Oscillating trophic control induces community reorganization in a marine ecosystem

Understanding how climate regulates trophic control may help to elucidate the causes of transitions between alternate ecosystem states following climate regime shifts. We used a 34-year time series of the abundance of Pacific cod ( Gadus macrocephalus ) and five prey species to show that the nature of trophic control in a North Pacific ecosystem depends on climate state. Rapid warming in the 1970s caused an oscillation between bottom–up and top–down control. This shift to top–down control apparently contributed to the transition from an initial, prey-rich ecosystem state to the final, prey-poor state. However, top–down control could not be detected in the final state without reference to the initial state and transition period. Complete understanding of trophic control in ecosystems capable of transitions between alternate states may therefore require observations spanning more than one state.     

The relative contributions of species richness and species composition to ecosystem functioning

How species diversity influences ecosystem functioning has been the subject of many experiments and remains a key question for ecology and conservation biology. However, the fact that diversity cannot be manipulated without affecting species composition makes this quest methodologically challenging. Here, we evaluate the relative importance of diversity and of composition on biomass production, by using partial Mantel tests for one variable while controlling for the other. We analyse two datasets, from the Jena (2002–2008) and the Grandcour (2008–2009) Experiments. In both experiments, plots were sown with different numbers of species to unravel mechanisms underlying the relationship between biodiversity and ecosystem functioning (BEF). Contrary to Jena, plots were neither mowed nor weeded in Grandcour, allowing external species to establish. Based on the diversity–ecosystem functioning and competition theories, we tested two predictions: 1) the contribution of composition should increase with time; 2) the contribution of composition should be more important in non‐weeded than in controlled systems. We found support for the second hypothesis, but not for the first. On the contrary, the contribution of species richness became markedly more important few years after the start of the Jena Experiment. This result can be interpreted as suggesting that species complementarity, rather than intraspecific competition, is the driving force in this system. Finally, we explored to what extent the estimated relative importance of both factors varied when measured on different spatial scales of the experiment (in this case, increasing the number of plots included in the analyses). We found a strong effect of scale, suggesting that comparisons between studies, and more generally the extrapolation of results from experiments to natural situations, should be made with caution.     

Effects of species evenness can be derived from species richness – ecosystem functioning relationships

Although species richness effects on ecosystem functioning have been studied thoroughly in countless experiments, the effects of the other side of diversity – species evenness – remain less identified, especially at high species richness. Due to the large number of different model ecosystems that need to be created, the explanatory power of the experimental approach for evenness is indeed limited. We show here that experimental studies on the influence of species richness on ecosystem functions contain hidden information on the influence of species evenness. Both the effects of maximum and minimum evenness, and of a key set of intermediate evenness levels, can be derived from species richness – ecosystem function curves, and that for every richness level, by using communities with low species richness as the equivalent of highly uneven communities with higher richness. We show that evenness effects on ecosystem functioning have the same direction as richness effects, however with increasing effect sizes at higher richness levels. We validated our technique for a wide range of ecosystem functions and applied it to the species richness – community biomass data from an existing biodiversity experiment. Our approach could provide a fast and easy alternative to resource‐intensive experiments in which evenness is experimentally varied, as we can build on the elaborate existing literature on species richness to assess its effects.     

Behavioural interactions between ecosystem engineers control community species richness

Behavioural interactions between ecosystem engineers may strongly influence community structure. We tested whether an invasive ecosystem engineer, the alga Caulerpa taxifolia , indirectly facilitated community diversity by modifying the behaviour of a native ecosystem engineer, the clam Anadara trapezia , in southeastern Australia . In this study, clams in Caulerpa -invaded sediments partially unburied themselves, extending >30% of their shell surface above the sediment, providing rare, hard substrata for colonization. Consequently, clams in Caulerpa had significantly higher diversity and abundance of epibiota compared with clams in unvegetated sediments. To isolate the role of clam burial depth from direct habitat influences or differential predation by habitat, we manipulated clam burial depth, predator exposure and habitat ( Caulerpa or unvegetated) in an orthogonal experiment. Burial depth overwhelmingly influenced epibiont species richness and abundance, resulting in a behaviourally mediated facilitation cascade. That Caulerpa controls epibiont communities by altering Anadara burial depths illustrates that even subtle behavioural responses of one ecosystem engineer to another can drive extensive community-wide facilitation.     

