does the depth‐specific provenance of water uptake differ among species and/or do interspecific differences in water source manifest with increasing community diversity? 2) Does the isotopic niche space occupied by plants change with increasing diversity? 3) Is plasticity in water uptake depth across a diversity gradient associated with functional plant responses? We found that the depth of soil water used by plants was inherently different among species when grown in monocultures. All species used less shallow soil water and more intermediate‐depth soil water in mixed assemblages than in monocultures, resulting in similar interspecific differences in water source across the diversity gradient. However, plasticity in the locations of water used were positively associated with increases in plant growth in higher diversity treatments. These results indicate that plasticity in water‐use may contribute to positive biodiversity–productivity relationships commonly observed in temperate grasslands.

Effect of forest structure on stand productivity in Central European forests depends on developmental stage and tree species diversity

Recently, many studies have found positive biodiversity–productivity relationships in forests. In contrast, different types of correlations have been identified in the analyses of tree diversity–structure–productivity relationships. We suspect that these conflicting conclusions might result from the different developmental stages of the investigated forest stands. We therefore analyzed the development of tree diversity–structure–productivity relationships at the stand level and individual tree level in 192 long-term experimental plots in Central Europe. As a measure of stand productivity, we analyzed stand volume growth (m3 ha−1 year−1). Tree species diversity was quantified by the Shannon index and structural heterogeneity was represented by the Gini coefficient of basal area. For a more detailed analysis at the tree level using a smaller portion of the dataset, the tree position–dependent indices, diameter differentiation index, and aggregation index were used. Whether the effect of structural heterogeneity on stand productivity was positive or negative depended on the stand development stage. In early developmental stages, high structural heterogeneity lowered productivity. In later developmental stages, however, stand structural heterogeneity had a positive effect on productivity. Our study might provide insights regarding the mechanisms underlying the contradictory findings obtained in recent studies dealing with tree diversity–structure–productivity relationships. This knowledge is vital for the adaptation of forest management to meet future demands on forest ecosystems.

Positive biodiversity–productivity relationships in forests: climate matters

While it is widely acknowledged that forest biodiversity contributes to climate change mitigation through improved carbon sequestration, conversely how climate affects tree species diversity-forest productivity relationships is still poorly understood. We combined the results of long-term experiments where forest mixtures and corresponding monocultures were compared on the same site to estimate the yield of mixed-species stands at a global scale, and its response to climatic factors. We found positive mixture effects on productivity using a meta-analysis of 126 case studies established at 60 sites spread across five continents. Overall, the productivity of mixed-species forests was 15% greater than the average of their component monocultures, and not statistically lower than the productivity of the best component monoculture. Productivity gains in mixed-species stands were not affected by tree age or stand species composition but significantly increased with local precipitation. The results should guide better use of tree species combinations in managed forests and suggest that increased drought severity under climate change might reduce the atmospheric carbon sequestration capacity of natural forests.

Positive biodiversity-productivity relationship predominant in global forests.

The biodiversity-productivity relationship (BPR) is foundational to our understanding of the global extinction crisis and its impacts on ecosystem functioning. Understanding BPR is critical for the accurate valuation and effective conservation of biodiversity. Using ground-sourced data from 777,126 permanent plots, spanning 44 countries and most terrestrial biomes, we reveal a globally consistent positive concave-down BPR, showing that continued biodiversity loss would result in an accelerating decline in forest productivity worldwide. The value of biodiversity in maintaining commercial forest productivity alone-US$166 billion to 490 billion per year according to our estimation-is more than twice what it would cost to implement effective global conservation. This highlights the need for a worldwide reassessment of biodiversity values, forest management strategies, and conservation priorities.

Tree species partition N uptake by soil depth in boreal forests.

