Mimicry

@article{huheey1961studies,
    author = "Huheey, James E.",
    title = "Studies in Warning Coloration and Mimicry. III. Evolution of Mullerian Mimicry",
    year = "1961",
    journal = "Evolution",
    url = "https://doi.org/10.2307/2406326",
    doi = "10.2307/2406326",
    number = "4",
    openalex = "W4231107665",
    pages = "567",
    volume = "15"
}

The natural color pattern of individuals of the unpalatable and mimetic butterfly Heliconius erato was altered to a unique nonmimetic pattern. When returned to natural populations, the nonmimetic individuals remained for shorter periods of time and received more wing damage indicative of predator attacks than did the controls. The results indicate that Müllerian mimicry was functioning to protect the butterflies from predation.

@article{benson1972natural,
    author = "Benson, Woodruff W.",
    title = "Natural Selection for Müllerian Mimicry in Heliconius erato in Costa Rica",
    year = "1972",
    journal = "Science",
    abstract = "The natural color pattern of individuals of the unpalatable and mimetic butterfly Heliconius erato was altered to a unique nonmimetic pattern. When returned to natural populations, the nonmimetic individuals remained for shorter periods of time and received more wing damage indicative of predator attacks than did the controls. The results indicate that Müllerian mimicry was functioning to protect the butterflies from predation.",
    url = "https://doi.org/10.1126/science.176.4037.936",
    doi = "10.1126/science.176.4037.936",
    number = "4037",
    pages = "936-939",
    volume = "176"
}

@misc{benson1972natural1,
    author = "Benson, W. W",
    title = "Natural selection for Mllerian mimicry in Heliconus erato in Costa Rica",
    year = "1972",
    howpublished = "Science, v. 176, p. 936-939",
    note = "talkorigins\_source = {true}; raw\_reference = {Benson, W. W., 1972, Natural selection for Mllerian mimicry in Heliconus erato in Costa Rica: Science, v. 176, p. 936-939.}"
}

Field studies of releases and recaptures of diurnal moths painted with yellow to resemble the edible tiger swallowtail and of black moths that resemble a toxic species of swallowtail produced these results: (i) A greater proportion of the black moths were recaptured; (ii) daily trapping for a week after each release showed that the black moths survived longer than the yellow-painted moths; (iii) an analysis of wing injuries shows that most attacks can be attributed to birds and that the yellow-painted moths were attacked more often, more vigorously, or more persistently than the black moths. These results are interpreted as showing a greater predation pressure on the yellow-painted than on the black moths and, therefore, as confirming the Batesian theory of mimicry.

@article{doi101126science1954279681,
    author = "Sternburg, J. G. and Waldbauer, G. P. and Jeffords, Michael R.",
    title = "Batesian Mimicry: Selective Advantage of Color Pattern",
    year = "1977",
    journal = "Science",
    abstract = "Field studies of releases and recaptures of diurnal moths painted with yellow to resemble the edible tiger swallowtail and of black moths that resemble a toxic species of swallowtail produced these results: (i) A greater proportion of the black moths were recaptured; (ii) daily trapping for a week after each release showed that the black moths survived longer than the yellow-painted moths; (iii) an analysis of wing injuries shows that most attacks can be attributed to birds and that the yellow-painted moths were attacked more often, more vigorously, or more persistently than the black moths. These results are interpreted as showing a greater predation pressure on the yellow-painted than on the black moths and, therefore, as confirming the Batesian theory of mimicry.",
    url = "https://doi.org/10.1126/science.195.4279.681",
    doi = "10.1126/science.195.4279.681",
    openalex = "W2019665968",
    references = "doi101126science1443615183, doi1023072406736, doi1023072406796"
}

@article{kingsland1978abbott,
    author = "Kingsland, Sharon",
    title = "Abbott thayer and the protective coloration debate",
    year = "1978",
    journal = "Journal of the History of Biology",
    url = "https://doi.org/10.1007/bf00389300",
    doi = "10.1007/bf00389300",
    number = "2",
    pages = "223-244",
    volume = "11"
}

