Effect of phytoplankton size diversity on primary productivity in the North Pacific: trait distributions under environmental variability

While most biodiversity and ecosystem functioning (BEF) studies have found positive effects of species richness on productivity, it remain unclear whether similar patterns hold for marine phytoplankton with high local richness. We use the continuous trait‐based modelling approach, which assumes infinite richness and represents diversity in terms of the variance of the size distribution, to investigate the effects of phytoplankton size diversity on productivity in a three‐dimensional ocean circulation model driven by realistic physics forcing. We find a slightly negative effect of size diversity on primary production, which we attribute to several factors including functional trait‐environment interactions, flexible stoichiometry and the saturation of productivity at low diversity levels. The benefits of trait optimisation, whereby narrow size distributions enhance productivity under relatively stable conditions, tend to dominate over those of adaptive capacity, whereby greater diversity enhances the ability of the community to respond to environmental variability.     

Biodiversity effects on ecosystem functioning respond unimodally to environmental stress

Understanding how biodiversity (B) affects ecosystem functioning (EF) is essential for assessing the consequences of ongoing biodiversity changes. An increasing number of studies, however, show that environmental conditions affect the shape of BEF relationships. Here, we first use a game‐theoretic community model to reveal that a unimodal response of the BEF slope can be expected along environmental stress gradients, but also how the ecological mechanisms underlying this response may vary depending on how stress affects species interactions. Next, we analysed a global dataset of 44 experiments that crossed biodiversity with environmental conditions. Confirming our main model prediction, the effect of biodiversity on ecosystem functioning tends to be greater at intermediate levels of environmental stress, but varies among studies corresponding to differences in stress‐effects on species interactions. Together, these results suggest that increases in stress from ongoing global environmental changes may amplify the consequences of biodiversity changes.     

浙江定海次生林内物种丰富度与生物量和生产力关系的环境依赖性

迄今生物多样性与生态系统功能关系的研究主要在物种组成随机配置的人工生态系统中进行, 在自然生态系统中研究较少, 且未考虑环境因子如何影响生态系统功能及其与生物多样性的关系。本研究选取亚热带广泛分布的次生林为研究对象, 利用模型拟合的方法, 探讨亚热带次生林中物种丰富度与生物量和生产力之间的关系, 以及环境因子(海拔、坡度、坡向、土层厚度)和次生林恢复时间(林龄)对生物量、生产力、物种丰富度与生物量和生产力间关系的影响。结果表明, 当不考虑环境因子时, 物种丰富度与生物量之间存在显著的线性正相关关系, 而与生产力之间存在显著的二次关系(先增加后减少的驼峰型)。当考虑环境因子时, 个体密度和土层厚度对生物量具有显著影响, 而环境因子对生产力并无显著效应。在坡度较陡、坡向朝南及土层较厚的环境条件下, 物种丰富度与生物量具有显著的线性正相关关系; 而在坡度较缓、坡向朝北及土层较薄的环境条件下, 物种丰富度不影响生物量。在较高海拔环境条件下, 生产力随物种丰富度先增加后减少(驼峰形状), 而在其他环境条件下, 生产力均不响应物种丰富度。以上结果说明自然森林生态系统中物种丰富度与生物量和生产力的关系存在差异, 且其相互间的关系依赖于环境因子。     

Functional richness outperforms taxonomic richness in predicting ecosystem functioning in natural phytoplankton communities

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Integrating community assembly and biodiversity to better understand ecosystem function: the Community Assembly and the Functioning of Ecosystems (CAFE) approach

The research of a generation of ecologists was catalysed by the recognition that the number and identity of species in communities influences the functioning of ecosystems. The relationship between biodiversity and ecosystem functioning (BEF) is most often examined by controlling species richness and randomising community composition. In natural systems, biodiversity changes are often part of a bigger community assembly dynamic. Therefore, focusing on community assembly and the functioning of ecosystems (CAFE), by integrating both species richness and composition through species gains, losses and changes in abundance, will better reveal how community changes affect ecosystem function. We synthesise the BEF and CAFE perspectives using an ecological application of the Price equation, which partitions the contributions of richness and composition to function. Using empirical examples, we show how the CAFE approach reveals important contributions of composition to function. These examples show how changes in species richness and composition driven by environmental perturbations can work in concert or antagonistically to influence ecosystem function. Considering how communities change in an integrative fashion, rather than focusing on one axis of community structure at a time, will improve our ability to anticipate and predict changes in ecosystem function.     

