Biodiversity and ecosystem stability across scales in metacommunities

Although diversity–stability relationships have been extensively studied in local ecosystems, the global biodiversity crisis calls for an improved understanding of these relationships in a spatial context. Here, we use a dynamical model of competitive metacommunities to study the relationships between species diversity and ecosystem variability across scales. We derive analytic relationships under a limiting case; these results are extended to more general cases with numerical simulations. Our model shows that, while alpha diversity decreases local ecosystem variability, beta diversity generally contributes to increasing spatial asynchrony among local ecosystems. Consequently, both alpha and beta diversity provide stabilising effects for regional ecosystems, through local and spatial insurance effects respectively. We further show that at the regional scale, the stabilising effect of biodiversity increases as spatial environmental correlation increases. Our findings have important implications for understanding the interactive effects of global environmental changes (e.g. environmental homogenisation) and biodiversity loss on ecosystem sustainability at large scales.     

Association between temporal and spatial beta diversity in phytoplankton

The rates of temporal and spatial species turnover have been compared in different organisms and scales, revealing that both are not independent but, rather, associated. However, the knowledge is limited for the association between spatial turnover and temporal turnover. Here, we performed two investigations of the phytoplankton composition in the lakes of the Yangtze River catchment in China in the spring and summer of 2012, which covered regional spatial scale and two‐season temporal scale. We analysed the association between temporal and spatial species turnover in phytoplankton. The results showed that 1) the two‐season temporal turnover of phytoplankton varied based on the mean values and the coefficient of variance of environmental variables, and pH was the most important variable negatively affecting the temporal turnover; 2) the spatial beta diversity of phytoplankton in summer was higher than that in spring, and the distance decay pattern was significant in summer, but not in spring; 3) the variation in spatial turnover in spring and summer was attributed to the primary environmental variables (nitrogen, phosphorus and underwater available light) and broader‐scale spatial variables; 4) the proportion of jointly explained variation of spatial Bray–Curtis dissimilarity by the environment and space increased from ~38% (spring) to ~55% (summer), which was mainly due to the variation in spatially structured environmental variables during the two‐season temporal turnover, such as pH and ion concentrations; 5) the community compositions in summer were more similar between the lakes with similar two‐season temporal turnover. These results indicate that the spatial turnover of phytoplankton composition in summer was partially predetermined by the variation in environmental variables and phytoplankton composition during the process of two‐season temporal turnover, and highlight the understanding of temporal variations in spatial beta diversity as well as the underlying assembly mechanisms in phytoplankton.     

Stability and synchrony across ecological hierarchies in heterogeneous metacommunities: linking theory to data

Understanding stability across ecological hierarchies is critical for landscape management in a changing world. Recent studies showed that synchrony among lower‐level components is key to scaling temporal stability across two hierarchical levels, whether spatial or organizational. But an extended framework that integrates both spatial scale and organizational level simultaneously is required to clarify the sources of ecosystem stability at large scales. However, such an extension is far from trivial when taking into account the spatial heterogeneities in real‐world ecosystems. In this paper, we develop a partitioning framework that bridges variability and synchrony measures across spatial scales and organizational levels in heterogeneous metacommunities. In this framework, metacommunity variability is expressed as the product of local‐scale population variability and two synchrony indices that capture the temporal coherence across species and space, respectively. We develop an R function ‘var.partition’ and apply it to five types of desert plant communities to illustrate our framework and test how diversity shapes synchrony and variability at different hierarchical levels. As the observation scale increased from local populations to metacommunities, the temporal variability of plant productivity was reduced mainly by factors that decreased species synchrony. Species synchrony decreased from local to regional scales, and spatial synchrony decreased from species to community levels. Local and regional species diversity were key factors that reduced species synchrony at the two scales. Moreover, beta diversity contributed to decreasing spatial synchrony among communities. We conclude that our new framework offers a valuable toolbox for future empirical studies to disentangle the mechanisms and pathways by which ecological factors influence stability at large scales.     