@article{kingsland1978abbott2,
    author = "Kingsland, S",
    title = "Abbott Thayer and the protective coloration debate",
    year = "1978",
    journal = "Journal of the History of Biology, v. 11, p. 233-244",
    note = "talkorigins\_source = {true}; raw\_reference = {Kingsland, S., 1978, Abbott Thayer and the protective coloration debate: Journal of the History of Biology, v. 11, p. 233-244.}"
}

@incollection{huheey1984warning,
    author = "Huheey, James E.",
    title = "Warning Coloration and Mimicry",
    year = "1984",
    booktitle = "Chemical Ecology of Insects",
    url = "https://doi.org/10.1007/978-1-4899-3368-3\_10",
    doi = "10.1007/978-1-4899-3368-3\_10",
    openalex = "W1822465461",
    pages = "257-297",
    references = "doi101111j109636421860tb00146x, doi101111j155856461964tb01674x, doi1023071437762, doi1023072063069, doi1023072341823, doi1023072576242, doi104039entm9745fv, doi105962bhltitle27468, openalexw2151993477, openalexw3133798068"
}

@misc{crossref1987erato,
    title = "Erato.",
    year = "1987",
    booktitle = "Renaissance Art",
    url = "https://doi.org/10.5040/9798216007562-422",
    doi = "10.5040/9798216007562-422",
    pages = "100-100"
}

▪ Abstract Mimicry and warning color are highly paradoxical adaptations. Color patterns in both Müllerian and Batesian mimicry are often determined by relatively few pattern-regulating loci with major effects. Many of these loci are “supergenes,” consisting of multiple, tightly linked epistatic elements. On the one hand, strong purifying selection on these genes must explain accurate resemblance (a reduction of morphological diversity between species), as well as monomorphic color patterns within species. On the other hand, mimicry has diversified at every taxonomic level; warning color has evolved from cryptic patterns, and there are mimetic polymorphisms within species, multiple color patterns in different geographic races of the same species, mimetic differences between sister species, and multiple mimicry rings within local communities. These contrasting patterns can be explained, in part, by the shape of a “number-dependent” selection function first modeled by Fritz Müller in 1879: Purifying selection against any warning-colored morph is very strong when that morph is rare, but becomes weak in a broad basin of intermediate frequencies, allowing opportunities for polymorphisms and genetic drift. This Müllerian explanation, however, makes unstated assumptions about predator learning and forgetting which have recently been challenged. Today's “receiver psychology” models predict that classical Müllerian mimicry could be much rarer than believed previously, and that “quasi-Batesian mimicry,” a new type of mimicry intermediate between Müllerian and Batesian, could be common. However, the new receiver psychology theory is untested, and indeed it seems to us unlikely; alternative assumptions could easily lead to a more traditional Müllerian/Batesian mimicry divide.

@article{doi101146annurevecolsys301201,
    author = "Mallet, James and Joron, Mathieu",
    title = "Evolution of Diversity in Warning Color and Mimicry: Polymorphisms, Shifting Balance, and Speciation",
    year = "1999",
    journal = "Annual Review of Ecology and Systematics",
    abstract = "▪ Abstract Mimicry and warning color are highly paradoxical adaptations. Color patterns in both Müllerian and Batesian mimicry are often determined by relatively few pattern-regulating loci with major effects. Many of these loci are “supergenes,” consisting of multiple, tightly linked epistatic elements. On the one hand, strong purifying selection on these genes must explain accurate resemblance (a reduction of morphological diversity between species), as well as monomorphic color patterns within species. On the other hand, mimicry has diversified at every taxonomic level; warning color has evolved from cryptic patterns, and there are mimetic polymorphisms within species, multiple color patterns in different geographic races of the same species, mimetic differences between sister species, and multiple mimicry rings within local communities. These contrasting patterns can be explained, in part, by the shape of a “number-dependent” selection function first modeled by Fritz Müller in 1879: Purifying selection against any warning-colored morph is very strong when that morph is rare, but becomes weak in a broad basin of intermediate frequencies, allowing opportunities for polymorphisms and genetic drift. This Müllerian explanation, however, makes unstated assumptions about predator learning and forgetting which have recently been challenged. Today's “receiver psychology” models predict that classical Müllerian mimicry could be much rarer than believed previously, and that “quasi-Batesian mimicry,” a new type of mimicry intermediate between Müllerian and Batesian, could be common. However, the new receiver psychology theory is untested, and indeed it seems to us unlikely; alternative assumptions could easily lead to a more traditional Müllerian/Batesian mimicry divide.",
    url = "https://doi.org/10.1146/annurev.ecolsys.30.1.201",
    doi = "10.1146/annurev.ecolsys.30.1.201",
    openalex = "W2155422373",
    references = "benson1972natural, doi1010160022519364900384, doi1010160022519371901895, doi101016s0022519384800041, doi10103712293000, doi101038384236a0, doi101086284581, doi101093oso97801985498330010001, doi101098rstb19850066, doi101098rstb19950033, doi101111j109583121984tb00143x, doi101111j109583121986tb01772x, doi101111j109583121987tb00435x, doi101111j109583121996tb01452x, doi101111j109636421860tb00146x, doi101111j155856461989tb04237x, doi101146annureven15010170000355, doi1015159780691207278, doi1023072411226, doi1023072420875, doi1023072531471, doi1023074510368, doi104159harvard9780674865327, doi105962bhltitle27468, eaton1940adaptive, openalexw1523652513, openalexw160989634, openalexw2151993477, openalexw2624262714, openalexw3133798068"
}