Environmental stress alters native-nonnative relationships at the community scale

The invasion of natural habitats by nonnative species is affected by both native biodiversity and environmental conditions; however few tests of facilitation between native community members and nonnative species have been conducted along disturbance and stress gradients. There is strong evidence for an increase in facilitation between native plant species with increasing levels of natural environmental stress, however it is unknown whether these same positive interactions occur between nonnative invaders and native communities. I investigated the effects of natural stress on community interactions between native heathland species and nonnative species with two field studies conducted at the landscape and community scale. At the landscape scale of investigation, nonnative species richness was positively related to native species richness. At the community level, nonnative invaders experienced facilitation with natives in the most stressful zones, whereas they experienced competition with native plants in the less stressful zones of the heathlands. Due to the observational nature of the landscape scale data, it is unclear whether nonnative diversity levels are responding positively to extrinsic factors or to native biodiversity. The experimental component of this research suggests that native community members may ameliorate stressful environmental conditions and facilitate invasion into high stress areas. I present a conceptual model which is a modification of the Shea and Chesson diversity-invasibility model and includes both facilitation as well as competition between the native community and nonnative invaders at the community level, summing to an overall positive relationship at the landscape scale.     

Tree species richness promotes productivity in temperate forests through strong complementarity between species

Understanding the link between biodiversity and ecosystem functioning (BEF) is pivotal in the context of global biodiversity loss. Yet, long-term effects have been explored only weakly, especially for forests, and no clear evidence has been found regarding the underlying mechanisms. We explore the long-term relationship between diversity and productivity using a forest succession model. Extensive simulations show that tree species richness promotes productivity in European temperate forests across a large climatic gradient, mostly through strong complementarity between species. We show that this biodiversity effect emerges because increasing species richness promotes higher diversity in shade tolerance and growth ability, which results in forests responding faster to small-scale mortality events. Our study generalises results from short-term experiments in grasslands to forest ecosystems and demonstrates that competition for light alone induces a positive effect of biodiversity on productivity, thus providing a new angle for explaining BEF relationships.     

Unifying the relationships of species richness to productivity and disturbance

Although species richness has been hypothesized to be highest at 'intermediate' levels of disturbance, empirical studies have demonstrated that the disturbance-diversity relationship can be either negative or positive depending on productivity On the other hand, hypothesized productivity diversity relationships can be positive, negative or unimodal, as confirmed by empirical studies. However, it has remained unclear under what conditions each pattern is realized, and there is little agreement about the mechanisms that generate these diverse patterns. In this study, I present a model that synthesizes these separately developed hypotheses and shows that the interactive effects of disturbance and productivity on the competitive outcome of multispecies dynamics can result in these diverse relationships of species richness to disturbance and productivity The predicted productivity diversity relationship is unimodal but the productivity level that maximizes species richness increases with increasing disturbance. Similarly, the predicted disturbance diversity relationship is unimodal but the peak moves to higher disturbance levels with increasing productivity Further, these patterns are well explained by the opposite effects of productivity and disturbance on competitive outcome that are suggested by the change in community composition along these two environmental gradients: higher productivity favours superior competitors while higher disturbance levels favour inferior competitors.     

The relationship between productivity and multiple aspects of biodiversity in six grassland communities

Biodiversity is a multifaceted concept but most studies examining the association between the biodiversity of a community and its productivity focus only on species richness. Consequently, studies are needed to examine how other facets of biodiversity vary with productivity if we want to have a better understanding of the distribution of biodiversity across our planet. We evaluated how a number of biodiversity measures (species richness, evenness, dominance, rarity, Simpson’s diversity, and Shannon–Weiner diversity) varied across natural productivity gradients at 6 grassland sites in the continental US. Variation in productivity did not account for a substantial amount of variation in any measure of biodiversity at small spatial scales (≈1 m²) at most sites. When productivity accounted for substantial variation in biodiversity, different measures of biodiversity responded to productivity in different ways. For example, dominance changed in a U-shaped fashion along a productivity gradient whereas richness increased in an asymptotic fashion. Consequently, diversity indices, which account for both species richness and evenness, varied in a hump-shaped fashion along the productivity gradient. Our results highlight that an exclusive focus on the association between species richness and productivity provides an incomplete picture of how a community’s biodiversity is related to its functioning.     