Effects of organism size and community composition on ecosystem functioning

We tested (1) if the size of dominant species influenced ecosystem functioning in food webs consisting of bacteria, algae, and protozoa; (2) whether those effects changed in importance through time; and (3) how those effects compared with differences in diversity among experimental food webs. We constructed food webs using two size fractions of organisms that differed in individual mass by approximately two orders of magnitude. We measured total biomass and respiration (total CO₂ production) as two aspects of ecosystem functioning. We also compared these size‐dependent patterns in functioning across two levels of species richness. Initially, organism size strongly influenced total community biomass. With time, however, biomass and respiration eventually converged in communities dominated by large or small species. We conclude that after sufficient time for community development any differences in ecosystem functioning resulted from differences in community composition, including species richness, but not the size of the dominant organisms.     

Plant species richness and shrub cover attenuate drought effects on ecosystem functioning across Patagonian rangelands

Drought is an increasingly common phenomenon in drylands as a consequence of climate change. We used 311 sites across a broad range of environmental conditions in Patagonian rangelands to evaluate how drought severity and temperature (abiotic factors) and vegetation structure (biotic factors) modulate the impact of a drought event on the annual integral of normalized difference vegetation index (NDVI-I), our surrogate of ecosystem functioning. We found that NDVI-I decreases were larger with both increasing drought severity and temperature. Plant species richness (SR) and shrub cover (SC) attenuated the effects of drought on NDVI-I. Grass cover did not affect the impacts of drought on NDVI-I. Our results suggest that warming and species loss, two important imprints of global environmental change, could increase the vulnerability of Patagonian ecosystems to drought. Therefore, maintaining SR through appropriate grazing management can attenuate the adverse effects of climate change on ecosystem functioning.     

Mussels as ecosystem engineers: Their contribution to species richness in a rocky littoral community

Mussels are important ecosystem engineers in marine benthic systems because they aggregate into beds, thus modifying the nature and complexity of the substrate. In this study, we evaluated the contribution of mussels (Brachidontes rodriguezii, Mytilus edulis platensis, and Perna perna) to the benthic species richness of intertidal and shallow subtidal communities at Cerro Verde (Uruguay). We compared the richness of macro-benthic species between mussel-engineered patches and patches without mussels but dominated by algae or barnacles at a landscape scale (all samples), between tidal levels, and between sites distributed along a wave exposition gradient. Overall, we found a net increase in species richness in samples with mussels (35 species), in contrast to samples where mussels were naturally absent or scarce (27 species). The positive trend of the effect did not depend upon tidal level or wave exposition, but its magnitude varied between sites. Within sites, a significant positive effect was detected only at the protected site. Within the mussel-engineered patches, the richness of all macro-faunal groups (total, sessile and mobile) was positively correlated with mussel abundance. This evidence indicates that the mussel beds studied here were important in maintaining species richness at the landscape-level, and highlights that beds of shelled bivalves should not be neglected as conservation targets in marine benthic environments.     

Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.     

Invasion rates increase with species richness in a marine epibenthic community by two mechanisms

It is widely believed that, when extrinsic conditions are similar, the likelihood of species invading established assemblages decreases with increasing species richness of the recipient community. Here we show that, for a sessile marine invertebrate community, invasion of patches increases with richness of the patch. We show that invasions can increase with local species richness by two distinct mechanisms. In the first, opportunistic colonisers with traits typical of invasive species colonise species-rich patches at higher rates because speciose patches are dominated by small colonies and mortality rates of small colonies are greater than that of large ones. Thus, mortality provides bare space for opportunists to colonise more frequently in species-rich patches. In the second, some species avoid colonising open areas of free space but preferentially associate with established colonies of particular other species, and a given preferred associate is more likely to occur in species-rich than in species-poor patches. These patterns are the result of particular properties of individual species and local species dynamics, and show that reduced risk of invasion is not necessarily an intrinsic property of species-rich communities. We conclude that resistance to invasion will be determined by the properties of the particular component species and emergent dynamics of the recipient community, and not by an aggregate community property such as richness.     

