Competitive Displacement and Agonistic Character Displacement, or the Ghost of Interference Competition.

AbstractInterference competition can drive species apart in habitat use through competitive displacement in ecological time and agonistic character displacement (ACD) over evolutionary time. As predicted by ACD theory, sympatric species of rubyspot damselflies (Hetaerina spp.) that respond more aggressively to each other in staged encounters differ more in microhabitat use. However, the same pattern could arise from competitive displacement if dominant species actively exclude subordinate species from preferred microhabitats. The degree to which habitat partitioning is caused by competitive displacement can be assessed with removal experiments. We carried out removal experiments with three species pairs of rubyspot damselflies. With competitive displacement, removing dominant species should allow subordinate species to shift into the dominant species' microhabitat. Instead, we found that species-specific microhabitat use persisted after the experimental removals. Thus, the previously documented association between heterospecific aggression and microhabitat partitioning in this genus is most likely a product of divergence in habitat preferences caused by interference competition in the evolutionary past.

Predicting evolutionary responses to interspecific interference in the wild.

Many interspecifically territorial species interfere with each other reproductively, and in some cases, aggression towards heterospecifics may be an adaptive response to interspecific mate competition. This hypothesis was recently formalised in an agonistic character displacement (ACD) model which predicts that species should evolve to defend territories against heterospecific rivals above a threshold level of reproductive interference. To test this prediction, we parameterised the model with field estimates of reproductive interference for 32 sympatric damselfly populations and ran evolutionary simulations. Asymmetries in reproductive interference made the outcome inherently unpredictable in some cases, but 80% of the model's stable outcomes matched levels of heterospecific aggression in the field, significantly exceeding chance expectations. In addition to bolstering the evidence for ACD, this paper introduces a new, predictive approach to testing character displacement theory that, if applied to other systems, could help in resolving long-standing questions about the importance of character displacement processes in nature.

A General Explanation for the Persistence of Reproductive Interference.

Reproductive interference is widespread, despite the theoretical expectation that it should be eliminated by reproductive character displacement (RCD). A possible explanation is that females of sympatric species are too similar phenotypically for males to distinguish between them, resulting in a type of evolutionary dilemma or catch-22 in which reproductive interference persists because male mate recognition (MR) cannot evolve until female phenotypes diverge further, and vice versa. Here we illustrate and test this hypothesis with data on rubyspot damselflies (Hetaerina spp.). First, reproductive isolation owing to male MR breaks down with increasing interspecific similarity in female phenotypes. Second, comparing allopatric and sympatric populations yielded no evidence for RCD, suggesting that parallel divergence in female coloration and male MR in allopatry determines the level of reproductive isolation on secondary contact. Whenever reproductive isolation depends on male MR and females of sympatric species are phenotypically similar, the evolutionary catch-22 hypothesis offers an explanation for the persistence of reproductive interference.