@incollection{crossref2009warning,
    title = "WARNING COLORATION AND MIMICRY",
    year = "2009",
    booktitle = "Darwinism",
    url = "https://doi.org/10.1017/cbo9780511693076.010",
    doi = "10.1017/cbo9780511693076.010",
    openalex = "W2497519154",
    pages = "232-267"
}

Wing pattern evolution in Heliconius butterflies provides some of the most striking examples of adaptation by natural selection. The genes controlling pattern variation are classic examples of Mendelian loci of large effect, where allelic variation causes large and discrete phenotypic changes and is responsible for both convergent and highly divergent wing pattern evolution across the genus. We characterize nucleotide variation, genotype-by-phenotype associations, linkage disequilibrium (LD), and candidate gene expression patterns across two unlinked genomic intervals that control yellow and red wing pattern variation among mimetic forms of Heliconius erato. Despite very strong natural selection on color pattern, we see neither a strong reduction in genetic diversity nor evidence for extended LD across either patterning interval. This observation highlights the extent that recombination can erase the signature of selection in natural populations and is consistent with the hypothesis that either the adaptive radiation or the alleles controlling it are quite old. However, across both patterning intervals we identified SNPs clustered in several coding regions that were strongly associated with color pattern phenotype. Interestingly, coding regions with associated SNPs were widely separated, suggesting that color pattern alleles may be composed of multiple functional sites, conforming to previous descriptions of these loci as "supergenes." Examination of gene expression levels of genes flanking these regions in both H. erato and its co-mimic, H. melpomene, implicate a gene with high sequence similarity to a kinesin as playing a key role in modulating pattern and provides convincing evidence for parallel changes in gene regulation across co-mimetic lineages. The complex genetic architecture at these color pattern loci stands in marked contrast to the single casual mutations often identified in genetic studies of adaptation, but may be more indicative of the type of genetic changes responsible for much of the adaptive variation found in natural populations.