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

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Species richness destabilizes ecosystem functioning in experimental aquatic microcosms

The relationship between species diversity and ecosystem stability has long interested ecologists, yet no consensus has been reached and the underlying mechanisms remain unclear. We used five unicellular algal species, cultured in all possible combinations, to assemble microcosms containing 1 to 5 algal species, on which a cold perturbation was imposed. Our aim was to find whether and how species richness begets ecosystem resistance and resilience. In the experiment, the species-rich communities produced more biomass than the species-poor ones, either in pre-, under- or post-perturbation conditions. The positive diversity–biomass relationship was weakened by the perturbation, and fully restored one week after the perturbation. The diverse communities showed greater absolute biomass reduction during the perturbation than did species-poor systems. Resistance of community, measured by the relative change in biomass from pre- to under-perturbation, decreased with species richness. All the species showed significant reduction in biomass when stressed, without any density compensation among species in diverse communities; and the ratio of biomass change in each species was independent of diversity. The overyielding effect, measured as relative yield total, remained constant from pre- to under-perturbation; and the selection and complementarity effects played equal roles for the biodiversity effect on biomass production, and their relative importance was not altered by the perturbation. These results suggest that similar responses of different species to environmental perturbations may limit the insurance effect of biodiversity, and lead to an inverse diversity–resistance relationship.     

Multi-level trophic cascades in a heavily exploited open marine ecosystem

Anthropogenic disturbances intertwined with climatic changes can have a large impact on the upper trophic levels of marine ecosystems, which may cascade down the food web. So far it has been difficult to demonstrate multi-level trophic cascades in pelagic marine environments. Using field data collected during a 33-year period, we show for the first time a four-level community-wide trophic cascade in the open Baltic Sea. The dramatic reduction of the cod (Gadus morhua) population directly affected its main prey, the zooplanktivorous sprat (Sprattus sprattus), and indirectly the summer biomass of zooplankton and phytoplankton (top-down processes). Bottom-up processes and climate-hydrological forces had a weaker influence on sprat and zooplankton, whereas phytoplankton variation was explained solely by top-down mechanisms. Our results suggest that in order to dampen the occasionally harmful algal blooms of the Baltic, effort should be addressed not only to control anthropogenic nutrient inputs but also to preserve structure and functioning of higher trophic levels.     

物种多样性与生态系统功能：影响机制及有关假说

本文主要介绍了有关物种多样性对生态系统功能的影响机制及有关假说,包括冗余种假说、铆钉假说、不确定假说、无效假说、补偿／关键种假说、非线性假说、单调／驼峰模型假说等。尽管这些假说的提出都有一定的理论或实验研究基础,但到目前为止,还没有一种假说被普遍认为反映了两者之间关系的实际情形（如果确实存在这样一种普适机制）。通过分析,我们发现在这些假说之间存在一定的内在联系,它们或者互相包含,或者互相补充,并且都可以归结到冗余种假说有关。     

Effects of foundation species genotypic diversity on subordinate species richness in an assembling community

Foundation (dominant or matrix) species play a key role in structuring plant communities, influencing processes from population to ecosystem scales. However, the effects of genotypic diversity of foundation species on these processes have not been thoroughly assessed in the context of assembling plant communities. We modified the classical filter model of community assembly to include genotypic diversity as part of the biotic filter. We hypothesized that the proportion of fit genotypes (i.e. competitively superior and dominant) affects niche space availability for subordinate species to establish with consequence for species diversity. To test this hypothesis, we used an individual‐based simulation model where a foundation species of varying genotypic diversity (number of genotypes and variability among genotypes) competes for space with subordinate species on a spatially heterogeneous lattice. Our model addresses a real and practical problem in restoration ecology: choosing the level of genetic diversity of re‐introduced foundation and subordinate species. Genotypic diversity of foundation species significantly affected equilibrium community diversity, measured as species richness, either positively or negatively, depending upon environmental heterogeneity. Increases in genotypic diversity gave the foundation species a wider niche breadth. Under conditions of high environmental heterogeneity, this wider niche breadth decreased niche space for other species, lowering species richness with increased genotypic diversity until the genotypes of the foundation species saturated the landscape. With a low level of environmental heterogeneity, increasing genotypic diversity caused the foundation species niche breadth to be overdispersed, resulting in a weak positive relationship with species richness. Under these conditions, some genotypes are maladapted to the environment lowering fitness of the foundation species. These effects of genotypic diversity were secondary to the larger effects of overall foundation species fitness and environmental heterogeneity. The novel aspect of incorporating genotype diversity in combination with environmental heterogeneity in community assembly models include predictions of either positive or negative relationships between species diversity and genotypic diversity depending on environmental heterogeneity, and the conditions under which these factors are potentially relevant. Mechanistically, differential niche availability is imposed by the foundation species.     

