Abstract

Müllerian mimicry is a classic example of adaptation, yet Müller's original theory does not account for the diversity often observed in mimicry rings. Here, we aimed to assess how well classical Müllerian mimicry can account for the colour polymorphism found in chemically defended Oreina leaf beetles by using field data and laboratory assays of predator behaviour. We also evaluated the hypothesis that thermoregulation can explain diversity between Oreina mimicry rings. We found that frequencies of each colour morph were positively correlated among species, a critical prediction of Müllerian mimicry. Predators learned to associate colour with chemical defences. Learned avoidance of the green morph of one species protected green morphs of another species. Avoidance of blue morphs was completely generalized to green morphs, but surprisingly, avoidance of green morphs was less generalized to blue morphs. This asymmetrical generalization should favour green morphs: indeed, green morphs persist in blue communities, whereas blue morphs are entirely excluded from green communities. We did not find a correlation between elevation and coloration, rejecting thermoregulation as an explanation for diversity between mimicry rings. Biased predation could explain within-community diversity in warning coloration, providing a solution to a long-standing puzzle. We propose testable hypotheses for why asymmetric generalization occurs, and how predators maintain the predominance of blue morphs in a community, despite asymmetric generalization.

Highlights

  • Aposematic signals that warn predators about prey defences have long occupied biologists’ attention as a prime example of adaptation (Poulton, 1890; Ruxton, Allen, Sherratt, & Speed, 2018; Wallace, 1867)

  • We modelled the ratio of blue:green beetles of “other” species at each site, with the proportion of blue O. cacaliae as a predictor, and the location of each site included as a random effect

  • Cox proportional hazards regression revealed that birds initially hesitated ~50 s longer in trial 1 to attack green beetles than blue beetles (Table 1a, Figure 5)

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Summary

Introduction

Aposematic signals that warn predators about prey defences have long occupied biologists’ attention as a prime example of adaptation (Poulton, 1890; Ruxton, Allen, Sherratt, & Speed, 2018; Wallace, 1867). Aposematism is subject to positive frequency-dependent selection (Fisher, 1958). When it is rare, predation will disfavour aposematism because few predators will have had the opportunity to learn to avoid the warning signal, so those few aposematic individuals present will suffer high per capita mortality Positive frequency dependence can be so strong that individuals of different defended species evolve to resemble one another, in a phenomenon called Müllerian mimicry (Müller, 1879)

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