#  Papers 

 



 [### Older publications here ...

 ](https://jamesmallet.github.io/publications.html) 

 

 [### Book chapters here...

 ](/book-chapters) 

 

 

 

 

 

 

##  Recent Publications 

 



  Download 122 citations  download- [BibTeX](/bibcite/export?pager_style=standard_pager&number_of_items=12&sort_field=bibcite_year--desc&taxonomy_filters=&&&format=bibtex)
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### Forthcoming

Seixas, F., Dendy, S., Guan, S., Rosser, N., Grishin, N., Davies, N., Trujano-Ortega, M., Myers, T., Counterman, B., Papa, R., Gilbert, L. , jr, McMillan, W., &amp; Mallet, J. (2026). [Ancient inversion polymorphisms are locally adaptive in a widespread butterfly species](/publication/ancient-inversion-polymorphisms-are-locally-adaptive-widespread-butterfly-species). *Nature Communications*.



 

 

Seixas, F., Dendy, S., Guan, S., Rosser, N., Grishin, N., Davies, N., Trujano-Ortega, M., Myers, T., Counterman, B., Papa, R., Gilbert, L. , jr, McMillan, W., &amp; Mallet, J. (2026). [Ancient inversion polymorphisms are locally adaptive in a widespread butterfly species](/publication/ancient-inversion-polymorphisms-are-locally-adaptive-widespread-butterfly-species). *Nature Communications*.



 

 

 

 

Ben Chehida, Y., van der Heijden, E., Page, E., C., P. S., Rosser, N., C., K. G., Sánchez-Prado, M., Sánchez-Carvajal, M., Chandi, F., Arias-Cruz, A., Radford, M., Lamas, G., Jiggins, C., Mallet, J., McClure, M., Salazar, C., Elias, M., Bacquet, C., Nadeau, N., … Meier, J. (2026). [Genetic parallelism underpins mimicry across a wide range of divergence times](/publication/genetic-parallelism-underpins-mimicry-across-wide-range-divergence-times). *PLoS Biology*.



 

 

Ben Chehida, Y., van der Heijden, E., Page, E., C., P. S., Rosser, N., C., K. G., Sánchez-Prado, M., Sánchez-Carvajal, M., Chandi, F., Arias-Cruz, A., Radford, M., Lamas, G., Jiggins, C., Mallet, J., McClure, M., Salazar, C., Elias, M., Bacquet, C., Nadeau, N., … Meier, J. (2026). [Genetic parallelism underpins mimicry across a wide range of divergence times](/publication/genetic-parallelism-underpins-mimicry-across-wide-range-divergence-times). *PLoS Biology*.



 

 

 

 

 



### 2026

Thawornwattana, Y., &amp; Mallet, J. (2026). [Utility of subspecies in mosquito taxonomy](https://doi.org/10.1093/zoolinnean/zlag033). *Zoological Journal of the Linnean Society*, *206*(4).



 

 

Thawornwattana, Y., &amp; Mallet, J. (2026). [Utility of subspecies in mosquito taxonomy](https://doi.org/10.1093/zoolinnean/zlag033). *Zoological Journal of the Linnean Society*, *206*(4).



 

 

 

 

 



### 2025

[Müllerian mimicry in Neotropical butterflies: one mimicry ring to bring them all and in the jungle bind them](https://doi.org/10.1111/geb.70127). (2025). *Global Ecology and Biogeography*, *34*(9), e70127. (Original work published 2025)



 

 

[Müllerian mimicry in Neotropical butterflies: one mimicry ring to bring them all and in the jungle bind them](https://doi.org/10.1111/geb.70127). (2025). *Global Ecology and Biogeography*, *34*(9), e70127. (Original work published 2025)



 

 

 

- [ downloadREPRINT](https://doi.org/10.1111/geb.70127)
 
- [ downloadREPRINT](https://doi.org/10.1111/geb.70127)
 
 

[Mimicry and gene flow among butterfly species: reply to Brower and Garzón-Orduña](/publication/mimicry-and-gene-flow-among-butterfly-species-reply-brower-and-garzon-orduna). (2025). *Antenna*, *49*(1), 3-7. (Original work published 2025)