Context dependence of marine ecosystem engineer invasion impacts on benthic ecosystem functioning

Introduced ecosystem engineers can severely modify the functioning on invaded systems. Species-level effects on ecosystem functioning (EF) are context dependent, but the effects of introduced ecosystem engineers are frequently assessed through single-location studies. The present work aimed to identify sources of context-dependence that can regulate the impacts of invasive ecosystem engineers on ecosystem functioning. As model systems, four locations where the bivalve Ruditapes philippinarum (Adams and Reeve) has been introduced were investigated, providing variability in habitat characteristics and community composition. As a measure of ecosystem engineering, the relative contribution of this species to community bioturbation potential was quantified at each site. The relevance of bioturbation to the local establishment of the mixing depth of marine sediments (used as a proxy for EF) was quantified in order to determine the potential for impact of the introduced species at each site. We found that R. philippinarum is one of the most important bioturbators within analysed communities, but the relative importance of this contribution at the community level depended on local species composition. The net contribution of bioturbation to the establishment of sediment mixing depths varied across sites depending on the presence of structuring vegetation, sediment granulometry and compaction. The effects of vegetation on sediment mixing were previously unreported. These findings indicate that the species composition of invaded communities, and the habitat characteristics of invaded systems, are important modulators of the impacts of introduced species on ecosystem functioning. A framework that encompasses these aspects for the prediction of the functional impacts of invasive ecosystem engineers is suggested, supporting a multi-site approach to invasive ecology studies concerned with ecosystem functioning.     

A multiscale analysis of herbaceous species richness in a Mediterranean ecosystem

Aims Studies of species distribution patterns traditionally have been conducted at a single scale, often overlooking species–environment relationships operating at finer or coarser scales. Testing diversity-related hypotheses at multiple scales requires a robust sampling design that is nested across scales. Our chief motivation in this study was to quantify the contributions of different predictors of herbaceous species richness at a range of local scales.Methods Here, we develop a hierarchically nested sampling design that is balanced across scales, in order to study the role of several environmental factors in determining herbaceous species distribution at various scales simultaneously. We focus on the impact of woody vegetation, a relatively unexplored factor, as well as that of soil and topography. Light detection and ranging (LiDAR) imaging enabled precise characterization of the 3D structure of the woody vegetation, while acoustic spectrophotometry allowed a particularly high-resolution mapping of soil CaCO 3 and organic matter contents.Important findings We found that woody vegetation was the dominant explanatory variable at all three scales (10, 100 and 1000 m 2), accounting for more than 60% of the total explained variance. In addition, we found that the species richness–environment relationship was scale dependent. Many studies that explicitly address the issue of scale do so by comparing local and regional scales. Our results show that efforts to conserve plant communities should take into account scale dependence when analyzing species richness–environment relationships, even at much finer resolutions than local vs. regional. In addition, conserving heterogeneity in woody vegetation structure at multiple scales is a key to conserving diverse herbaceous communities.     

Effects of macroalgal species identity and richness on primary production in benthic marine communities

Plant biodiversity can enhance primary production in terrestrial ecosystems, but biodiversity effects are largely unstudied in the ocean. We conducted a series of field and mesocosm experiments to measure the relative effects of macroalgal identity and richness on primary productivity (net photosynthetic rate) and biomass accumulation in hard substratum subtidal communities in North Carolina, USA. Algal identity consistently and strongly affected production; species richness effects, although often significent, were subtle. Partitioning of the net biodiversity effect indicated that complementarity effects were always positive and species were usually more productive in mixtures than in monoculture. Surprisingly, slow growing species performed relatively better in the most diverse treatments than the most productive species, thus selection effects were consistently negative. Our results suggest that several basic mechanisms underlying terrestrial plant biodiversity effects also operate in algal-based marine ecosystems, and thus may be general.     