Intraspecific phenotypic variation in a fish predator affects multitrophic lake metacommunity structure

Contemporary insights from evolutionary ecology suggest that population divergence in ecologically important traits within predators can generate diversifying ecological selection on local community structure. Many studies acknowledging these effects of intraspecific variation assume that local populations are situated in communities that are unconnected to similar communities within a shared region. Recent work from metacommunity ecology suggests that species dispersal among communities can also influence species diversity and composition but can depend upon the relative importance of the local environment. Here, we study the relative effects of intraspecific phenotypic variation in a fish predator and spatial processes related to plankton species dispersal on multitrophic lake plankton metacommunity structure. Intraspecific diversification in foraging traits and residence time of the planktivorous fish alewife (Alosa pseudoharengus) among coastal lakes yields lake metacommunities supporting three lake types which differ in the phenotype and incidence of alewife: lakes with anadromous, landlocked, or no alewives. In coastal lakes, plankton community composition was attributed to dispersal versus local environmental predictors, including intraspecific variation in alewives. Local and beta diversity of zooplankton and phytoplankton was additionally measured in response to intraspecific variation in alewives. Zooplankton communities were structured by species sorting, with a strong influence of intraspecific variation in A. pseudoharengus. Intraspecific variation altered zooplankton species richness and beta diversity, where lake communities with landlocked alewives exhibited intermediate richness between lakes with anadromous alewives and without alewives, and greater community similarity. Phytoplankton diversity, in contrast, was highest in lakes with landlocked alewives. The results indicate that plankton dispersal in the region supplied a migrant pool that was strongly structured by intraspecific variation in alewives. This is one of the first studies to demonstrate that intraspecific phenotypic variation in a predator can maintain contrasting patterns of multitrophic diversity in metacommunities.     

Regional invariance among microbial communities

Microbial ecology has focused much on causes of between-site variation in community composition. By analysing five data-sets each of aquatic bacteria and phytoplankton, we demonstrated that microbial communities show a large degree of similarity in community composition and that abundant taxa were widespread, a typical pattern for many metazoan metacommunities. The regional abundance of taxa explained on average 85 and 41% of variation in detection frequency and 58 and 31% of variation in local abundances for bacteria and phytoplankton, respectively. However, regional abundance explained less variation in local abundances with increasing environmental variation between sites within data-sets. These findings indicate that the studies of microbial assemblages need to consider similarities between communities to better understand the processes underlying the assembly of microbial communities. Finally, we propose that the degree of regional invariance can be linked to the evolution of microbes and the variation in ecosystem functions performed by microbial communities.
Ecology Letters (2010) 13: 118–127     

Spatial scaling of arbuscular mycorrhizal fungal diversity is affected by farming practice

Evidence suggests that microbial communities show patterns of spatial scaling which can be driven by geographical distance and environmental heterogeneity. Here we demonstrate that human management can have a major impact on microbial distribution patterns at both the local and landscape scale. Mycorrhizal fungi are vital components of terrestrial ecosystems, forming a mutualistic symbiosis with plant roots which has a major impact on above ground ecology and productivity. We used contrasting agricultural systems to investigate the spatial scaling of the most widespread mycorrhizal fungus group, the arbuscular mycorrhizal fungi (AMF). Using multiple sampling sites with a maximum separation of 250 km we describe for the first time the roles which land management, environmental heterogeneity and geographical distance play in determining spatial patterns of microbial distribution. Analysis of AMF taxa–area relationships at each sampling site revealed that AMF diversity and spatial turnover was greater under organic relative to conventional farm management. At the regional scale (250 km) distance–decay analyses showed that there was significant change in AMF community composition with distance, and that this was greater under organic relative to conventional management. Environmental heterogeneity was found to be the major factor determining turnover of AMF taxa at the landscape scale. Overall we demonstrate that human management can play a key role in determining the turnover of microbial communities at both the local and regional scales.     