@article{doi101371journalpgen1000796,
    author = "Counterman, Brian A. and Araújo-Pérez, Félix and Hines, Heather M. and Baxter, Simon W. and Morrison, Clay and Lindstrom, Daniel P. and Papa, Riccardo and Ferguson, Laura and Joron, Mathieu and ffrench‐Constant, Richard H. and Smith, Christopher P. and Nielsen, Dahlia M. and Chen, Rui and Jiggins, Chris D. and Reed, Robert D. and Halder, Georg and Mallet, Jim and McMillan, W. Owen",
    title = "Genomic Hotspots for Adaptation: The Population Genetics of Müllerian Mimicry in Heliconius erato",
    year = "2010",
    journal = "PLoS Genetics",
    abstract = {Wing pattern evolution in Heliconius butterflies provides some of the most striking examples of adaptation by natural selection. The genes controlling pattern variation are classic examples of Mendelian loci of large effect, where allelic variation causes large and discrete phenotypic changes and is responsible for both convergent and highly divergent wing pattern evolution across the genus. We characterize nucleotide variation, genotype-by-phenotype associations, linkage disequilibrium (LD), and candidate gene expression patterns across two unlinked genomic intervals that control yellow and red wing pattern variation among mimetic forms of Heliconius erato. Despite very strong natural selection on color pattern, we see neither a strong reduction in genetic diversity nor evidence for extended LD across either patterning interval. This observation highlights the extent that recombination can erase the signature of selection in natural populations and is consistent with the hypothesis that either the adaptive radiation or the alleles controlling it are quite old. However, across both patterning intervals we identified SNPs clustered in several coding regions that were strongly associated with color pattern phenotype. Interestingly, coding regions with associated SNPs were widely separated, suggesting that color pattern alleles may be composed of multiple functional sites, conforming to previous descriptions of these loci as "supergenes." Examination of gene expression levels of genes flanking these regions in both H. erato and its co-mimic, H. melpomene, implicate a gene with high sequence similarity to a kinesin as playing a key role in modulating pattern and provides convincing evidence for parallel changes in gene regulation across co-mimetic lineages. The complex genetic architecture at these color pattern loci stands in marked contrast to the single casual mutations often identified in genetic studies of adaptation, but may be more indicative of the type of genetic changes responsible for much of the adaptive variation found in natural populations.},
    url = "https://doi.org/10.1371/journal.pgen.1000796",
    doi = "10.1371/journal.pgen.1000796",
    openalex = "W2047862598",
    references = "doi105962p203304"
}

Mimetic wing coloration evolves in butterflies in the context of predator confusion. Unless butterfly eyes have adaptations for discriminating mimetic color variation, mimicry also carries a risk of confusion for the butterflies themselves. Heliconius butterfly eyes, which express recently duplicated ultraviolet (UV) opsins, have such an adaptation. To examine bird and butterfly color vision as sources of selection on butterfly coloration, we studied yellow wing pigmentation in the tribe Heliconiini. We confirmed, using reflectance and mass spectrometry, that only Heliconius use 3-hydroxy-DL-kynurenine (3-OHK), which looks yellow to humans but reflects both UV- and long-wavelength light, whereas butterflies in related genera have chemically unknown yellow pigments mostly lacking UV reflectance. Modeling of these color signals reveals that the two UV photoreceptors of Heliconius are better suited to separating 3-OHK from non-3-OHK spectra compared with the photoreceptors of related genera or birds. The co-occurrence of potentially enhanced UV vision and a UV-reflecting yellow wing pigment could allow unpalatable Heliconius private intraspecific communication in the presence of mimics. Our results are the best available evidence for the correlated evolution of a color signal and color vision. They also suggest that predator visual systems are error prone in the context of mimicry.

@article{doi101086663192,
    author = "Bybee, Seth and Yuan, Furong and Ramstetter, Monica D. and Llorente-Bousquets, Jorge and Reed, Robert D. and Osorio, Daniel and Briscoe, Adriana D.",
    title = "UV Photoreceptors and UV-Yellow Wing Pigments in Heliconius Butterflies Allow a Color Signal to Serve both Mimicry and Intraspecific Communication",
    year = "2011",
    journal = "The American Naturalist",
    abstract = "Mimetic wing coloration evolves in butterflies in the context of predator confusion. Unless butterfly eyes have adaptations for discriminating mimetic color variation, mimicry also carries a risk of confusion for the butterflies themselves. Heliconius butterfly eyes, which express recently duplicated ultraviolet (UV) opsins, have such an adaptation. To examine bird and butterfly color vision as sources of selection on butterfly coloration, we studied yellow wing pigmentation in the tribe Heliconiini. We confirmed, using reflectance and mass spectrometry, that only Heliconius use 3-hydroxy-DL-kynurenine (3-OHK), which looks yellow to humans but reflects both UV- and long-wavelength light, whereas butterflies in related genera have chemically unknown yellow pigments mostly lacking UV reflectance. Modeling of these color signals reveals that the two UV photoreceptors of Heliconius are better suited to separating 3-OHK from non-3-OHK spectra compared with the photoreceptors of related genera or birds. The co-occurrence of potentially enhanced UV vision and a UV-reflecting yellow wing pigment could allow unpalatable Heliconius private intraspecific communication in the presence of mimics. Our results are the best available evidence for the correlated evolution of a color signal and color vision. They also suggest that predator visual systems are error prone in the context of mimicry.",
    url = "https://doi.org/10.1086/663192",
    doi = "10.1086/663192",
    openalex = "W1982684335",
    references = "doi101111j109583121996tb01452x"
}

