RESEARCH: MIMICRY AND SPECIATION IN PAPILIO SWALLOWTAIL BUTTERFLIES

Swallowtail butterflies (genus Papilio) have undergone one of the most spectacular radiations in the animal world. With over 200 species and probably twice that many subspecies described, it makes a very diverse taxonomic group. Its members include such large, colorful and prominent species as the Ulysses Swallowtail (Papilio ulysses), Common Yellow Swallowtail (P. machaon), Eastern Tiger Swallowtail (P. glaucus), Mocker Swallowtail (Papilio dardanus), and Paris Peacock (P. paris).

Mormon swallowtails (Menelaides) make up the largest subgenus of Papilio, with approximately 50 species (25% of the genus) and 200 subspecies distributed over the Indo-Australian Region. The geographic mosaic of their distributional ranges is characterized by endemism and diversification in regional hotspots on one hand and subspeciation at sometimes micro-spatial scales on the other. This makes them a promising group in which to study speciation and subspeciation, and biogeographic processes such as dispersal, isolation and vicariance. Many of them also show a spectacular diversity of Batesian mimicry and wing patterns, and the subgenus includes such iconic mimetic species as Papilio memnon, P. polyes and P. aegeus. Mimicry in some species is female-limited and polymorphic, i.e., females in these species are mimetic and appear in several forms, whereas males are monomorphic and non-mimetic, representing ancestral color patterns of the species. This makes them ideal subjects for testing theories of natural, sexual and frequency-dependent selection. Finally, the evolution of mimicry is believed to have driven genomic re-arrangements and positive selection on wing-patterning genes, which offer opportunities to study the molecular genetic aspects of evolution. Together, these attributes make mormon swallowtails a model system to study biodiversity and selection at various scales.

Currently we are working towards building foundations on which to develop Papilio as a model clade to study morphological diversification, evolution of wing color patterns and mimicry, and speciation. With Felix Sperling and Adam Cotton we are preparing a species-level molecular phylogeny that will cover all the species in this genus, with analyses of biogeography, wing color pattern evolution and mimicry closely following. We are investigating the molecular genetic and chemical bases of female-limited mimicry in Papilo polytes. We are also currently doing and planning several behavioral experiments and field observational studies to test theories about Batesian mimicry, female-limited mimicry, predation, natural selection and sexual selection using several species of Menelaides. Details and updates will be provided on this page as major projects are completed.

Publications:

Deshmukh, R., S. Baral, M. Kuwalekar, A. G. Kizhakke, and K. Kunte. 2025. Reproductive barriers and genomic hotspots of adaptation during allopatric species divergence. Molecular Ecology, 34:e17703. PDF file (1.3 MB; Open Access). Published in the special issue, Genomics of Speciation.

Deshmukh, R., S. Baral, A. G. Kizhakke, M. Kuwalekar, and K. Kunte. 2024. Evolution of novel mimicry polymorphisms through Haldane’s sieve and rare recombination. eLife, 13:RP101346.

Dharmaraaj, B., R. Venkatesan, and K. Kunte. 2024. Spectral variation and pigmentary basis of ornamental and mimetic wing colour patches of swallowtail butterflies. Evolutionary Journal of the Linnean Society, kzae018. PDF File (6 MB; Open Access).

Basu, D. N., V. Bhaumik, and K. Kunte. 2023. The tempo and mode of character evolution in the assembly of mimetic communities. Proceedings of the National Academy of Sciences, USA, 120:e2203724120. PDF file (8 MB; now Open Access). Featured on the journal cover, in a PNAS Commetary, 'Butterfly mimicry rings run in circles', and in news articles on NCBS News (The Art of Subterfuge), Deccan Herald, and The Hindu.

Condamine, F. L., R. Allio, E. L. Reboud, J. R. Dupuis, E. F. A. Toussaint, N. Mazet, S.-J. Hue, D. S. Lewis, K. Kunte†, A. M. Cotton†, and F. A. H. Sperling†. 2023. A comprehensive phylogeny and revised taxonomy illuminate the origin and diversification of the global radiation of Papilio (Lepidoptera: Papilionidae). Molecular Phylogenetics and Evolution, 183:107758. † equal senior authors. PDF file (9 MB).

