Evidence for the evolution of limiting similarity in diving beetle communities

Remi Vergnon, Remko Leijs, Egbert H. van Nes, and Marten Scheffer. (2013) Repeated Parallel Evolution Reveals Limiting Similarity in Subterranean Diving Beetles. The American Naturalist, Vol. 182, pp. 67-75.

In 2006, Marten Scheffer and Egbert van Nes published a very nice paper showing the outcome of simulated evolution of competing species. Their results showed how patterns of evenly-spaced clusters of species along a niche axis could evolve to minimize competition via limiting similarity. 

From Scheffer and van Nes (2006): Evenly spaced clusters of species along a niche axis (x-axis) evolved in response to competition.

Within any cluster along the niche axis, species tended to be more similar than expected. The results suggested that complex self-organizing clustered patterns might result from simple competitive limitations. Interestingly, although the original paper suggested that clustered patterns in size distributions are common, only now are these theoretical expectations about the evolution of limiting similarity being tested with data. In fact, though theory has long suggested that patterns of limiting similarity should evolve to allow coexistence between competing species, empirical evidence is rather lacking. Despite this, limiting similarity and competition are staples of ecological thought: for example, patterns of overdispersion in traits or relatedness are often used as evidence for the importance of competition.

The follow-up paper -Vergnon et al. (2013)- tests for the pattern predicted in Scheffer and van Nes (2006) using communities of subterranean diving beetles (Coleoptera, Dytiscidae) in Australia. These species have evolved for over 5 million years in isolated aquifers. If limiting similarity structured beetle communities, the authors predicted that there should be regularity in the spacing of species along a niche axis. If competitive interactions determine species' positions on the niche axis, then their absolute positions on the niche axis could vary between communities so long as their relative positions are evenly spaced. If, in contrast, niches are driven by environmental conditions, species in different communities/aquifers should have similar absolute positions along the niche axis.

The authors used a nice combination of statistics, modelling and observational data (34 communities of beetles representing 75 total species) to test for these predicted patterns. They used beetle size as the measure of niche position, since size is often an indicator of niche position and food availability and identity. For almost all aquifers, co-occurring beetles were significantly different in size. Further, species in different aquifers classified as occurring in the same size classes (small, medium, large), had different absolute sizes (i.e. the largest beetle in one 2-species aquifer was not similar in size to the largest beetle in another 2-species aquifer).  

From Vergnon et al. (2013): Absolute sizes of diving beetles in aquifers with 3 species present. The absolute size in a size class (large - black; medium - white; small - grey) varies between aquifers.

Although the absolute size of species differed between aquifers, the ratio of sizes (regularity of spacing on the niche axis) was highly consistent. Further, simulations of evolution of body size due to competition were capable of reproducing the observed size structure of the diving beetles.

From Vergnon et al. (2013): regularity of spacing between competing diving beetles (measured as the body size ratio). 

This paper does a nice job of integrating theory and data, and combining pattern and process. The focus is on testing contrasting predictions, and the authors use complementary approaches to test statistically for the presence of patterns and to demonstrate with simulations the relationship between the evolution of limiting similarity and the observed pattern. The evidence is suggestive that limiting similarity and not pre-existing environmental niches explains the size structure of communities of competing diving beetles. There are still questions about how far these inferences can be extended. For example, do we expect that predefined environmental niches are really the same across aquifers? How important is competition in these communities - at the moment, the authors only have minimal evidence of gut content overlap from a single aquifer. Further the low diversity of aquifer communities (~1-5 diving beetle species) means that the prediction of clusters of multiple similar species made in the original Scheffer and van Nes paper can't be tested. But the fact that aquifer diving beetle communities have low diversity and are very simplistic is beneficial for the authors. Patterns in diverse communities where multiple processes (predation, migration, etc) are important may be too complex to show clear evidence in observational data. Simple systems (including microcosms) are a good place to find evidence that a process of interest actually occurs. Whether or not that process is important across many systems is of course a more difficult question to answer.