The importance of metacommunity ecology for environmental assessment research in the freshwater realm

Most bioassessment programs rest on the assumption that species have different niches, and that abiotic environmental conditions and changes therein determine community structure. This assumption is thus equivalent to the species sorting perspective (i.e. that species differ in their responses to environmental variation) in metacommunity ecology. The degree to which basing bioassessment on the species sorting perspective is reasonable is likely to be related to the spatial extent of a study and the characteristics of the organism groups (e.g. dispersal ability) with which the effects of anthropogenic changes are assessed. Recent findings in metacommunity research have stressed that community structure is determined not only by local abiotic environmental conditions but also by biotic interactions and dispersal‐related effects. For example, dispersal limitation may prevent community structure recovery from the effects of a putative stressor, as organisms may not be able to disperse to all sites in a region. Mass effects (i.e. the presence of species in environmentally suboptimal sites due to high dispersal rates from environmentally suitable sites) may, in turn, obscure the effects of a stressor, as dispersal from source sites (e.g. an unaltered site) allows persistence at sink sites (e.g. an anthropogenically altered site). Better bioassessment should thus take both niche‐ and dispersal‐related processes simultaneously into consideration, which can be accomplished by explicitly modelling spatial location as a proxy for dispersal effects. Such an integrated approach should be included in bioassessment programs using general multivariate approaches, predictive modelling, and multimetric indices.     

Integrating environmental and spatial processes in ecological community dynamics

The processes controlling the abundances of species across multiple sites form the cornerstone of modern ecology. In these metacommunities, the relative importance of local environmental and regional spatial processes is currently hotly debated, especially in terms of the validity of neutral model. I collected 158 published data sets with information on community structure, environmental and spatial variables. I showed that approximately 50% of the variation in community composition is explained by both environmental and spatial variables. The majority of the data sets were structured by species-sorting dynamics (SS), followed by a combination of SS and mass-effect dynamics. While neutral processes were the only structuring process in 8% of the collected natural communities, disregarding neutral dispersal processes would result in missing important patterns in 37% of the studied communities. Moreover, metacommunity characteristics such as dispersal type, habitat type and spatial scale predicted part of the detected variation in metacommunity structure.     

Meta-ecosystems: a theoretical framework for a spatial ecosystem ecology

This contribution proposes the meta‐ecosystem concept as a natural extension of the metapopulation and metacommunity concepts. A meta‐ecosystem is defined as a set of ecosystems connected by spatial flows of energy, materials and organisms across ecosystem boundaries. This concept provides a powerful theoretical tool to understand the emergent properties that arise from spatial coupling of local ecosystems, such as global source–sink constraints, diversity–productivity patterns, stabilization of ecosystem processes and indirect interactions at landscape or regional scales. The meta‐ecosystem perspective thereby has the potential to integrate the perspectives of community and landscape ecology, to provide novel fundamental insights into the dynamics and functioning of ecosystems from local to global scales, and to increase our ability to predict the consequences of land‐use changes on biodiversity and the provision of ecosystem services to human societies.     

Temporal dynamics of a local fish community are strongly affected by immigration from the surrounding metacommunity

A 5‐year time series of annual censuses was collected from a large floodplain lake to determine how dynamics of the local fish community were affected by changes in hydrological connectivity with the surrounding metacommunity. The lake was disconnected from the metacommunity for 1 year prior to our study and remained disconnected until 3 months before our third annual census, when a flood reconnected the lake to the metacommunity. We determined how changes in connectivity affected temporal dynamics of (1) local community composition and (2) the population composition, condition, and growth of catfish, to shed light on how immigration of other species might affect local population dynamics. Before reconnection, the community was likely shaped by interactions between the local environment and species traits. The reconnection caused significant immigration and change in community composition and correlated with a significant and abrupt decline in catfish condition, growth, and abundance; effects likely due to the immigration of a competitor with a similar trophic niche: carp. The community was slow to return to its preconnection state, which may be due to dispersal traits of the fishes, and a time‐lag in the recovery of the local catfish population following transient intensification of species interactions. The dynamics observed were concordant with the species sorting and mass‐effects perspectives of metacommunity theory. Floods cause episodic dispersal in floodplain fish metacommunities, and so, flood frequency determines the relative importance of regional and local processes. Local processes may be particularly important to certain species, but these species may need sufficient time between floods for population increase, before the next flood‐induced dispersal episode brings competitors and predators that might cause population decline. Accordingly, species coexistence in these metacommunities may be facilitated by spatiotemporal storage effects, which may in turn be regulated by flood frequency.     