It is recognized that the coexistence of herbaceous species in N-depleted habitats can be facilitated by N partitioning; however, the existence of such a phenomenon for trees has not yet been demonstrated. Here, we show from both foliage and soil 15N natural abundance values and from a 12-year in situ 15N addition experiment, that black spruce (Picea mariana) and jack pine (Pinus banksiana), two widespread species of the Canadian boreal forest, take up N at different depths. While black spruce takes up N from the organic soil, jack pine acquires it deeper within the highly N-depleted mineral soil. Systematic difference in foliar 15N natural abundance between the two species across seven sites distributed throughout the eastern Canadian boreal forest shows that N spatial partitioning is a widespread phenomenon. Distinct relationships between delta15N and N concentration in leaves of both species further emphasize their difference in N acquisition strategies. This result suggests that such complementary mechanisms of N acquisition could facilitate tree species coexistence in such N-depleted habitats and could contribute to the positive biodiversity-productivity relationship recently revealed for the eastern Canadian boreal forest, where jack pine is present. It also has implications for forest management and provides new insights to interpret boreal forest regeneration following natural or anthropogenic perturbations.

Evidence from the real world: 15N natural abundances reveal enhanced nitrogen use at high plant diversity in Central European grasslands

Summary Complementarity that leads to more efficient resource use is presumed to be a key mechanism explaining positive biodiversity–productivity relationships but has been described solely for experimental set‐ups with controlled environmental settings or for very short gradients of abiotic conditions, land‐use intensity and biodiversity. Therefore, we analysed plant diversity effects on nitrogen dynamics across a broad range of Central European grasslands. The 15N natural abundance in soil and plant biomass reflects the net effect of processes affecting ecosystem N dynamics. This includes the mechanism of complementary resource utilization that causes a decrease in the 15N isotopic signal. We measured plant species richness, natural abundance of 15N in soil and plants, above‐ground biomass of the community and three single species (an herb, grass and legume) and a variety of additional environmental variables in 150 grassland plots in three regions of Germany. To explore the drivers of the nitrogen dynamics, we performed several analyses of covariance treating the 15N isotopic signals as a function of plant diversity and a large set of covariates. Increasing plant diversity was consistently linked to decreased δ15N isotopic signals in soil, above‐ground community biomass and the three single species. Even after accounting for multiple covariates, plant diversity remained the strongest predictor of δ15N isotopic signals suggesting that higher plant diversity leads to a more closed nitrogen cycle due to more efficient nitrogen use. Factors linked to increased δ15N values included the amount of nitrogen taken up, soil moisture and land‐use intensity (particularly fertilization), all indicators of the openness of the nitrogen cycle due to enhanced N‐turnover and subsequent losses. Study region was significantly related to the δ15N isotopic signals indicating that regional peculiarities such as former intensive land use could strongly affect nitrogen dynamics. Synthesis. Our results provide strong evidence that the mechanism of complementary resource utilization operates in real‐world grasslands where multiple external factors affect nitrogen dynamics. Although single species may differ in effect size, actively increasing total plant diversity in grasslands could be an option to more effectively use nitrogen resources and to reduce the negative environmental impacts of nitrogen losses.

Cadmium pollution triggers a positive biodiversity–productivity relationship: evidence from a laboratory microcosm experiment

Summary 1. Human activities are greatly changing the conditions of ecosystems. One of these major changes is that more and more ecosystems are facing increasing concentrations of highly toxic heavy metals like cadmium (Cd), lead (Pb) and mercury (Hg). Previous studies have shown this type of pollution may be associated with the loss of biodiversity, leaving some pollution‐tolerant species to dominate the polluted ecosystems. To date, however, little is known about the extent to which the pollution caused by these toxins may affect the relationship between biodiversity and ecosystem function. 2. In this study, we aimed to assess the role of Cd pollution in determining biodiversity–productivity relationships in aquatic ecosystems, by using data from a laboratory microcosm experiment that tested 110 algal communities with three levels of Cd pollution. 3. Our results showed that the productivities of algal communities growing in non‐Cd‐polluted media did not significantly increase with increasing species richness, and this pattern did not change over time. In contrast, significantly positive biodiversity–productivity relationships were consistently found in algal communities growing in Cd‐polluted media. 4. We revealed that the putative facilitation among algal species in response to Cd pollution was the main driver of the observed significantly positive biodiversity–productivity relationships. Most importantly, we found that not only Cd‐tolerant algal species but also Cd‐sensitive algal species were able to dominate Cd‐polluted polycultures and contribute to the increased productivities of these polycultures. 5. Synthesis and applications. Collectively, our results provide the first explicit experimental evidence that Cd pollution can trigger a positive biodiversity–productivity relationship. We identify two profound implications: (i) conservation of biodiversity in all environments may reduce the future impacts of increasing environmental stresses (like Cd pollution) on ecosystem function (such as primary productivity); (ii) one possible approach to maintain or improve algal primary productivity in a polluted aquatic ecosystem may be to construct some suitable diverse algal assemblages that include not only pollution‐tolerant species but also pollution‐sensitive ones.

