Maisonneuve L, Chouteau M, Joron M, Llaurens V (submitted) How heterozygote advantage can promote the evolution of disassortative mating and shape the underlying genetic architecture.
ddoi:10.1101/616409 [BioRxiv]

Saenko SV, Chouteau M, Piron-Prunier F, Blugeon C, Joron M & Llaurens V (submitted). Unravelling the genes forming the wing pattern supergene in the polymorphic butterfly.
Heliconius numata.

Mérot C, Debat V, Le Poul Y, Merrill R, Naisbit R, Tholance A, Jiggins C, Joron M (submitted) Hybridization and transgressive exploration of wing morphology in Heliconius butterflies.
PDF | doi:xxx [BioRxiv]

Aubier T, Kokko H & Joron M (submitted) Coevolution of male and female mate choice can destabilize reproductive isolation. BioRxiv preprint
PDF | doi:10.1101/246355 [BioRxiv]

Edelman NB, Frandsen P, Miyagi M, Clavijo B, Davey J, Dikow R, García-Accinelli G, van Belleghem S, Patterson N, Neafsey D, Challis , Kumar S, Moreira G, Salazar C, Chouteau M, Counterman B, Papa R, Blaxter M, Reed RD, Dasmahapatra K, Kronforst, M, Joron M, Jiggins CD, McMillan WO, Di Palma F, Blumberg AJ , Wakeley J, Jaffe D & Mallet J (submitted) Genomic architecture and introgression shape a butterfly radiation. BioRxiv preprint
PDF | doi:10.1101/466292 [BioRxiv]

Kozak KM, Joron M, Jiggins C & McMillan WO (submitted). Rampant genome-wide admixture across the Heliconius radiation. BioRxiv preprint
PDF | doi:10.1101/414201 [BioRxiv]

Arias M, Griffiths D, Joron M, Davey J, Martin SH, Jiggins CD, Nadeau N, and Llaurens V (preprint). Generalisation behaviour of predators toward warning signals displayed by harmful prey: answers from a videogame played by humans. BioRxiv preprint: 409557
PDF | doi:10.1101/409557

Chouteau M, Dezeure J, Sherratt TN, Llaurens V & Joron M. (2019) Similar predator aversion for natural prey with diverse toxicity levels. Animal Behaviour 153:49–59
PDF | doi:10.1016/j.anbehav.2019.04.017


Jay P, Whibley A, Frézal L, Rodríguez de Cara MA, Nowell RW, Mallet J, Dasmahapatra KK & Joron M (2018). Supergene evolution triggered by the introgression of a chromosomal inversion. Current Biology 28:1839–1845
PDF | Supplementary information | doi:10.1016/j.cub.2018.04.072

Rosser N, Freitas AVL, Huertas B, Joron M, Lamas G, Mérot C, Simpson F, Willmott K, Mallet J & Dasmahapatra KK (2018). Cryptic speciation associated with geographic and ecological divergence in two Amazonian Heliconius butterflies . Zoological Journal of the Linnean Society
PDF | Supplementary Material | doi:10.1093/zoolinnean/zly046 – view online


Mérot C, Salazar C, Merrill RM, Jiggins C & Joron M (2017). What shapes the continuum of reproductive isolation? Lessons from Heliconius butterflies. Proceedings of the Royal Society B
PDF | doi:10.1098/rspb.2017.0335

Chouteau M, Llaurens V, Piron-Prunier F & Joron M (2017). Polymorphism at a mimicry supergene maintained by opposing frequency-dependent selection pressures. Proceedings of the National Academy of Sciences, USA 114(31):8325–8329
PDF | doi:10.1073/pnas.1702482114

de Silva DL, Motas L, Chazot N, Mallarino R, Silva-Brandão KL, Gomez LM, Freitas AVL, Lamas G, Joron M, Mallet J, Giraldo CE, Uribe S, Särkinen T, Knapp S, Jiggins CD, Willmott KR, Elias M (2017). North Andean origin and diversification of the largest ithomiine butterfly genus. Scientific Reports 7:45966
PDF | doi:10.1038/srep45966

Llaurens V, Whibley A & Joron M (2017). Genetic architecture and balancing selection: the life and death of differentiated variants (review). Molecular Ecology 26(9):2430–2448
PDF | doi:10.1111/mec.14051