The influence of niche and neutral processes on a neotropical anuran metacommunity

One of the most important questions in ecology is the relative importance of local conditions (niche processes) and dispersal ability (neutral processes) in driving metacommunity structure. Although many studies have been conducted in recent years, there is still much debate. We evaluated the processes (niche and neutral) responsible for variation in anuran composition in 28 lentic water bodies in southeastern Brazil. Because anurans depend heavily on environmental conditions, we hypothesized that environmental variables (niche processes) are the most important drivers of community composition. Additionally, as anurans have limited dispersal abilities, and the study region presents harsh conditions (high forest fragmentation, low rainfall and long dry season), we expected a lower, but significant, spatial signature in metacommunity structure, due to neutral dynamics. We used a partial redundancy analysis with variation partitioning to evaluate the relative influence of environmental and spatial variables as drivers of metacommunity structure. Additionally, we used a recently developed spatial autocorrelation analysis to test if neutral dynamics can be attributed to the pure spatial component. This analysis is based on predictions that species abundances are independent but similarly spatially structured, with correlograms similar in shape. Therefore, under neutral dynamics there is no expectation of a correlation between the pairwise distance of spatial correlograms and the pairwise correlation of species abundances predicted by the pure spatial component. We found that the environmental component explained 21.5%, the spatial component 10.2%, and the shared component 6.4% of the metacommunity structure. We found no correlation between correlograms and correlation of abundances predicted by the pure spatial component (Mantel test = ?0.109, P = 0.961). In our study, niche‐based processes are the dominant process that explained community composition. However, neutral processes are important because spatial variation can be attributed to pure neutral dynamics rather than to missing spatially structured environmental factors.     

Spatial and environmental effects on a rock‐pool metacommunity depend on landscape setting and dispersal mode

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Mechanisms determining the fate of dispersed bacterial communities in new environments

Recent work has shown that dispersal has an important role in shaping microbial communities. However, little is known about how dispersed bacteria cope with new environmental conditions and how they compete with local resident communities. To test this, we implemented two full-factorial transplant experiments with bacterial communities originating from two sources (freshwater or saline water), which were incubated, separately or in mixes, under both environmental conditions. Thus, we were able to separately test for the effects of the new environment with and without interactions with local communities. We determined community composition using 454-pyrosequencing of bacterial 16S rRNA to specifically target the active fraction of the communities, and measured several functional parameters. In absence of a local resident community, the net functional response was mainly affected by the environmental conditions, suggesting successful functional adaptation to the new environmental conditions. Community composition was influenced both by the source and the incubation environment, suggesting simultaneous effects of species sorting and functional plasticity. In presence of a local resident community, functional parameters were higher compared with those expected from proportional mixes of the unmixed communities in three out of four cases. This was accompanied by an increase in the relative abundance of generalists, suggesting that competitive interactions among local and immigrant taxa could explain the observed ‘functional overachievement''. In summary, our results suggest that environmental filtering, functional plasticity and competition are all important mechanisms influencing the fate of dispersed communities.     

