- Evolution et Diversité Biologique lab, University of Toulouse III, Toulouse, France
- Biodiversity, Biogeography, Climate change, Community ecology, Conservation biology, Evolutionary ecology, Human impact, Macroecology, Meta-analyses, Spatial ecology, Metacommunities & Metapopulations, Species distributions, Statistical ecology
Community size affects the signals of ecological drift and selection on biodiversity
Toward an empirical synthesis on the niche versus stochastic debateRecommended by Eric Harvey based on reviews by Kevin Cazelles and Romain Bertrand
As far back as Clements  and Gleason , the historical schism between deterministic and stochastic perspectives has divided ecologists. Deterministic theories tend to emphasize niche-based processes such as environmental filtering and species interactions as the main drivers of species distribution in nature, while stochastic theories mainly focus on chance colonization, random extinctions and ecological drift . Although the old days when ecologists were fighting fiercely over null models and their adequacy to capture niche-based processes is over , the ghost of that debate between deterministic and stochastic perspectives came back to haunt ecologists in the form of the ‘environment versus space’ debate with the development of metacommunity theory . While interest in that question led to meaningful syntheses of metacommunity dynamics in natural systems , it also illustrated how context-dependant the answer was . One of the next frontiers in metacommunity ecology is to identify the underlying drivers of this observed context-dependency in the relative importance of ecological processus [7, 8].
Reflecting on seminal work by Robert MacArthur emphasizing different processes at different spatial scales [9, 10] (the so-called ‘MacArthur paradox’), Chase and Myers proposed in 2011 that a key in solving the deterministic versus stochastic debate was probably to turn our attention to how the relative importance of local processes changes across spatial scales . Scale-dependance is a well-acknowledged challenge in ecology, hampering empirical syntheses and comparisons between studies [11-14]. Embracing the scale-dependance of ecological processes would not only lead to stronger syntheses and consolidation of current knowledge, it could also help resolve many current debates or apparent contradictions [11, 15, 16].
The timely study by Siqueira et al.  fits well within this historical context by exploring the relative importance of ecological drift and selection across a gradient of community size (number of individuals in a given community). More specifically, they tested the hypothesis that small communities are more dissimilar among each other because of ecological drift compared to large communities, which are mainly structured by niche selection . That smaller populations or communities should be more affected by drift is a mathematical given , but the main questions are i) for a given community size how important is ecological drift relative to other processes, and ii) how small does a community have to be before random assembly dominates? The authors answer these questions using an extensive stream dataset with a community size gradient sampled from 200 streams in two climatic regions (Brazil and Finland). Combining linear models with recent null model approaches to measure deviations from random expectations , they show that, as expected based on theory and recent experimental work, smaller communities tend to have higher β-diversity, and that those β-diversity patterns could not be distinguished from random assembly processes . Spatial turnover among larger communities is mainly driven by niche-based processes related to species sorting or dispersal dynamics . Given the current environmental context, with many anthropogenic perturbations leading to reduced community size, it is legitimate to wonder, as the authors do, whether we are moving toward a more stochastic and thus less predictable world with obvious implications for the conservation of biodiversity .
The real strength of the study by Siqueira et al. , in my opinion, is in the inclusion of stream data from boreal and tropical regions. Interestingly and most importantly, the largest communities in the tropical streams are as large as the smallest communities in the boreal streams. This is where the study should really have us reflect on the notions of context-dependency in observed patterns because the negative relationship between community size and β-diversity was only observed in the tropical streams, but not in the boreal streams . This interesting nonlinearity in the response means that a study that would have investigated the drift versus niche-based question only in Finland would have found very different results from the same study in Brazil. Only by integrating such a large scale gradient of community sizes together could the authors show the actual shape of the relationship, which is the first step toward building a comprehensive synthesis on a debate that has challenged ecologists for almost a century.
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