Persistently rare species experience stronger negative frequency dependence than common species: A statistical attractor that is hard to avoid

Aim Why are so many species rare, yet persistent? Possibly, rare species experience strong negative frequency dependence (NFD, i.e. strong intraspecific competition relative to interspecific competition), which both makes them rare and buffers them against extinction. A second, not mutually exclusive, possibility is that rare species that experience weak NFD go extinct quickly due to demographic and environmental stochasticity. Both possibilities predict that persistent rare species will experience stronger NFD than common ones. A previous study confirmed this prediction in a range of mostly terrestrial communities. Here, we test that prediction in lake zooplankton and explore its theoretical basis. Location Fifty-three temperate lakes. Time period 1970-2011. Major taxa studied Cladocerans, copepods. Methods We used long-term time series data to estimate the covariance between strength of NFD and mean frequency (relative abundance) for crustacean zooplankton. We used a randomization test to ask whether the covariance between NFD and mean frequency is stronger than expected, given sampling error. We also calculated the covariance between NFD and mean frequency in simulated communities from three different ecological models. Results Rare species experience significantly stronger NFD than common species in over half of the communities. The distribution of associations between NFD and rarity is skewed towards weak associations, which always occur in communities with high evenness. All three theoretical models reproduce these empirical results, even though they are based on different mechanisms (demographic or environmental stochasticity, and mathematical constraints on possible NFD-frequency relationships). Main conclusions Rare species typically experience stronger NFD than common ones because there are many different ecological scenarios in which they will do so, and only a few scenarios in which they won't. Like several other macroecological patterns, the tendency for rare species to experience stronger NFD than common ones is a statistical attractor that is hard to avoid.