Consistent Richness-Biomass Relationship across Environmental Gradients in a Marine Macroalgal-Dominated Subtidal Community on the Western Antarctic Peninsula

Biodiversity loss has spurred the biodiversity-ecosystem functioning research over a range of ecosystems. In Antarctica, however, the relationship of taxonomic and functional diversity with ecosystem properties (e.g., community biomass) has received less attention, despite the presence of sharp and dynamic environmental stress gradients that might modulate these properties. Here, we investigated whether the richness-biomass relationship in macrobenthic subtidal communities is still apparent after accounting for environmental stress gradients in Fildes Bay, King George Island, Antarctica. Measurements of biomass of mobile and sessile macrobenthic taxa were conducted in the austral summer 2013/4 across two environmental stress gradients: distance from nearest glaciers and subtidal depth (from 5 to 30 m). In general, community biomass increased with distance from glaciers and water depth. However, generalised additive models showed that distance from glaciers and depth accounted for negligible proportions of variation in the number of functional groups (i.e., functional richness) and community biomass when compared to taxonomic richness. Functional richness and community biomass were positive and saturating functions of taxonomic richness. Large endemic, canopy-forming brown algae of the order Desmarestiales dominated the community biomass across both gradients. Accordingly, differences in the composition of taxa accounted for a significant and large proportion (51%) of variation in community biomass in comparison with functional richness (10%). Our results suggest that the environmental factors here analysed may be less important than biodiversity in shaping mesoscale (several km) biomass patterns in this Antarctic system. We suggest that further manipulative, hypothesis-driven research should address the role of biodiversity and species’ functional traits in the responses of Antarctic subtidal communities to environmental variation.     

Trophic complementarity drives the biodiversity–ecosystem functioning relationship in food webs

The biodiversity–ecosystem functioning (BEF) relationship is central in community ecology. Its drivers in competitive systems (sampling effect and functional complementarity) are intuitive and elegant, but we lack an integrative understanding of these drivers in complex ecosystems. Because networks encompass two key components of the BEF relationship (species richness and biomass flow), they provide a key to identify these drivers, assuming that we have a meaningful measure of functional complementarity. In a network, diversity can be defined by species richness, the number of trophic levels, but perhaps more importantly, the diversity of interactions. In this paper, we define the concept of trophic complementarity (TC), which emerges through exploitative and apparent competition processes, and study its contribution to ecosystem functioning. Using a model of trophic community dynamics, we show that TC predicts various measures of ecosystem functioning, and generate a range of testable predictions. We find that, in addition to the number of species, the structure of their interactions needs to be accounted for to predict ecosystem productivity.     

Productivity–species richness relationship changes from unimodal to positive linear with increasing spatial scale in the Inner Mongolia steppe

Productivity–species diversity relationships have been a controversial research topic in ecology with scale believed to be among the main reasons for discovering different relationships. We collected data on species diversity (richness) and productivity (peak above-ground biomass) of the Stipa breviflora association in the Inner Mongolia grassland to examine spatial scale dependency and possible underlying mechanisms responsible for the relationships found. One local and seven different landscape scales (the first level corresponds in extent to a 100 × 100 km area, which is increased consecutively by 100 km resulting in the 700 × 700 km area at the highest level) were considered. We found that: (1) unimodal relationships dominated the local scale, but this varied depending on the position along successional gradients; (2) a positive linear relationship was common at larger spatial scales; (3) biotic processes were the most likely primary factor underlying local scale unimodal relationships, but environmental heterogeneity (precipitation patterns) was the main determinant of relationships found at larger spatial scales; (4) our study contributed to other empirical evidence and predictions of theoretical models regarding scale dependency of productivity–species richness relationships; (5) while earlier research demonstrated positive linear species richness–productivity relationships across a number of ecological scales in the Inner Mongolia steppe, our study specifically tested a spectrum of geographical scales to confirm the scale-dependency of this relationship. Lastly, our study emphasized the critical role played by precipitation patterns in controlling biodiversity and grassland ecosystem functioning, which maintains the relatively high level of biodiversity and stable ecosystem processes.     