Temporal variation in phytoplankton beta diversity patterns and metacommunity structures across subtropical reservoirs

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A comparative analysis of metacommunity types in the freshwater realm

Most metacommunity studies have taken a direct mechanistic approach, aiming to model the effects of local and regional processes on local communities within a metacommunity. An alternative approach is to focus on emergent patterns at the metacommunity level through applying the elements of metacommunity structure (EMS; Oikos, 97, 2002, 237) analysis. The EMS approach has very rarely been applied in the context of a comparative analysis of metacommunity types of main microbial, plant, and animal groups. Furthermore, to our knowledge, no study has associated metacommunity types with their potential ecological correlates in the freshwater realm. We assembled data for 45 freshwater metacommunities, incorporating biologically highly disparate organismal groups (i.e., bacteria, algae, macrophytes, invertebrates, and fish). We first examined ecological correlates (e.g., matrix properties, beta diversity, and average characteristics of a metacommunity, including body size, trophic group, ecosystem type, life form, and dispersal mode) of the three elements of metacommunity structure (i.e., coherence, turnover, and boundary clumping). Second, based on those three elements, we determined which metacommunity types prevailed in freshwater systems and which ecological correlates best discriminated among the observed metacommunity types. We found that the three elements of metacommunity structure were not strongly related to the ecological correlates, except that turnover was positively related to beta diversity. We observed six metacommunity types. The most common were Clementsian and quasi‐nested metacommunity types, whereas Random, quasi‐Clementsian, Gleasonian, and quasi‐Gleasonian types were less common. These six metacommunity types were best discriminated by beta diversity and the first axis of metacommunity ecological traits, ranging from metacommunities of producer organisms occurring in streams to those of large predatory organisms occurring in lakes. Our results showed that focusing on the emergent properties of multiple metacommunities provides information additional to that obtained in studies examining variation in local community structure within a metacommunity.     

Local, among-site, and regional diversity patterns of benthic macroinvertebrates in high altitude waterbodies: do ponds differ from lakes?

In this study we aimed at comparing invertebrate diversity of high altitude lakes and ponds along hierarchical spatial scales. We compared local, among-site, and regional diversity of benthic macroinvertebrates in 25 ponds and 34 lakes in the Tatra Mountains, central Europe. The ponds showed significantly lower local diversity, higher among-site diversity and similar regional diversity than the lakes. The species–area relationships (SAR), habitat heterogeneity, and environmental harshness are assumed as drivers for the local diversity patterns. An ecological threshold separating pond and lake systems emerged at an area of 2 ha, where the SAR pattern changed significantly. Differences in species turnover between these systems were likely driven by greater environmental variability and isolation of the ponds. High altitude ponds neither significantly support greater regional diversity nor higher number of unique taxa than lakes. The higher among-site diversity of ponds relative to lakes highlights the relevance of ponds for regional diversity in mountain areas.     

Effect of (a)synchronous light fluctuation on diversity,functional and structural stability of a marine phytoplankton metacommunity

Disentangling the mechanisms that maintain the stability of communities and ecosystem properties has become a major research focus in ecology in the face of anthropogenic environmental change. Dispersal plays a pivotal role in maintaining diversity in spatially subdivided communities, but only a few experiments have simultaneously investigated how dispersal and environmental fluctuation affect community dynamics and ecosystem stability. We performed an experimental study using marine phytoplankton species as model organisms to test these mechanisms in a metacommunity context. We established three levels of dispersal and exposed the phytoplankton to fluctuating light levels, where fluctuations were either spatially asynchronous or synchronous across patches of the metacommunity. Dispersal had no effect on diversity and ecosystem function (biomass), while light fluctuations affected both evenness and community biomass. The temporal variability of community biomass was reduced by fluctuating light and temporal beta diversity was influenced interactively by dispersal and fluctuation, whereas spatial variability in community biomass and beta diversity were barely affected by treatments. Along the establishing gradient of species richness and dominance, community biomass increased but temporal variability of biomass decreased, thus highest stability was associated with species-rich but highly uneven communities and less influenced by compensatory dynamics. In conclusion, both specific traits (dominance) and diversity (richness) affected the stability of metacommunities under fluctuating conditions.     

