Niche or Neutral? Why size matters.

Metacommunity dynamics (i.e. the effects of dispersal among connected communities) have become an increasingly common lens through which to explain community structure. For example, competition-colonization models explain the coexistence of superior and inferior competitors as the result of a trade-off in colonization and competitive ability. Species are either superior competitors, with high probabilities of establishing in patches, but low ability to move between patches, or superior colonizers, which have tend to lose in competitive interactions but can travel easily between patches. Under this framework, the ability of superior colonizers to reach and maintain populations in patches where their superior competitors are absent allows them to avoid extinction.

One problem with these types of models is that they rarely acknowledge the importance of ecological drift – that is, that chance events also affect species interactions. This despite the fact that we know that in “real life”, chance events likely play a major role in producing assemblages different than those we might predict based on theory. One of the strengths of the Hubbell’s neutral model is that it recognizes and embraces the importance of randomness.

A recent paper by Orrock and Watling (2010) examines how chance events can alter the predictions of the classic competition-colonization model. Orrock and Watling show that the size of communities in a metacommunity (which is assumed to correlate with the strength of ecological drift) determines whether community dynamics are niche-structured or neutral in nature. In large communities, predictions agree closely with those of the classic competition-colonization model, and niche-based interactions (i.e. competitive hierarchies) dominate. It’s in small communities that things get interesting: ecological drift becomes more important, so that differences in competitive ability between species are effectively neutralized. As a result, small communities begin to resemble neutral assemblages in which species abundances don’t relate to differences in competitive ability. An interesting consequence of this outcome is that species who are poor competitors but good colonizers have an additional refuge – simply by escaping to small communities, even if these communities contain superior competitors, they can persist in a metacommunity.

Beyond the theoretical implications of this model, the applied implications are what really matter. Habitat destruction and fragmentation are an growing problem due to human activities. Habitat patches are often smaller, and of lower quality, decreasing the size of the community each patch can support. Even if these patches are still connected and functioning as a metacommunity, species which rely on their strong competitive ability for persistence will lose this advantage as assemblages become increasingly neutral. Under this model, community diversity declines even more as habitat is lost than in the traditional competition-colonization model, and superior competitors face even greater extinction risk than previously predicted.

Since in reality, metacommunities are likely to consist of patches of different sizes, rather than all large or all small patches, the predictions here remain to be extended to more realistic metacommunities. However, Orrock and Watling have produced a useful model for understanding how ecological drift can affect diversity in a metacommunity and alter the expectations of traditional competition-colonization models.


Orrock, J.L. and Watling, J.I. (2010) Local community size mediates ecological drift and competition in metacommunities. Proc. R. Soc. B.

Being a clover isn’t always so lucky

Happy St. Patrick’s Day! I thought that covering an article about Trifolium (clover) seemed very appropriate. In a recent paper, Matthias Schleuning and colleagues examine the population dynamics of Trifolium montanum, a species in decline in central Germany. They examined the relative threats of habitat fragmentation and degradation on T. montanum’s population dynamics. They found that both degradation and fragmentation were having serious negative impacts. Degraded habitats in this system mean the shift away from nutrient-poor conditions and include the invasion of taller species that are better light competitors. T. montanum is a poor light competitor and maintains larger populations in mown or grazed habitats that keep taller invaders out. This species also faces the double whammy of fragmented habitats resulting in isolated populations. These isolates have lower reproductive output likely due to greater inbreeding and less genetic transfer, via pollinators, among different populations.

I always think of Trifolium species as being particularly common and widely distributed, but there are some that are threatened and potentially tell us about the threats faced by imperiled plant populations. In fact, while a number of North American Trifolium species have successfully invaded North America, but T. montanum is not, according to the USDA Plants Database. These results reveal that these negative effects affect plants at different stages of their life cycle (growth to maturity vs. recuitment) and that log-term persistence of these populations requires management activities that ameliorate both of these effects.

SCHLEUNING, M., NIGGEMANN, M., BECKER, U., & MATTHIES, D. (2009). Negative effects of habitat degradation and fragmentation on the declining grassland plant Trifolium montanum Basic and Applied Ecology, 10 (1), 61-69 DOI: 10.1016/j.baae.2007.12.002