Developing X-ray Computed Tomography to non-invasively image 3-D root systems architecture in soil

Background

The positive relationship between biodiversity and ecosystem functioning (BEF) is due mainly to complementarity between species. Most BEF studies primarily focused on plant interactions; however, plants are embedded in a dense network of multitrophic interactions above and below the ground, which are likely to play a crucial role in BEF relationships.

Scope

In the present review I point out the relevance of aboveground–belowground interactions as a source of complementarity effects in grassland biodiversity experiments. A review of the current knowledge on the role of decomposers, arbuscular mycorrhizal fungi, rhizobia, plant growth promoting rhizobacteria, invertebrate ecosystem engineers, herbivores, pathogens and predators in biodiversity experiments, indicates that soil biota can drive both positive and negative complementarity between plant species via a multitude of mechanisms.

Conclusions

I pose four main processes by which aboveground–belowground interactions determine positive complementarity effects: enlarging biotope space, mediating legume effects, increasing plant community resistance, and maintaining plant diversity. By contrast, soil biota may also reinforce negative complementarity effects by competing with plants for nutrients or by exerting herbivore or pathogen pressure, thereby reducing community productivity. Thus, considering aboveground–belowground interactions as well as interactions between antagonistic and mutualistic consumers may improve the mechanistic understanding of complementarity effects in plant diversity–ecosystem functioning experiments and should inspire future research.     

How community adaptation affects biodiversity–ecosystem functioning relationships

Evidence is growing that evolutionary dynamics can impact biodiversity–ecosystem functioning (BEF) relationships. However the nature of such impacts remains poorly understood. Here we use a modelling approach to compare random communities, with no trait evolutionary fine‐tuning, and co‐adapted communities, where traits have co‐evolved, in terms of emerging biodiversity–productivity, biodiversity–stability and biodiversity–invasion relationships. Community adaptation impacted most BEF relationships, sometimes inverting the slope of the relationship compared to random communities. Biodiversity–productivity relationships were generally less positive among co‐adapted communities, with reduced contribution of sampling effects. The effect of community‐adaptation, though modest regarding invasion resistance, was striking regarding invasion tolerance: co‐adapted communities could remain very tolerant to invasions even at high diversity. BEF relationships are thus contingent on the history of ecosystems and their degree of community adaptation. Short‐term experiments and observations following recent changes may not be safely extrapolated into the future, once eco‐evolutionary feedbacks have taken place.     

The relationship between species richness and productivity in four typical grasslands of northern China

The relationship between plant species richness and primary productivity has long been acentral topic in biodiversity research.In this paper,we examine the relationship between species richness and productivity in four typical grasslands of Northern China at different spatial scales.At the community scale,a positive correlation was found for six of seven communities.A unimodal pattern was found only for one community (Stipa glareosa community),while at a large scale (vegetation type or landscape/region),the relationship was also found significantly positive.Species richness ranged from 4 to 35 species,and community aboveground productiand aboveground productivity were found in alpine meadow,followed by meadow steppe,typical steppe and desert steppe.     

Determinants of species evenness in a neotropical bat ensemble

Evenness is an important property of communities. Species richness alone does not capture the fact that one or a few species may dominate total abundance and biomass of a community. This in turn has important consequences for ecosystem functioning and species interactions. Evenness has been observed to vary systematically along environmental and productivity gradients. However, a truly general theory about which factors control evenness in a community has yet to emerge. Prior research on evenness has suggested that high richness, biomass and abundance should lead to lower community evenness in our study system of bats in Panama. However, only few empirical studies examine the simultaneous effects of species richness, biomass or abundance on evenness. For the first time, we applied path analysis in the study of evenness to tease apart the relative importance and direction (positive or negative) of causality among these three factors. As predicted, we found that evenness decreases with increasing species richness, abundance and biomass. The negative effect of abundance was mediated by the positive joint effect of biomass and richness. The selected models varied in the strength of the correlation between the three variables with evenness but their direction was consistent. Overall, we argue that rarity, high mobility and differences in resource availability at sites with lower environmental stress can explain the negative effects of richness on evenness.     