Nearly a decade‐long repeatable seasonal diversity patterns of bacterioplankton communities in the eutrophic Lake Donghu (Wuhan,China)

Uncovering which environmental factors govern community diversity patterns and how ecological processes drive community turnover are key questions related to understand the community assembly. However, the ecological mechanisms regulating long‐term variations of bacterioplankton communities in lake ecosystems remain poorly understood. Here we present nearly a decade‐long study of bacterioplankton communities from the eutrophic Lake Donghu (Wuhan, China) using 16S rRNA gene amplicon sequencing with MiSeq platform. We found strong repeatable seasonal diversity patterns in terms of both common (detected in more than 50% samples) and dominant (relative abundance >1%) bacterial taxa turnover. Moreover, community composition tracked the seasonal temperature gradient, indicating that temperature is a key environmental factor controlling observed diversity patterns. Total phosphorus also contributed significantly to the seasonal shifts in bacterioplankton composition. However, any spatial pattern of bacterioplankton communities across the main lake areas within season was overwhelmed by their temporal variabilities. Phylogenetic analysis further indicated that 75%–82% of community turnover was governed by homogeneous selection due to consistent environmental conditions within seasons, suggesting that the microbial communities in Lake Donghu are mainly controlled by niche‐based processes. Therefore, dominant niches available within seasons might be occupied by similar combinations of bacterial taxa with modest dispersal rates throughout different lake areas.     

Dispersal scales up the biodiversity–productivity relationship in an experimental source-sink metacommunity

The influence of biodiversity on ecosystem functioning is a major concern of ecological research. However, the biodiversity–ecosystem functioning relationship has very often been studied independently from the mechanisms allowing coexistence. By considering the effects of dispersal and niche partitioning on diversity, the metacommunity perspective predicts a spatial scale-dependence of the shape of the relationship. Here, we present experimental evidence of such scale-dependent patterns. After approximately 500 generations of diversification in a spatially heterogeneous environment, we measured functional diversity (FD) and productivity at both local and regional scales in experimental source-sink metacommunities of the bacterium Pseudomonas fluorescens SBW25. At the regional scale, environmental heterogeneity yielded high levels of FD and we observed a positive correlation between diversity and productivity. At the local scale, intermediate dispersal increased local FD through a mass effect but there was no correlation between diversity and productivity. These experimental results underline the importance of considering the mechanisms maintaining biodiversity and the appropriate spatial scales in understanding its relationship with ecosystem functioning.     

Responses of epilimnetic phytoplankton to experimental nutrient enrichment in three small seepage lakes

This paper describes the responses of three epilimnetic phytoplanktoncommunities to experimental nitrogen and phosphorus enrichmentas compared to the phytoplankton community in a fourth, unmanipulated,lake. Increased nutrient inputs increased total phytoplanktonbiomass, primary productivity, chlorophytes, cryptomonads andspecies turnover rates in all three enriched lakes; cyanobacteriaincreased in two of the three enriched lakes. However, nutrientaddition also led to declines in previously dominant dinoflagellatesand chrysophytes, and in species diversity. At the species level,there were large changes in community composition from yearto year in both enriched and reference lakes, suggesting thatphytoplankton community composition is highly dynamic even inthe absence of enrichment. Overall, changes in total biomass,productivity and species diversity were consistent among theenriched lakes, while changes in species composition differeddue to variation in the physical, chemical and biotic environmentof each lake. This suggests that aggregated variates are moreuseful for quantitative prediction of nutrient effects, whilespecies responses can be used to signal qualitative differencesin environmental conditions among lakes. ³Present address: Department of Biological Sciences, DartmouthCollege, 6044 Gilman Laboratory, Hanover, NH 03755-3576, USA     