Towards a multi‐trophic extension of metacommunity ecology

Metacommunity theory provides an understanding of how spatial processes determine the structure and function of communities at local and regional scales. Although metacommunity theory has considered trophic dynamics in the past, it has been performed idiosyncratically with a wide selection of possible dynamics. Trophic metacommunity theory needs a synthesis of a few influential axis to simplify future predictions and tests. We propose an extension of metacommunity ecology that addresses these shortcomings by incorporating variability among trophic levels in ‘spatial use properties’. We define ‘spatial use properties’ as a set of traits (dispersal, migration, foraging and spatial information processing) that set the spatial and temporal scales of organismal movement, and thus scales of interspecific interactions. Progress towards a synthetic predictive framework can be made by (1) documenting patterns of spatial use properties in natural food webs and (2) using theory and experiments to test how trophic structure in spatial use properties affects metacommunity dynamics.     

The challenges that spatial context present for synthesizing community ecology across scales

Accurately characterizing spatial patterns on landscapes is necessary to understand the processes that generate biodiversity, a problem that has applications in ecological theory, conservation planning, ecosystem restoration, and ecosystem management. However, the measurement of biodiversity patterns and the ecological and evolutionary processes that underlie those patterns is highly dependent on the study unit size, boundary placement, and number of observations. These issues, together known as the modifiable areal unit problem, are well known in geography. These factors limit the degree to which results from different metacommunity and macro‐ecological studies can be compared to draw new inferences, and yet these types of comparisons are widespread in community ecology. Using aquatic community datasets, we demonstrate that spatial context drives analytical results when landscapes are sub‐divided. Next, we present a framework for using resampling and neighborhood smoothing to standardize datasets to allow for inferential comparisons. We then provide examples for how addressing these issues enhances our ability to understand the processes shaping ecological communities at landscape scales and allows for informative meta‐analytical synthesis. We conclude by calling for greater recognition of issues derived from the modifiable areal unit problem in community ecology, discuss implications of the problem for interpreting the existing literature, and identify tools and approaches for future research.     

The role of recruitment and dispersal limitation in tree community assembly in Amazonian forests

Background: Species composition of plant communities is shaped by the interplay between dispersal limitation, environmental filters and stochastic events.

Aims: The aim of this work was to investigate the effects of dispersal limitation and environmental filtering on tree recruitment. To accomplish this, we employed the unified neutral theory of biodiversity and biogeography to examine migration within the metacommunity, defined as a set of interacting local communities linked by the dispersal of multiple potentially interacting species.

Methods: We sampled 12,975 individuals with dbh ≥ 1 cm in 26 1-ha permanent plots, including habitats of terra firme, transitional forests, várzea and campinarana, on the upper Madeira River, Brazilian Amazon.

Results: Campinarana drew individuals from outside the metacommunity species pool at a mean probability of recruitment of 0.06, a much lower probability than terra firme (0.31), transitional (0.21) and várzea forests (0.22). Environmental variables, such as water table depth, soil texture and fertility, were related to differences in community assembly.

Conclusions: Species abundance distribution and diversity patterns of plant assemblages in a large river landscape in the Amazon highlight the importance of environmental heterogeneity that conditions beta-diversity. The high variation in recruitment probabilities from the metacommunity species pool to local communities suggests high habitat variability in the process of maintaining patterns of local diversity.     

The relative importance of dispersal and local processes in structuring phytoplankton communities in a set of highly interconnected ponds

1. The recognition that both local and regional processes act together in shaping local communities makes determining their relative roles in natural communities central to understanding patterns in community structure. 2. We investigated the relative influence of these processes on the phytoplankton communities of a highly interconnected pond system. We sampled the phytoplankton communities of 28 ponds concurrently with 20 local environmental variables. 3. We found that phytoplankton community variation, in terms of both phytoplankton community composition (PCC) and diversity, was only significantly explained by local environmental variables. These were mainly associated with the contrasting clear‐water and turbid ecological states of the shallow ponds studied. Clear‐water conditions favoured only a few taxa, resulting in a significantly lower taxon diversity and richness under these conditions. 4. The failure to explain variation in PCC by a dispersal model based on the water flow between ponds points at very effective species sorting. This is attributed to the high population turn‐over rates and sensitivity to environmental conditions of phytoplankton communities. Some evidence was found, however, that dispersal influences local communities through mass effects between neighbouring ponds. 5. Overall, our results emphasize both the strong selection pressure that components of the food web exert on phytoplankton communities and the high potential of these communities to respond to such environmental change, thereby effectively opposing the homogenizing effects of continuous dispersal.     