@misc{lynch2013rudd,
    author = "Lynch, Timothy",
    title = "Rudd, Abbott and the American debate analogy",
    year = "2013",
    url = "https://doi.org/10.64628/aa.fedjpky64",
    doi = "10.64628/aa.fedjpky64",
    openalex = "W4413657367"
}

Convergent evolution, the repeated evolution of similar phenotypes in response to the same selective pressures across multiple lineages, is widespread in nature. The extent to which the same genetic mechanisms contribute to convergent evolution could reveal whether the pathway towards these optimal endpoints is flexible or constrained to follow a particular route. Although mimicry of aposematic colour patterns is well known in Lepidoptera, our knowledge of the genetic basis of these convergent patterns is mostly restricted to a few closely-related species. Here we study the genetic basis of mimicry across seven species of Ithomiini and Heliconius butterflies and a day-flying Chetone moth, representing lineages that diverged between ∼1-120 Mya, each presenting similar colour pattern switches. In all the butterfly species, the genetic variants most strongly associated with convergent colour pattern switches are similarly located in non-coding regions near the genes ivory and optix. Colour pattern variation in the moth is associated with a ∼1 Mb inversion around ivory paralleling the supergene architecture of the co-mimic Heliconius numata. In contrast to previous studies in Heliconius, there is limited evidence of alleles shared by means of hybridization in convergence among closely-related ithomiine species. Repeated parallel evolution of regulatory switches via reuse of the same two genes suggests that convergent colour pattern evolution is highly constrained, even across large evolutionary timescales.

@misc{benchehida2025genetic,
    author = "Ben Chehida, Yacine and van der Heijden, Eva S.M. and Page, Edward J. and Salazar C., Patricio A. and Rosser, Neil and Gabriela Gavilanes Córdova, Kimberly and Sánchez-Prado, Mónica and Sánchez-Carvajal, María José and Chandi, Franz and Arias-Cruz, Alex P and Radford, Maya and Lamas, Gerardo and Jiggins, Chris and Mallet, James and McClure, Melanie and Salazar, Camilo and Elias, Marianne and Bacquet, Caroline N. and Nadeau, Nicola J. and Dasmahapatra, Kanchon K. and Meier, Joana I.",
    title = "Genetic parallelism underpins convergent mimicry coloration across Lepidoptera",
    year = "2025",
    abstract = "Convergent evolution, the repeated evolution of similar phenotypes in response to the same selective pressures across multiple lineages, is widespread in nature. The extent to which the same genetic mechanisms contribute to convergent evolution could reveal whether the pathway towards these optimal endpoints is flexible or constrained to follow a particular route. Although mimicry of aposematic colour patterns is well known in Lepidoptera, our knowledge of the genetic basis of these convergent patterns is mostly restricted to a few closely-related species. Here we study the genetic basis of mimicry across seven species of Ithomiini and Heliconius butterflies and a day-flying Chetone moth, representing lineages that diverged between ∼1-120 Mya, each presenting similar colour pattern switches. In all the butterfly species, the genetic variants most strongly associated with convergent colour pattern switches are similarly located in non-coding regions near the genes ivory and optix. Colour pattern variation in the moth is associated with a ∼1 Mb inversion around ivory paralleling the supergene architecture of the co-mimic Heliconius numata. In contrast to previous studies in Heliconius, there is limited evidence of alleles shared by means of hybridization in convergence among closely-related ithomiine species. Repeated parallel evolution of regulatory switches via reuse of the same two genes suggests that convergent colour pattern evolution is highly constrained, even across large evolutionary timescales.",
    url = "https://doi.org/10.1101/2025.06.26.661542",
    doi = "10.1101/2025.06.26.661542",
    openalex = "W4411744351"
}

@misc{crossrefNoneerato,
    title = "Erato",
    year = "None",
    booktitle = "Brill’s New Pauly",
    url = "https://doi.org/10.1163/1574-9347\_bnp\_e400750",
    doi = "10.1163/1574-9347\_bnp\_e400750"
}