Positive biodiversity–productivity relationship due to increased plant density

1. Positive effects of biodiversity on plant productivity may result from diversity-induced changes in the size or density of individual plants, yet these two possibilities have never been tested at the same time in a biodiversity experiment with a large species pool. Here, we distinguish between size effects and density effects on plant productivity, using data from 198 experimental grassland communities that contained 1–16 species. Plant modules such as tillers or rosettes were defined as relevant units, being equivalent to plant individuals in the majority of species. 2. In agreement with previous studies, we found positive effects of species richness on above-ground productivity. We show that this positive biodiversity effect resulted from diversity-induced increases in module density rather than from increases in module size. In contrast, variation in productivity within diversity levels was related to module size rather than module density. 3. The size–density relationships varied among plant functional groups and among species but their average response to increasing species richness paralleled the pattern observed at the level of the entire plant communities: species richness had a positive effect on above-ground species biomass and species module density but not on species module size. Twenty-four out of 26 overyielding species had denser populations and 25 out of 28 underyielding species had smaller modules in mixtures than in monocultures. 4. Synthesis: In grasslands, an increase in community productivity must involve an increase in plant size or density. We found that diversity-induced increases in productivity were related to diversity-induced increases in density, whereas diversity-independent increases in productivity were related to increases in plant size. Our results suggest that increased density of overyielding species in mixtures was the main driver of the positive biodiversity–productivity relationship in our experiment. We conclude that the mechanisms leading to enhanced productivity of species-rich as compared with species-poor communities cannot be derived from mechanisms explaining high productivity within communities that contain a particular number of species.

Belowground nitrogen partitioning in experimental grassland plant communities of varying species richness

Partitioning of soil nitrogen (N) by niche separation among species may be an important mechanism explaining species coexistence and positive biodiversity-productivity relationships in terrestrial plant communities. However, there is little experimental evidence for such partitioning, in particular, as assessed across a gradient of species richness. In experimental communities of one, three, and six temperate grassland species in the field, we tested whether increasing species richness (1) decreases niche breadths of individual species, (2) decreases niche overlap among species, and (3) increases niche breadth of whole communities. Six N sources consisting of three different chemical forms of 15N-labeled N (15NO3(-), 15NH4+, 13C2-15N-glycine) injected at two soil depths (3 and 12 cm) were applied to each community. The chemical form and the soil depth of N characterize the niches for which niche breadth (Levins' B) and overlap (proportional similarity) were measured. After 48 hours, aboveground plant material was harvested to measure 15N enrichment. As expected, niche breadth of single species and niche overlap among species decreased with increased species richness, but community niche breadth did not increase. The decrease in niche breadth and niche overlap mostly occurred among subordinate species or pairs of subordinate and dominant species, rather than among dominant species. Species in the six-species mixtures mostly preferred NO3(-) from shallow soil. This may be partly explained by the presence of legumes in all six-species mixtures which allowed "N sparing" (i.e., increased availability of soil N since legumes rely more on atmospheric N2 than on soil N). Niche separation with respect to N uptake from different chemical forms and soil depths did not contribute much to facilitating the coexistence of dominant species, nor do our results suggest it as a major driver of positive diversity-ecosystem functioning relationships. However, partitioning of N may be important for the persistence of subordinate species.