Deshmukh, R., D. Lakhe, and K. Kunte. 2020. Tissue-specific developmental regulation and isoform usage underlie the role of doublesex in sex differentiation and mimicry in Papilio swallowtails. Royal Society Open Science, 7:200792. PDF file (1.3 MB; Open Access).

Baral, S., Gandhimathi A., R. Deshmukh, and K. Kunte. 2019. Genetic architecture and sex-specific selection govern modular, male-biased evolution of doublesex. Science Advances, 5:eaau3753. PDF file (5.5MB).

Gaitonde, N., J. Joshi, and K. Kunte. 2018. Evolution of ontogenic change in color defenses of swallowtail butterflies. Ecology and Evolution, online early. PDF file (1.2MB).

Deshmukh, R., S. Baral, A. Gandhimathi, M. Kuwalekar, and K. Kunte. 2018. Mimicry in butterflies: co-option and a bag of magnificent developmental genetic tricks. WIREs Developmental Biology, 7:e291. PDF file (8.5MB, has colour figures).

Arnold, M. L. and K. Kunte. 2017. Adaptive genetic exchange: a tangled history of admixture and evolutionary innovation. Trends in Ecology and Evolution, 32:601–611. PDF file (1.8MB, has colour figures). Recommended by Faculty of 1000 (F1000Prime) (download the recommendation if you do not have a subscription).

Joshi, J., A. Prakash, and K. Kunte. 2017. Evolutionary assembly of communities in butterfly mimicry rings. The American Naturalist, 189:E58–E76. PDF file (12MB, has colour figures). Winner of the American Society of Naturalists' 2018 Presidential Award, "honoring an outstanding article published in The American Naturalist in the previous year". See coverage on NCBS News.

Su, S., M. Lim, and K. Kunte. 2015. Prey from the eyes of predators: colour discriminability of aposematic and mimetic butterflies from an avian visual perspective. Evolution, 69:2985–2994. PDF file (650KB; or PDF file with supporting information, 1.1MB). This work was featured on the cover of the journal issue, and in several popular science news stories in PhysOrg, EurekAlert, The Hindu, The Statesman, and NCBS News, among others.

Kunte, K., W. Zhang, A. Tenger-Trolander, D. H. Palmer, A. Martin, R. D. Reed, S. P. Mullen, and M. R. Kronforst. 2014. doublesex is a mimicry supergene. Nature, 507:229–232. PDF file (1.2MB). Recommended by Faculty of 1000 (F1000Prime) (download the recommendation if you do not have a subscription). Read more about the story on this webpage. See popular science coverage of this paper in Nature, Nature India, Science, NCBS News, The Scientist, National Geographic's Phenomena blog, Mongabay, New York Times, The Hindu, and ScienceDaily.

Kunte, K., C. Shea, M. L. Aardema, J. M. Scriber, T. E. Juenger, L. E. Gilbert, and M. R. Kronforst. 2011. Sex chromosome mosaicism and hybrid speciation among tiger swallowtail butterflies. PLoS Genetics, 7:e1002274. See the full paper online on the journal website, or its news coverage on the National Science Foundation (NSF, USA) website, the University of Texas at Austin and the College of Natural Sciences websites, in Harvard Gazette, or on MSNBC, and myscience. Read more about this work ...

Kunte, K. 2009. Female-limited mimetic polymorphism: A review of theories and a critique of sexual selection as balancing selection. Animal Behaviour, 78:1029–1036. PDF file (586 KB, includes a color figure). Read more about this work ...

Kunte, K. 2009. The diversity and evolution of Batesian mimicry in Papilio swallowtail butterflies. Evolution, 63:2707–2716. PDF file (987KB, includes a color figure).

Kunte, K. 2008. Mimetic butterflies support Wallace's model of sexual dimorphism. Proceedings of the Royal Society, B, 275:1617–1624 PDF file (404 KB). Recommended by Faculty of 1000 (F1000Prime) (download the recommendation if you do not have a subscription). Featured on Science website, and also in Roughgarden, J. 2009. The Genial Gene: Deconstructing Darwinian Selfishness, Univ. of California Press, Berkeley, pp. 38-39.