 

 

[Mimicry and gene flow among butterfly species: reply to Brower and Garzón-Orduña](/publication/mimicry-and-gene-flow-among-butterfly-species-reply-brower-and-garzon-orduna). (2025). *Antenna*, *49*(1), 3-7. (Original work published 2025)



 

 

 

- add\_circle\_outline do\_not\_disturb\_on Abstract
- [ picture\_as\_pdfMallet &amp; Dasmahapatra 202...](/sites/g/files/omnuum6606/files/2025-04/Mallet%20%26%20Dasmahapatra%202025%20proof.pdf)
- [ descriptionA reply to: Brower, A.V.Z., &amp; Garzón-Orduña, I.J. 2024. Evolution of mimicry in Heliconius butterflies: historical hypotheses meet modern models. Antenna 178:171-176. https://www.royensoc.co.uk/publications/antenna/antenna-volume-48-4-2024/](https://www.royensoc.co.uk/publications/antenna/antenna-volume-48-4-2024/)
 
Once again, Brower and Garzón- Orduña criticize the majority of their fellow students of the butterfly genus Heliconius, especially the current authors. Once again, they fail to provide any new data or even reanalyses of the copious data we have...



 

 

- [ picture\_as\_pdfMallet &amp; Dasmahapatra 202...](/sites/g/files/omnuum6606/files/2025-04/Mallet%20%26%20Dasmahapatra%202025%20proof.pdf)
- [ descriptionA reply to: Brower, A.V.Z., &amp; Garzón-Orduña, I.J. 2024. Evolution of mimicry in Heliconius butterflies: historical hypotheses meet modern models. Antenna 178:171-176. https://www.royensoc.co.uk/publications/antenna/antenna-volume-48-4-2024/](https://www.royensoc.co.uk/publications/antenna/antenna-volume-48-4-2024/)
 
 

Thawornwattana, Y., Flouri, T., Mallet, J., &amp; Yang, Z. (2025). [Inference of gene flow between species from genomic data when the mode, direction, and lineages are misspecified](https://doi.org/10.1093/molbev/msaf121). *Molecular Biology and Evolution*, *42*(6), msaf121.



 

 

Thawornwattana, Y., Flouri, T., Mallet, J., &amp; Yang, Z. (2025). [Inference of gene flow between species from genomic data when the mode, direction, and lineages are misspecified](https://doi.org/10.1093/molbev/msaf121). *Molecular Biology and Evolution*, *42*(6), msaf121.



 

 

 

 

 



### 2024

Peñalba, J., Runemark, A., Meier, J., Singh, P., Wogan, G., Sánchez-Guillén, R., Mallet, J., Rometsch, S., Menon, M., Seehausen, O., Kulmuni, J., &amp; Pereira, R. (2024). [The role of hybridization in species formation and persistence](/publications/role-hybridization-species-formation-and-persistence). *Cold Spring Harbor Perspectives in Biology*, a041445. (Original work published 2024)



 

 

Peñalba, J., Runemark, A., Meier, J., Singh, P., Wogan, G., Sánchez-Guillén, R., Mallet, J., Rometsch, S., Menon, M., Seehausen, O., Kulmuni, J., &amp; Pereira, R. (2024). [The role of hybridization in species formation and persistence](/publications/role-hybridization-species-formation-and-persistence). *Cold Spring Harbor Perspectives in Biology*, a041445. (Original work published 2024)



 

 

 

- add\_circle\_outline do\_not\_disturb\_on Abstract
- [ picture\_as\_pdfpenalba\_et\_al\_2024\_hybrid...](/sites/g/files/omnuum6606/files/malletlab/files/penalba_et_al_2024_hybridizationreview.pdf)
 
 Hybridization, or interbreeding between different taxa, was traditionally considered to be rare and to have a largely detrimental impact on biodiversity, sometimes leading to the breakdown of reproductive isolation and even to the reversal of speciation... 

