A comparative analysis of the evolution of imperfect mimicry.

Although exceptional examples of adaptation are frequently celebrated, some outcomes of natural selection seem far from perfect. For example, many hoverflies (Diptera: Syrphidae) are harmless (Batesian) mimics of stinging Hymenoptera. However, although some hoverfly species are considered excellent mimics, other species bear only a superficial resemblance to their models and it is unclear why this is so. To evaluate hypotheses that have been put forward to explain interspecific variation in the mimetic fidelity of Palearctic Syrphidae we use a comparative approach. We show that the most plausible explanation is that predators impose less selection for mimetic fidelity on smaller hoverfly species because they are less profitable prey items. In particular, our findings, in combination with previous results, allow us to reject several key hypotheses for imperfect mimicry: first, human ratings of mimetic fidelity are positively correlated with both morphometric measures and avian rankings, indicating that variation in mimetic fidelity is not simply an illusion based on human perception; second, no species of syrphid maps out in multidimensional space as being intermediate in appearance between several different hymenopteran model species, as the multimodel hypothesis requires; and third, we find no evidence for a negative relationship between mimetic fidelity and abundance, which calls into question the kin-selection hypothesis. By contrast, a strong positive relationship between mimetic fidelity and body size supports the relaxed-selection hypothesis, suggesting that reduced predation pressure on less profitable prey species limits the selection for mimetic perfection.

Background evolution in camouflage systems: a predator-prey/pollinator-flower game.

A common predator or anti-predator strategy involves camouflage based on background matching. In some systems, the background is an organism whose fitness is affected by the predator-prey interaction. In these cases, the phenotype of the background species may evolve to affect the degree of background matching in the predator-prey interaction. For example, some flower species (the background) are inhabited by camouflaged ambush predators that attack visiting pollinators. These flowers have a fitness interest in the outcome of the predator-prey interaction because flowers depend on pollinator visitations for reproduction. Therefore, floral colour might evolve relative to predator colour so as to influence the detectability of resident predators. I have created a three-player game, based on Signal Detection Theory, to model the co-evolution of predator and prey/pollinator behavioural strategies with floral colour. This model makes two general predictions: (1) Constraints on predator distributions favour the evolution of flowers that match the predators' colour because they prevent predators from overexploiting these flowers; (2) factors that produce less discriminating pollinators also favour the evolution of flowers that match the predators' colour because these pollinators are willing to land on these flowers even if the safety of the flower is in doubt.