Aubier T, Joron M and Sherratt TN (2017). Mimicry among unequally defended prey should be mutualistic when predators sample optimally. The American Naturalist 189(3):267-282
PDF | doi:10.1086/690121


Chazot N, Willmott K, de Silva DL, Condamine FL, Freitas AVL, Lamas G, Morlon H, Joron M, Jiggins C, Uribe S, Mallet J, Giraldo-Sanchez CE and Elias M (2016). Into the Andes: multiple independent colonizations drive montane diversity in the Neotropical clearwing butterflies Godyridina. Molecular Ecology 25:5765–5784
PDF | doi:10.1111/mec.13773

Nadeau NJ, Pardo-Diaz C, Whibley A, Supple MA, Saenko SV, Wallbank RWR, Wu GC, Maroja L, Ferguson L, Hanly JJ, Hines H, Salazar C, Merrill RM, Dowling AJ, ffrench-Constant RH, Llaurens V, Joron M, McMillan OW, and Jiggins C (2016). The gene cortex controls mimicry and crypsis in butterflies and moths. Nature 534:106–110.
PDF | Full text | doi:10.1038/nature17961

Piron Prunier F, Chouteau M, Whibley A, Joron M and Llaurens V (2016). Selection of valid reference genes for reverse transcription quantitative PCR analysis in Heliconius numata. Journal of Insect Science 16(1):50
PDF | doi:10.1093/jisesa/iew034

Mérot C, Le Poul Y, Théry M and Joron M (2016). Mimicry refinement: Phenotypic variations tracking the local optimum. Journal of Animal Ecology, 85:1056-1069.
PDF | doi:10.1111/1365-2656.12521
Cover of the virtual issue on Evolutionary Ecology (Photo G. Vernade)

Chouteau M, Arias M and Joron M (2016). Warning signals are under positive frequency-dependent selection in nature. Proceedings of the National Academy of Sciences, USA 113(8):2164–2169
PDF (with supporting information) | doi:10.1073/pnas.1519216113

Davey JW, Chouteau M, Barker SL, Maroja L, Baxter SW, Simpson F, Joron M, Mallet J, Dasmahapatra KK and Jiggins CD (2016). Major improvements to the Heliconius melpomene genome assembly used to confirm 10 chromosome fusion events in 6 million years of butterfly evolution. G3: Genes Genomes Genetics 6(3):695-708
PDF | doi:10.1534/g3.115.023655

Wallbank RWR, Baxter SW, Pardo-Diaz C, Hanley JJ, Martin SH, Mallet J, Dasmahapatra KK, Salazar C, Reed R, Joron M, Nadeau NJ, McMillan OW, and Jiggins CD (2016). Evolutionary novelty in a butterfly wing pattern through enhancer shuffling. PLOS Biology 14(1): e1002353.
PDF | Supporting information | doi:10.1371/journal.pbio.1002353


Llaurens V, Joron M and Billiard S (2015). The evolution of dominance and its underlying molecular mechanisms in Müllerian mimicry. Evolution 69(12):3097–3108.
PDF | doi:10.1111/evo.12810

Mérot C, Frérot B, Leppik E and Joron M (2015). Beyond magic traits: multimodal mating cues in Heliconius butterflies. Evolution 69(11):2891-2904.
PDF | doi:10.1111/evo.12789

Merrill RM, Dasmahapatra KK, Davey J, Dell’Aglio D, Hanley JJ, Jiggins C, Joron M, Kozak KM, Llaurens V, Martin SH, Montgomery SH, Morris J, Nadeau NJ, Pinharanda A, Rosser N, Thompson D, Vanjari S, Wallbank RWR, and Yu Q (2015). The diversification of Heliconius butterflies: What have we learned in 150 years? Journal of Evolutionary Biology 28:1417–1438.
PDF | doi:10.1111/jeb.12672

Huber B, Le Poul Y, Whibley A, Navarro N, Martin A, Baxter S, Shah AB, Gilles B, Wirth T, McMillan OW, and Joron M (2015). Conservatism and novelty in the genetic architecture of adaptation in Heliconius butterflies. Heredity, 114:515–524.
PDF, Supplementary information | doi: 10.1038/hdy.2015.22

