Abstract

Heliconius are unpalatable butterflies that exhibit remarkable intra‐ and interspecific variation in wing color pattern, specifically warning coloration. Species that have converged on the same pattern are often clustered in Müllerian mimicry rings. Overall, wing color patterns are nearly identical among co‐mimics. However, fine‐scale differences exist, indicating that factors in addition to natural selection may underlie wing phenotype. Here, we investigate differences in shape and size of the forewing and the red band in the Heliconius postman mimicry ring (H. erato phyllis and the co‐mimics H. besckei, H. melpomene burchelli, and H. melpomene nanna) using a landmark‐based approach. If phenotypic evolution is driven entirely by predation pressure, we expect nonsignificant differences among co‐mimics in terms of wing shape. Also, a reinforcement of wing pattern (i.e., greater similarity) could occur when co‐mimics are in sympatry. We also examined variation in the red forewing band because this trait is critical for both mimicry and sexual communication. Morphometric results revealed significant but small differences among species, particularly in the shape of the forewing of co‐mimics. Although we did not observe greater similarity when co‐mimics were in sympatry, nearly identical patterns provided evidence of convergence for mimicry. In contrast, mimetic pairs could be distinguished based on the shape (but not the size) of the red band, suggesting an “advergence” process. In addition, sexual dimorphism in the red band shape (but not size) was found for all lineages. Thus, we infer that natural selection due to predation by birds might not be the only mechanism responsible for variation in color patterns, and sexual selection could be an important driver of wing phenotypic evolution in this mimicry ring.

Highlights

  • Neotropical Heliconius butterflies represent a conspicuously variable group from a morphological perspective (Holzinger & Holzinger, 1994), generally considered as a good model of evolutionary studies

  • The analysis of variance (ANOVA) results comparing the effect of sex and geographic group on red band size suggest that both mimicry convergence and sexual dimorphism act in H. erato phyllis (F5,54 = 2.60, r2 = .12, p = .03) and that there is an interaction between these factors

  • As sexual dimorphism in the shape of the red band was found in all lineages explored in this study, the results suggest that sexual selection may be involved in the evolution of this trait

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Summary

| INTRODUCTION

Neotropical Heliconius butterflies represent a conspicuously variable group from a morphological perspective (Holzinger & Holzinger, 1994), generally considered as a good model of evolutionary studies. The existence of phenotypic variation may be advantageous to bird predators, because those individuals that better associate the unpalatability signal on the butterfly wing may have greater survivorship (Mallet & Barton, 1989) Learning in this case may be associated with several phenotypic, behavioral, and ecological factors associated with the butterflies, for example: (1) color (Svádová et al, 2009) and pattern (Ihalainen, Lindström, Mappes, & Puolakkainen, 2008) of the wings; (2) level of unpalatability (Ihalainen, 2006); and (3) relative frequency of the co-­mimics (Mérot et al, 2016; Rowland, Wiley, Ruxton, Mappes, & Speed, 2010; Speed, 2001). If phenotypic resemblance among co-­mimics resulted mainly by predation pressure (i.e., convergence for mimicry; Kapan (2001); Mallet and Barton (1989)), we expected (1) nonsignificant differences among species (i.e., nearly identical co-­mimics) considering the size and shape of whole forewing and (2) reinforcement of wing patterning when co-­mimics are in sympatry. In particular, because it is responsible for red wing pattern variation across Heliconius species (Reed et al, 2011)

| MATERIALS AND METHODS
| DISCUSSION
Findings
| CONCLUSION
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