Connectivity, scale-dependence, and the productivity–diversity relationship

We surveyed freshwater ponds (localities) nested within watersheds (regions) to evaluate the relationship between productivity and animal species richness at different spatial scales. In watersheds where the ponds were relatively distant from one another (likely reducing the level of interpond dispersal of many organisms), we found a scale‐dependent productivity–diversity relationship; at local scales (among ponds), diversity was a hump‐shaped function of productivity, whereas at regional scales (among watersheds), diversity monotonically increased with productivity. Furthermore, this relationship emerged because there was a strong relationship between productivity and pond‐to‐pond species compositional differences. Alternatively, in watersheds where ponds were relatively close together (likely leading to higher rates of dispersal of many organisms), we found no scale‐dependence; diversity was a hump‐shaped function of productivity at both local and regional scales. Here, the relationship between species compositional dissimilarity and productivity was much weaker. We conclude that whether or not scale‐dependence is observed in productivity–diversity relationships will depend, at least in part, on the degree of connectivity among localities within regions.     

Dispersal ability links to cross‐scale species diversity patterns across the Eurasian Arctic tundra

Aim The role of dispersal in structuring biodiversity across spatial scales is controversial. If dispersal controls regional and local community assembly, it should also affect the degree of spatial species turnover as well as the extent to which regional communities are represented in local communities. Here we provide the first integrated assessment of relationships between dispersal ability and local‐to‐regional spatial aspects of species diversity across a large geographical area. Location Northern Eurasia. Methods Using a cross‐scale analysis covering local (0.64 m²) to continental (the Eurasian Arctic biome) scales, we compared slope parameters of the dissimilarity‐to‐distance relationship in species composition and the local‐to‐regional relationship in species richness among three plant‐like groups that differ in dispersal ability: lichens with the highest dispersal ability; mosses and moss allies with intermediate dispersal ability; and seed plants with the lowest dispersal ability. Results Diversity patterns generally differed between the three groups according to their dispersal ability, even after controlling for niche‐based processes. Increasing dispersal ability is linked to decreasing spatial species turnover and an increasing ratio of local to regional species richness. All comparisons supported our expectations, except for the slope of the local‐to‐regional relationship in species richness for mosses and moss allies which was not significantly steeper than that of seed plants. Main conclusions The negative link between dispersal ability and spatial species turnover and the corresponding positive link between dispersal ability and the ratio of local‐to‐regional species richness support the idea that dispersal affects community structure and diversity patterns across spatial scales.     

Warming induces synchrony and destabilizes experimental pond zooplankton metacommunities

The spatial insurance hypothesis predicts that intermediate rates of dispersal between patches in a metacommunity allow species to track favourable conditions, preserving diversity and stabilizing biomass at local and regional scales. However, theory is unclear as to whether dispersal will provide spatial insurance when environmental conditions are changing directionally. In particular, increased temperatures as a result of climate change are expected to cause synchronous growth or decline across species and communities, and this has the potential to erode the stabilizing compensatory dynamics facilitated by dispersal. Here we report on an experimental test of how dispersal affects the diversity and stability of metacommunities under warming using replicate two‐patch pond zooplankton metacommunities. Initial differences in local community composition and abiotic conditions were established by seeding each patch in the metacommunities with plankton and sediment from one of two natural ponds that differed in water chemistry and species composition. We exposed metacommunities to a 2°C increase in average ambient temperature, crossed with three rates of dispersal (none, intermediate, high). In ambient conditions, intermediate dispersal rates preserved diversity and stabilized metacommunities by promoting spatially asynchronous fluctuations in biomass, especially between local populations of the dominant genus, Ceriodaphnia. However, warming synchronized their populations so that these effects of dispersal were lost. Furthermore, because the stabilizing effect of dispersal was primarily due to asynchronous fluctuations between populations of a single genus, metacommunity biomass was stabilized, but dispersal did not stabilize local community biomass. Our results show that dispersal can preserve diversity and provide stability to metacommunities, but also show that this benefit can be eroded when warming is directional and synchronous across patches of a metacommunity, as is expected with climate warming.     