 

 

- [ picture\_as\_pdfpenalba\_et\_al\_2024\_hybrid...](/sites/g/files/omnuum6606/files/malletlab/files/penalba_et_al_2024_hybridizationreview.pdf)
 
 

Rosser, N., Seixas, F., Queste, L. M., Cama, B., Mori-Pezo, R., Kryvokhyzha, D., Nelson, M., de Figueiredo, C., Freitas, A., Joron, M., Kozak, K., Lamas, G., Martins, A., McMillan, O., Ready, J., Rueda-Muñoz, N., Salazar, C., Salazar, P., Schulz, S., … Dasmahapatra, K. K. (2024). [Hybrid speciation driven by multilocus introgression of ecological traits](/publications/hybrid-speciation-driven-multilocus-introgression-ecological-traits). *Nature*, *628*(25 April 2024), 811-817.



 

 

Rosser, N., Seixas, F., Queste, L. M., Cama, B., Mori-Pezo, R., Kryvokhyzha, D., Nelson, M., de Figueiredo, C., Freitas, A., Joron, M., Kozak, K., Lamas, G., Martins, A., McMillan, O., Ready, J., Rueda-Muñoz, N., Salazar, C., Salazar, P., Schulz, S., … Dasmahapatra, K. K. (2024). [Hybrid speciation driven by multilocus introgression of ecological traits](/publications/hybrid-speciation-driven-multilocus-introgression-ecological-traits). *Nature*, *628*(25 April 2024), 811-817.



 

 

 

- add\_circle\_outline do\_not\_disturb\_on Abstract
- [ descriptionPublisher's Version](https://doi.org/10.1038/s41586-024-07263-w)
- [ picture\_as\_pdfrosser\_et\_al.\_2024\_hybrid...](/sites/g/files/omnuum6606/files/malletlab/files/rosser_et_al._2024_hybrid_speciation.printed.pdf)
 
 Hybridisation allows adaptations to be shared among lineages and may trigger the evolution of new species. However, convincing examples of homoploid hybrid speciation remain rare because it is challenging to demonstrate that hybridisation was critical in... 

 

 

- [ descriptionPublisher's Version](https://doi.org/10.1038/s41586-024-07263-w)
- [ picture\_as\_pdfrosser\_et\_al.\_2024\_hybrid...](/sites/g/files/omnuum6606/files/malletlab/files/rosser_et_al._2024_hybrid_speciation.printed.pdf)
 
 

Mallet, J. (2024). *[Colour patterns in Lepidoptera evolved mainly as signals to predators. Heliconius are indeed Müllerian mimics. Reply to Mouy (2022).](https://www.heliconius.org/2024/no-evidence-for-henri-mouys-views-aposematism-and-mimicry/)* Heliconius.Org.



 

 

Mallet, J. (2024). *[Colour patterns in Lepidoptera evolved mainly as signals to predators. Heliconius are indeed Müllerian mimics. Reply to Mouy (2022).](https://www.heliconius.org/2024/no-evidence-for-henri-mouys-views-aposematism-and-mimicry/)* Heliconius.Org.



 

 

 

- add\_circle\_outline do\_not\_disturb\_on Abstract
- [ descriptionPublisher's Version](https://www.heliconius.org/2024/no-evidence-for-henri-mouys-views-aposematism-and-mimicry/)
 
 A recent paper in the journal Evolutionary Ecology by Henri Mouy, an independent researcher and theoretician who gives his address as “City of Westminster, London, UK,” argues that aposematism (warning colour) and Müllerian mimicry are problematic... 

 

 

- [ descriptionPublisher's Version](https://www.heliconius.org/2024/no-evidence-for-henri-mouys-views-aposematism-and-mimicry/)
 
 

Page, E., Queste, L. M., Rosser, N., Salazar, P., Nadeau, N., Mallet, J., Srygley, R., McMillan, O., &amp; Dasmahapatra, K. K. (2024). [Pervasive mimicry in flight behavior among aposematic butterflies](/publications/pervasive-mimicry-flight-behavior-among-aposematicbutterflies). *Proceedings of the National Academy of Sciences, USA*, *121*(11), e2300886121.



