Metacommunity data and theory: the tortoise and the hare

Empirical approaches to metacommunities: a review and comparison with theory
Logue et al. 2011

The recognition that community composition is a function of both local and regional-scale processes, meaning that a community cannot be understood in isolation from the network of communities with which it interacts, is the fundamental idea behind metacommunity ecology. In a relatively short period of time, metacommunity ecology has integrated concepts from spatial ecology, metapopulation ecology, and community ecology with novel ideas, and developed a strong body of theory. However, metacommunity theory has advanced much more rapidly than empirical tests of that theory. In an interesting review in TREE, Logue et al. examine whether empirical data needs to catch up with the pace of theory development, or whether theory is moving too fast to incorporate the information available from empirical data.

The types of systems used in the 34 experimental and 74 observational studies that Logue et al. found were very limited – the most common experimental approach involved setting up aquatic microcosms of unicellular organisms.* Observational studies similarly tested microorganisms, usually in aquatic systems. The organisms so beloved in the rest of community ecology (plants? vertebrates?) barely feature. Most studies focus on aquatic systems composed of multiple patches (such as microcosms, ponds, pitcher plant communities) because systems with discrete boundaries are more amenable to testing current theory. However, natural systems are rarely configured into a clear “patch” versus “matrix” dichotomy. Instead they are complex and heterogeneous, and may lack clear boundaries.

Dynamics in metacommunities are generally described using four dominant paradigms: mass-effects, species sorting, neutral perspective, or patch-dynamics. These paradigms reflect the most important processes structuring communities, that is, either dispersal between communities, environmental differences between communities, dynamics driven by the tenets of neutral theory, or extinction and colonization, respectively. Strikingly, experimental studies mostly tested for mass-effects or patch dynamics, and observational studies mostly tested for species-sorting and mass effects paradigms. The neutral paradigm was rarely tested in any type of study. Logue et al. found that many studies had difficulty designing experiments that tested for evidence of specific paradigms, because natural communities are much more complex than the simple paradigms suggest. Most studies that did test for evidence of particular paradigms found evidence for multiple paradigms or had difficulty disentangling different mechanisms.

The metacommunity theory that has developed in the last five years is among the most exciting and interesting work in ecology. However, the slower pace of experimental work means that theory has developed with little feedback. For example, Logue et al. make a strong argument that the results from these studies suggest that it is time to integrate the four-paradigm system into a single, comprehensive framework (see figure). Theory is only valuable if it’s useful - this paper is an important reminder that there is an important feedback loop between theory and data, and successful science requires input from both.

*Important disclaimer: at this very moment I'm running aquatic microcosms of microscopic protists in the lab. We all have room for improvment. :)

Don’t miss the mechanism when testing for biodiversity effects

Variation in the strength of diversity effects among experimental studies raise the question when and where consequences of diversity loss is strongest. As in grassland experiments, diversity effects on plant biomass production can be observed in systems with marine macroalgae. However, even among marine macroalgae experiments variation in the strength of the diversity effect cannot be explained because of largely differing experimental set-ups (i.e. long-termed vs. short-termed studies, mesocosms vs. field experiments, using inter- or subtidal habitats). From literature Stachowicz et al. assumed that short termed factors regulating diversity effects in such systems could be attributed to spatial complementarity in photosynthesis rates or different limiting nutrients. Long-term regulating factors could be attributed to habitat differentiation, temporal complementarity, fascilitation, recruitment and natural heterogeneity of substrate. In a very elegant way Stachowicz and his co-workers tested whether mechanisms responsible for diversity effects change with experimental procedure and/or study type within the same marine algae system. In a series of three experiments, that is a short-termed mesocosm with transplanted thalli, a short-termed (two month) field-experiment with naturally recruited thalli and heterogeneous substrate, and in a long-term (three years) field-experiment, the authors were able to show that strong diversity effects are positively correlated with experimental duration, environmental heterogeneity and population responses (recruitment). Whereas in the mesocosm species identity affected biomass production, in the field studies it was species richness but not identity. Fractional change of species biomass could be explained by species identity in the mesocosm, and by both identity and richness in the field. The authors are making an important point by showing that mechanisms for diversity effects are not exclusive but occur together and become stronger over time. They conclude that the absence or the detection of only weak diversity effects in short-termed experiments does not necessarily mean that there is no effect because such approaches detect only a limited number of potential mechanisms.


John J. Stachowicz, Rebecca J. Best, Matthew E. S. Bracken, Michael H. Graham (2008). Complementarity in marine biodiversity manipulations: Reconciling divergent evidence from field and mesocosm experiments. Proceedings of the National Academy of Sciences DOI: 10.1073/pnas.0806425105

Small experimental plots predict entire ecosystem responses! (if you work in peatlands…)

The possibility of extrapolating results from experimental plots to larger (or “real”) scales is a major issue in ecology. For several reasons ecologists conduct manipulative experiments in relatively small experimental units. This that has been suggested to be a big problem since the effect of the studied factor could change with spatial scale. An example of this can be found in biological invasions where there is some evidence that the more species you have at a small scale (e.g. a plot), the less likely an exotic can invade; but, at the regional level, the more species there are, the more likely that exotics can invade, so invasion has a scale-dependent response to species richness. However, if you work on peatlands you are very lucky! A recent paper by Magdalena Wiedermann and collaborators found that in peatlands, experiments in 2 x 2 meter plots represented really well what was happening at the entire ecosystem level. They compared a manipulative experiment where they added nitrogen at different concentrations, with an observational study in a region with gradient of nitrogen concentrations similar to the ones used in the experiment. They found that cover of Sphagnum and vascular plants could be explained by the levels of nitrogen equally well at plot and regional scales

Magdalena M. Wiedermann, Urban Gunnarsson, Mats B. Nilsson, Annika Nordin, Lars Ericson (2009). Can small-scale experiments predict ecosystem responses? An example from peatlands Oikos DOI: 10.1111/j.1600-0706.2008.17129.x