Spatial averaging and disturbance lead to high productivity in aquatic metacommunities

Dispersal in heterogeneous ecosystems, such as coastal metacommunities, is a major driver of diversity and productivity. According to theory, both species richness and spatial averaging shape a unimodal relationship of productivity with dispersal. We experimentally tested the hypothesis that disturbances acting on local patches would buffer the loss of productivity at high dispersal by preventing synchronized species oscillations. To simulate these disturbances, our experimental assemblages involved species that self‐organized in isolation under three inflow pulsing frequencies, where hydraulic displacement and nutrient loading affected assemblage diversity and composition. At steady‐state, the emerging isolated assemblages were connected at three levels of dispersal creating three metacommunities of different connectivity. Consistent with theory, as dispersal increased, species richness in the metacommunity declined; productivity however remained high. This occurred because the most productive species in our study (which dominated the isolated patch of intermediate inflow pulsing frequency) dominated all three patches (low, intermediate and high inflow pulsing frequencies) after dispersal commenced in our metacommunities. This experimental result provides empirical support for the mechanism of spatial averaging. Furthermore, disturbances, in the form of localized pulsed inflows, prevented population oscillation synchrony caused by homogenization. Overall, our observations suggest that localized environmental fluctuations and the identity of species seem to be more influential than dispersal in shaping the diversity and composition of phytoplankton assemblages and stabilizing productivity.     

Frog community composition-environment relationships vary over time: Are snapshot studies of metacommunity dynamics useful?

There have been important advances in understanding the relative importance of environmental and spatial processes for the variation in species composition across a set of local communities linked by dispersal (i.e. metacommunities). However, community composition-environment relationships change over time, and the mechanisms shaping such temporal variation in metacommunities encompassing large environmental gradients remain poorly understood. If the ability of statistical models to predict community composition-environment relationships depends on the sampling year, snapshot metacommunity studies would have limited implications, both theoretical and applied. Here, we partitioned the variation in compositional data of frog communities and asked if the relative importance of environmental and spatial components change over years at broad spatial scales (hereafter, protected areas in coastal and inland regions) of southeastern Brazil. These regions have marked differences in environmental characteristics as well as the size and composition of their regional species pool. Our results showed that the factors explaining the temporal variability in community composition-environment relationships were congruent for the inland region, which is less productive and characterized by harsh environmental conditions. In contrast, the relative importance of environmental and spatial components changed over years in the coastal region, which has more productive environments and benign conditions. Although snapshot studies will continue to provide important information about metacommunity dynamics, researchers have to be better able to incorporate the temporal variation inherent in community composition-environment relationships, which may be especially important in productive environments.     

Environmental heterogeneity among lakes promotes hyper β‐diversity across phytoplankton communities

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Predators temper the relative importance of stochastic processes in the assembly of prey metacommunities

Communities assemble through a combination of stochastic processes, which can make environmentally similar communities divergent (high β-diversity), and deterministic processes, which can make environmentally similar communities convergent (low β-diversity). Top predators can influence both stochasticity (e.g. colonization and extinction events) and determinism (e.g. size of the realized species pool), in community assembly, and thus their net effect is unknown. We investigated how predatory fish influenced the scaling of prey diversity in ponds at local and regional spatial scales. While fish reduced both local and regional richness, their effects were markedly more intense at the regional scale. Underlying this result was that the presence of fish made localities within metacommunities more similar in their community composition (lower β-diversity), suggesting that fish enhance the deterministic, relative to the stochastic, components of community assembly. Thus, the presence of predators can alter fundamental mechanisms of community assembly and the scaling of diversity within metacommunities.