 

 

Page, E., Queste, L. M., Rosser, N., Salazar, P., Nadeau, N., Mallet, J., Srygley, R., McMillan, O., &amp; Dasmahapatra, K. K. (2024). [Pervasive mimicry in flight behavior among aposematic butterflies](/publications/pervasive-mimicry-flight-behavior-among-aposematicbutterflies). *Proceedings of the National Academy of Sciences, USA*, *121*(11), e2300886121.



 

 

 

- add\_circle\_outline do\_not\_disturb\_on Abstract
- [ descriptionPublisher's Version](https://doi.org/10.1073/pnas.2300886121)
- [ picture\_as\_pdfpage\_et\_al.\_2024\_flight\_m...](/sites/g/files/omnuum6606/files/malletlab/files/page_et_al._2024_flight_mimicry_pnas.pdf)
 
 Flight was a key innovation in the adaptive radiation of insects. However, it is a complex  
trait influenced by a large number of interacting biotic and abiotic factors, making it  
difficult to unravel the evolutionary drivers. We investigate flight patterns... 

 

 

- [ descriptionPublisher's Version](https://doi.org/10.1073/pnas.2300886121)
- [ picture\_as\_pdfpage\_et\_al.\_2024\_flight\_m...](/sites/g/files/omnuum6606/files/malletlab/files/page_et_al._2024_flight_mimicry_pnas.pdf)
 
 

Seixas, F., Dendy, S., Guan, S., Rosser, Grishin, N., Davies, N., Gilbert, L. , jr., McMillan, W., &amp; Mallet. (2024). [Ancient inversion polymorphisms are locally adaptive in a widespread butterfly species](/publications/ancient-inversion-polymorphisms-are-locally-adaptive-widespread-butterfly). *BioRxiv*, 2024.09.20.614156.



 

 

Seixas, F., Dendy, S., Guan, S., Rosser, Grishin, N., Davies, N., Gilbert, L. , jr., McMillan, W., &amp; Mallet. (2024). [Ancient inversion polymorphisms are locally adaptive in a widespread butterfly species](/publications/ancient-inversion-polymorphisms-are-locally-adaptive-widespread-butterfly). *BioRxiv*, 2024.09.20.614156.



 

 

 

- add\_circle\_outline do\_not\_disturb\_on Abstract
- [ descriptionPublisher's Version](http://biorxiv.org/content/early/2024/09/24/2024.09.20.614156.abstract)
 
 Wide-ranging species are subject to varying biotic and abiotic selection pressures across their distribution. While local adaptation does not manifest in obvious morphological changes, population genomic studies can reveal cryptic diversity and provide... 

 

 

- [ descriptionPublisher's Version](http://biorxiv.org/content/early/2024/09/24/2024.09.20.614156.abstract)
 
 

Seixas, F., Guerrero, S., Xiong, T., &amp; Mallet. (2024). [Temperature-dependent and sex-specific phenotypic plasticity in Heliconius charithonia](/publications/temperature-dependent-and-sex-specific-phenotypic-plasticity-heliconius). *BioRxiv*, 2024.10.22.619625.



 

 

Seixas, F., Guerrero, S., Xiong, T., &amp; Mallet. (2024). [Temperature-dependent and sex-specific phenotypic plasticity in Heliconius charithonia](/publications/temperature-dependent-and-sex-specific-phenotypic-plasticity-heliconius). *BioRxiv*, 2024.10.22.619625.



 

 

 

- [ descriptionPublisher's Version](https://www.biorxiv.org/content/biorxiv/early/2024/10/22/2024.10.22.619625.full.pdf)
 
- [ descriptionPublisher's Version](https://www.biorxiv.org/content/biorxiv/early/2024/10/22/2024.10.22.619625.full.pdf)
 
 

 



 

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##  New papers from us 

**Hybrid speciation in the Amazon.** Rosser, N., Seixas, F., et al. 2024. Hybrid speciation driven by multilocus introgression of ecological traits. Nature 628:811-817. <https://doi.org/10.1038/s41586-024-07263-w>

   ![hybrid  speciation](/sites/g/files/omnuum6606/files/styles/hwp_1_1__720x720_scale/public/malletlab/files/untitled-1sm.jpg?itok=bmtW3QgY) 

 

**Halloween cover of PNAS, 31 October 2023.** [Tianzhu Xiong, et al.](/file_url/293) Hybrid sterility in female butterflies is best explained by a polygenic model of inheritance on Z chromosome and autosomes.