Placing biodiversity and ecosystem functioning in context: environmental perturbations and the effects of species richness in a stream field experiment

Greater biodiversity is often associated with increased ecosystem process rates, and is expected to enhance the stability of ecosystem functioning under abiotic stress. However, these relationships might themselves be altered by environmental factors, complicating prediction of the effects of species loss in ecosystems subjected to abiotic stress. In boreal streams, we investigated effects of biodiversity and two abiotic perturbations on three related indices of ecosystem functioning: leaf decomposition, detritivore leaf processing efficiency (LPE) and detritivore growth. Replicate field enclosures containing leaves and detritivore assemblages were exposed to liming and nutrient enrichment, raising pH and nutrient levels. Both treatments constitute perturbations for our naturally acidic and nutrient-poor streams. We also varied detritivore species richness and density. The effects of the abiotic and diversity manipulations were similar in magnitude, but whereas leaf decomposition increased by 18% and 8% following liming and nutrient enrichment, respectively, increased detritivore richness reduced leaf decomposition (6%), detritivore LPE (19%) and detritivore growth (12%). The detritivore richness effect on growth was associated with negative trait-independent complementarity, indicating interspecific interference competition. These interactions were apparently alleviated in both enriched and limed enclosures, as trait-independent complementarity became less negative. LPE increased with detritivore density in the monocultures, indicating benefits of intra-specific aggregation that outweighed the costs of intra-specific competition, and dilution of these benefits probably contributed to lowered leaf decomposition in the species mixtures. Finally, the effects of liming were reduced in most species mixtures relative to the monocultures. These results demonstrate how environmental changes might regulate the consequences of species loss for functioning in anthropogenically perturbed ecosystems, and highlight potential influences of biodiversity on functional stability. Additionally, the negative effects of richness and positive effects of density in our field study were opposite to previous laboratory observations, further illustrating the importance of environmental context for biodiversity–ecosystem functioning relationships.     

Effects of woody plant species richness on mammal species richness in southern Africa

Aim To determine the relationship between the species richness of woody plants and that of mammals after accounting for the effect of environmental variables. Location Southern Africa, including Namibia, South Africa, Lesotho, Swaziland, Botswana, Zimbabwe, and part of Mozambique. Methods We used a comprehensive dataset including the species richness of mammals and of woody plants and environmental variables for 118 quadrats (each of 25,000 km²) across southern Africa, and used structural equation models (SEMs) and spatial regressions to examine the relationship between the species richness of woody plants and of mammal trophic guilds (herbivores, insectivores, carni/omnivores) and habitat guilds (aquatic/fossorial, ground‐living, climbers, aerial), after controlling for environment. We compared the results of SEMs with those of single‐predictor regressions (without controlling for environment) and of spatial regressions (controlling for both environment and residual spatial autocorrelation). Results The geographical variation of mammal species richness in southern Africa was strongly and positively related to that of woody plant species richness, and this relationship held for most mammal guilds even when the influence of environment and spatial autocorrelation had been accounted for. However, the effect of woody plant species richness on the richness of aquatic/fossorial species almost disappeared after controlling for environment, suggesting that the congruence in species richness patterns between these two groups results from similar responses to the same environmental variables. For many mammal guilds, the relative role of environmental predictors as measured by standardized partial regression coefficients changed depending on whether non‐spatial single‐predictor regressions, non‐spatial SEMs, or spatial regressions were used. Main conclusions Woody plants are important determinants of the species richness of most mammal guilds in southern Africa, even when controlling for environment and residual spatial autocorrelation. Environmental correlates with animal species richness as measured by simple correlations or single‐predictor regressions might not always reflect direct effects; they might, at least to some degree, result from indirect effects via woody plants. Interpretations of the strength of the effect of environmental variables on mammal species richness in southern Africa depend largely on whether spatial or non‐spatial models are used. We therefore stress the need for caution when interpreting environmental ‘effects’ on broad‐scale patterns of species richness if spatial and non‐spatial methods yield contrasting results.