Review
Empirical approaches to metacommunities: a review and comparison with theory

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Metacommunity theory has advanced understanding of how spatial dynamics and local interactions shape community structure and biodiversity. Here, we review empirical approaches to metacommunities, both observational and experimental, pertaining to how well they relate to and test theoretical metacommunity paradigms and how well they capture the realities of natural ecosystems. First, we show that the species-sorting and mass-effects paradigms are the most commonly tested and supported paradigms. Second, the dynamics observed can often be ascribed to two or more of the four non-exclusive paradigms. Third, empirical approaches relate only weakly to the concise assumptions and predictions made by the paradigms. Consequently, we suggest major avenues of improvement for empirical metacommunity approaches, including the integration across theoretical approaches and the incorporation of evolutionary and meta-ecosystem dynamics. We hope for metacommunity ecology to thereby bridge existing gaps between empirical and theoretical work, thus becoming a more powerful framework to understand dynamics across ecosystems.

Section snippets

Metacommunity theory: a mechanistic approach towards the understanding of local community dynamics

Understanding the mechanisms that underlie patterns of species distribution, abundance and interactions is central to community ecology. Traditionally, community ecology focused on either local processes [1] or dynamics at the regional scale 2, 3. The acknowledgement that community composition within a local habitat is affected by both local interactions and regional processes is one of the major achievements in community ecology within the past 50 years.

This idea of processes at different

Experimental approaches

Experimental studies were diverse in their approaches towards metacommunities (Box 2). The majority of the 34 experimental studies related to aquatic environments (Figure 2a). Some mimicked certain marine 8, 9 or freshwater 10, 11 habitat types, whereas others established artificial aquatic microcosms as model communities without referring to a specific type of aquatic system 12, 13. The latter approach was most common (see category ‘none’ in Figure 2a) and corresponded with the preponderance

Conformity and coherence between the reviewed empirical studies and the four paradigms of metacommunity theory

The two key publications that have synthesised approaches to metacommunity theory 5, 6 have provided the theoretical foundation for most of the empirical work reviewed here. The organisation of approaches to metacommunity theory into four paradigms has been both implicitly and explicitly implemented by the empirical studies analysed. This has, however, led to the introduction of a certain degree of incongruence between theoretical and empirical metacommunity research. First, processes of

Linking empirical metacommunity literature with processes in nature

Applying metacommunity paradigms to nature, which is not only patchy and heterogeneous [41] but also stochastic [34], is not straightforward [5]. Different organisms respond differently to processes (e.g. at different scales) and local communities often lack discrete boundaries, rendering a direct implemenation of simple theoretical approaches to natural systems difficult. With regard to the latter, most empirical studies addressed permanent habitat patches with discrete boundaries and focused

Need for integration

The original framework 5, 6 provided four main paradigms to metacommunity ecology that are well delineated and separate the main mechanisms of community assembly from a theoretical point of view (Box 1). Yet, as our review illustrates, this classification is not directly operational because it is difficult to link mechanisms to a single paradigm and because natural assemblages are structured by a combination of processes ascribed to different paradigms. In an attempt to reflect processes in

Need for empirical advancements

The metacommunity concept is a recent addition to community ecology theory. Progress in understanding factors and processes that structure metacommunities has been made beyond doubt. However, we have identified a series of areas within empirical work that need attention and, thus, hope that our review will initiate developments in empirical metacommunity research.

First, the types of habitat and groups of organism focused on in empirical approaches need to be extended. This necessity refers not

Conclusions

Metacommunity theory has enabled a big leap forward in understanding mechanisms of spatial community ecology. We have highlighted this in our review of empirical approaches and their subsequent comparison with metacommunity theory. We have nonetheless revealed major gaps in approaches to metacommunities and discrepancies between theory and empirical approaches. These limitations led to concise recommendations for future metacommunity studies. First, we suggest the extension of empirical

Acknowledgements

We are grateful to Eva S. Lindström and Silke Langenheder for organising a workshop on metacommunity ecology, held in Uppsala in 2009. Special thanks go to Eva, Silke and Yvonne Meyer-Lucht for constructive criticism on an earlier version of the manuscript. We thank Christine Meynard for help with Box 3. We further thank Marc Cadotte, Kendi Davies, Lars Gamfeldt, Andrew Gonzalez, Birte Matthiessen, Luc de Meester, Pablo Munguia and Patrick Venail for kindly supplying us with pictures of the

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