   ![Butterfly oocyte](/sites/g/files/omnuum6606/files/styles/hwp_1_1__360x360_scale/public/malletlab/files/papilio_ovary.jpg?itok=5i6HYFFK) 

 

**Cover of Science, Nov. 1, 2019**.  
The issue features [our paper on genomic evidence for gene flow among species](https://mallet.oeb.harvard.edu/publications/genomic-architecture-and-introgression-shape-butterfly-radiation-science). Photo by Andrew Neild.

   ![Science cover, Nov. 1, 2019](/sites/g/files/omnuum6606/files/styles/hwp_1_1__360x360_scale/public/malletlab/files/cover_med_v2.jpg?itok=TMqWWGPM) 

 

**A cryptic species discovered in** ***Heliconius*** **!** It is not always the case that mtDNA 'barcode' differences correctly delimit separate species. However, we recently found two cryptic *Heliconius* species that co-occur in sympatry in a narrow zone of overlap in Amazonia, initially via barcoding. Furthermore, the two taxa are co-mimics, so no mimicry switch led to speciation here, although we had thought that mimicry switches typically accompanied speciation in the genus *Heliconius.* [Rosser et al. 2019](https://doi.org/10.1093/zoolinnean/zly046).

   ![Heliconius demeter vs. eratosignis](/sites/g/files/omnuum6606/files/styles/hwp_1_1__360x360_scale/public/malletlab/files/demeter_widget.jpg?itok=grOI8byU) 

 

  
**What prevents hybrid zones from moving?** (a), top above. Could be there's a density trough (b), or a cline at another ecologically relevant locus that interferes (c)? We argue that it's more likely that a high rainfall peak (d), bottom at the edge of the Andes traps butterfly hybrid zones, to form a massive suture zone in many butterfly taxa. (See [Rosser et al. 2014](http://www.ucl.ac.uk/taxome/jim/pap/Rosser%20et%20al%202014%20hybrid%20zones.pdf)).

   ![moving clines](/sites/g/files/omnuum6606/files/styles/hwp_1_1__360x360_scale/public/malletlab/files/fig3_50.jpg?itok=T4XrPVSn) 

 

  
**Evidence for adaptive introgression at the B/D mimicry locus**  
**a**, Genetic divergence between *H. melpomene races aglaope* (rayed) and *amaryllis* (postman) across a hybrid zone in northeast Peru. Divergence, *FST*, measured along the *B/D* genomic region peaks in the locus that controls red wing pattern elements between the genes *kinesin* and *optix*. **b**, **c**, Distribution of fixed ABBA and BABA sites (see above) along *B/D*region for two comparisons between *H. melpomene* and *H. timareta*. Excesses of ABBA in **b** and BABA in **c** are highly significant, indicating introgression of colour pattern control region. From Fig. 4 of [Heliconius Genome Consortium 2012](https://doi.org/10.1038/nature11041)

   ![from Heliconius Nature 2012 Fig. 4](/sites/g/files/omnuum6606/files/styles/hwp_1_1__360x360_scale/public/malletlab/files/natfigred.jpg?itok=ObkXv1s-) 

 

  
**Selection against melanism in UK after the Clean Air Act.** Survival of peppered moth morphs (+/- SE) over the course of six years of the predation experiment. Unfilled diamonds with dashed lines, non-melanic; filled squares with solid lines, melanic. From Fig. 1 of [Laurence Cook et al. 2012](//doi.org/10.1098/rsbl.2011.1136)

   ![fig1_sm.jpeg](/sites/g/files/omnuum6606/files/styles/hwp_1_1__360x360_scale/public/malletlab/files/fig1_sm.jpeg?itok=tqFwUcpQ)