Reciprocal causality between marine macroalgal diversity and productivity in an upwelling area

The relationship between biodiversity and ecosystem functioning has become a prominent topic in the ecological literature. However, the contemporary approach that species diversity controls primary productivity contrasts with the historical perspective that species diversity responds to productivity. Moreover, previous experimental results have not been consistent with the patterns observed in nature. To resolve these questions, the multivariate productivity–diversity (MPD) hypothesis proposes a bidirectional relationship between diversity and productivity. It predicts that the resource supply, expressed in terms of resource availability and imbalance, establishes the number of species that can locally coexist. Simultaneously, the resource supply also indirectly affects biomass production, determining the form and cause of the effects of species richness on resource use and biomass. To test the MPD hypothesis, we conducted three field experiments with a subtidal marine macroalgal community using a seasonal upwelling process as a driver of distinct levels of nutrient supply. Seasonally, macroalgal species richness and biomass were assessed and experimental manipulations conducted to investigate the relative importance of species richness and identity effects on biomass production and the mechanisms underlying these. Changes in macroalgal biomass and species richness were observed in response to the nutrient supply. Stronger effects of species identity were detected for all periods investigated, although species richness effects also occurred to some extent. The magnitudes of the net biodiversity and of the complementarity effects were a unimodal function of nutrient supply, whereas a concave‐up curve was observed for selection effects. The nutrient supply directly affected the number of species that dominated the local community and, consequently, determined the efficiency with which resources were exploited and converted to biomass. Our results provide evidence consistent with the MPD hypothesis and aids in explaining the discrepancies between experimental results and natural patterns through the merging of two contrasting perspectives in ecology.     

On the importance of the negative selection effect for the relationship between biodiversity and ecosystem functioning

Much of our knowledge on biodiversity and ecosystem functioning comes from studies examining the effects of biodiversity on biomass production within a trophic group. A large number of these studies have found that increasing biodiversity tends to increase biomass production, leading many ecologists to believe that there exists a general positive relationship between biodiversity and ecosystem functioning. Here we argue that such a positive relationship may not be general, particularly for ecosystem functions other than biomass. Our argument centers on the potential importance of the negative selection effect, which operates where competitively dominant species do not contribute significantly to the function of interest. We suggest that negative selection effects may be potentially common for non-biomass functions, for which species competitive ability may often be a poor indictor of its functional impact. We conclude that diverse (positive, negative, and neutral) BEF relationships are possible for non-biomass functions and that for a particular function, the exact form of the BEF relationship may depend on how species functional impacts relate to their competitive abilities in the community.     

生物多样性与生态系统生产力：为什么野外观测和受控实验结果不一致？

 人们担心生物多样性的空前丧失会危及到生态系统的服务功能,因此有关生物多样性-生态系统生产力这一古老命题的讨论成为当今生态学的热点议题之一。20世纪90年代以David Tilman和Shahid Naeem为代表的生态学家利用大规模的受控实验,对物种多样性与生态系统功能的诸多方面进行     

Using root traits to understand temporal changes in biodiversity effects in grassland mixtures

Biodiversity–ecosystem functioning (BEF) studies typically show that species richness enhances community biomass, but the underlying mechanisms remain debated. Here, we combine metrics from BEF research that distinguish the contribution of dominant species (selection effects, SE) from those due to positive interactions such as resource partitioning (complementarity effects, CE) with a functional trait approach in an attempt to reveal the functional characteristics of species that drive community biomass in species mixtures. In a biodiversity experiment with 16 plant species in monocultures, 4‐species and 16‐species mixtures, we used aboveground biomass to determine the relative contributions of CE and SE to biomass production in mixtures in the second, dry year of the experiment. We also measured root traits (specific root length, root length density, root tissue density and the deep root fraction) of each species in monocultures and linked the calculated community weighted mean (CWM) trait values and trait diversity of mixtures to CE and SE. In the second year of the experiment, community biomass, CE and SE increased compared to the first year. The contribution of SE to this positive effect was greater than that of CE. The increased contribution of SE was associated with root traits: SE increased most in communities with high abundance of species with deep, thick and dense roots. In contrast, changes in CE were not related to trait diversity or CWM trait values. Together, these results suggest that increased positive effects of species richness on community biomass in a dry year were mainly driven by increased dominance of deep‐rooting species, supporting the insurance hypothesis of biodiversity. Positive CE indicates that other positive interactions did occur, but we could not find evidence that belowground resource partitioning or facilitation via root trait diversity was important for community productivity in our biodiversity experiment.