A general framework for neutral models of community dynamics

Neutral models of community dynamics are a powerful tool for ecological research, but their applications are currently limited to unrealistically simple types of dynamics and ignore much of the complexity that characterize natural ecosystems. Here, we present a new analytical framework for neutral models that unifies existing models of neutral communities and extends the applicability of existing models to a much wider spectrum of ecological phenomena. The new framework extends the concept of neutrality to fitness equivalence and in spite of its simplicity explains a wide spectrum of empirical patterns of species diversity including positive, negative and unimodal productivity–diversity relationships; gradual and highly delayed declines in species diversity with habitat loss; and positive and negative responses of species diversity to habitat heterogeneity. Surprisingly, the abundance distribution in all of these cases is given by the dispersal limited multinomial (DLM), the abundance distribution in Hubbell's zero-sum model, showing DLM's robustness and demonstrating that it cannot be used to infer the underlying community dynamics. These results support the hypothesis that ecological communities are regulated by a limited set of fundamental mechanisms much simpler than could be expected from their immense complexity.     

Trade-offs in community ecology: linking spatial scales and species coexistence

Trade‐offs in species performances of different ecological functions is one of the most common explanations for coexistence in communities. Despite the potential for species coexistence occurring at local or regional spatial scales, trade‐offs are typically approached at a single scale. In recent years, ecologists have increasingly provided evidence for the importance of community processes at both local and regional spatial scales. This review summarizes the theoretical predictions for the traits associated with trade‐offs under different conditions and at different spatial scales. We provide a spatial framework for understanding trade‐offs, coexistence and the supportive empirical evidence. Predictions are presented that link the patterns of diversity observed to the patterns of trade‐offs that lead to coexistence at different spatial scales. Recent evidence for the evolution of trade‐offs under different conditions is provided which explores both laboratory microcosm studies and phylogenetic tests. Examining trade‐offs within a spatial framework can provide a strong approach to understanding community structure and dynamics, while explaining patterns of species diversity.     

The community ecology of pathogens: coinfection,coexistence and community composition

Disease and community ecology share conceptual and theoretical lineages, and there has been a resurgence of interest in strengthening links between these fields. Building on recent syntheses focused on the effects of host community composition on single pathogen systems, we examine pathogen (microparasite) communities using a stochastic metacommunity model as a starting point to bridge community and disease ecology perspectives. Such models incorporate the effects of core community processes, such as ecological drift, selection and dispersal, but have not been extended to incorporate host–pathogen interactions, such as immunosuppression or synergistic mortality, that are central to disease ecology. We use a two‐pathogen susceptible‐infected (SI) model to fill these gaps in the metacommunity approach; however, SI models can be intractable for examining species‐diverse, spatially structured systems. By placing disease into a framework developed for community ecology, our synthesis highlights areas ripe for progress, including a theoretical framework that incorporates host dynamics, spatial structuring and evolutionary processes, as well as the data needed to test the predictions of such a model. Our synthesis points the way for this framework and demonstrates that a deeper understanding of pathogen community dynamics will emerge from approaches working at the interface of disease and community ecology.     

集合群落及其在河流鱼类群聚研究中的应用

集合群落(metacommunity)是指多个潜在相互作用的物种通过它们之间的扩散而连接在一起的一组局域群落,目前已成为斑块生境下生物群落结构、格局和动态的重要理论基础之一。斑块动态、物种排序、群体效应和中性模型等4种理论模型,可用于解释不同情形下集合群落内物种的迁移状况,描述集合群落的动态。可采用群落结构或生态学机制等途径,来阐述所研究的群落是属于哪一种特定的集合群落类型。集合群落可用于研究河流鱼类群聚,解释鱼类的群落结构等问题。另外本文还结合我国水域生态环境及水生生物现状,对今后集合群落的研究作了展望。