Chouteau M, Whibley A, Angers B, & Joron M (2015). The development of microsatellite loci from a reference genome in the Neotropical butterfly Heliconius numata and its close relatives. Entomological Science 18(2):283-287.
PDF | doi:10.1111/ens.12121


Le Poul Y, Whibley A, Chouteau M, Prunier F, Llaurens V, & Joron M (2014). Evolution of dominance mechanisms at a butterfly mimicry supergene. Nature Communications 5:5644.
PDF, Supplementary information (PDF) | Full text (Open Access) | doi:10.1038/ncomms6644

Timmermans MJTN, Baxter SW, Clark R, Heckel DG, Vogel H, Collins S, Papanicolaou A, Fuková I, Joron M, Thompson M, Jiggins CD, ffrench-Constant RH, & Vogler AP (2014). Comparative genomics of the mimicry switch in Papilio dardanus. Proceedings of the Royal Society B 281(1787):20140465.
PDF | doi:10.1098/rspb.2014.0465

Llaurens V, M Joron & M Théry (2014). Cryptic differences in color among Müllerian mimics: how can the visual capacities of predators and prey shape the evolution of wing colors?. Journal of Evolutionary Biology 27(3):531-540.
PDF | doi:10.1111/jeb.12317


Llaurens V, S Billiard & M Joron (2013). The effect of dominance on polymorphism in Müllerian mimicry. Journal of Theoretical Biology, 337:101-110.
PDF | doi: 10.1016/j.jtbi.2013.08.006

Jones R, Y Le Poul, A Whibley, C Mérot, R ffrench-Constant & M Joron (2013). Wing shape variation associated with mimicry in butterflies. Evolution 67(8):2323-2334.
PDF | doi: 10.1111/evo.12114

Mérot C, J Mavárez, KK Dasmahapatra, J Mallet, G Lamas M., and M Joron (2013). Genetic differentiation without mimicry shift in a pair of hybridizing Heliconius species. Biological Journal of the Linnean Society 109(4):830-847.
PDF | doi: 10.1111/bij.12091


The Heliconius Genome Consortium (2012). Butterfly genome reveals promiscuous exchange of mimicry adaptations among species. Nature 487:94-98
PDF | Full text (Open Access) | Supplementary information | doi: 10.1038/nature11041
Communiqué de presse (CNRS) | UK press release (UCL site).
Faculty of 1000 « Must read » | News of the Week at Science | Le Figaro newspaper report

Pardo-Diaz C, C Salazar, SW Baxter, C Mérot, W Figueiredo-Ready, M Joron, WO McMillan, and CD Jiggins (2012). Adaptive introgression across species boundaries in Heliconius butterflies. PLoS Genetics 8(6):e1002752.
PDF | doi: 10.1371/journal.pgen.1002752 | PLoS Perspective by Marcus Kronforst

Jones RT, PA Salazar, RH ffrench-Constant, CD Jiggins & M Joron (2012). Evolution of a mimicry supergene from a multilocus architecture. Proceedings of the Royal Society B 279:316-325.
PDF | doi: 10.1098/rspb.2011.0882

Nadeau NJ, A Whibley, RT Jones, J Davey, KK Dasmahapatra, SW Baxter, M Joron, RH ffrench-Constant, ML Blaxter, J Mallet & CD Jiggins (2012). Genomic islands in hybridizing Heliconius butterflies identified by large-scale targeted sequencing. Philosophical Transactions of the Royal Society B 367:343-353
PDF | doi: 10.1098/rstb.2011.0198


Joron M, L Frezal, RT Jones, N Chamberlain, SF Lee, CR Haag, A Whibley, M Becuwe, SW Baxter, L Ferguson, PA Wilkinson, C Salazar, C Davidson, R Clark, MA Quail, H Beasley, R Glithero, C Lloyd, S Sims, M Jones, J Rogers, C Jiggins, and RH ffrench-Constant (2011) Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry. Nature 477:203-206
PDF | Supplementary information | doi: 10.1038/nature10341
Faculty of 1000 top 2% in biology and medicine | Great 3-min NatureVideo overview | New York Times article | More press coverage

Joron M & A Whibley (2011). Stripes, sex and sparrows: what processes underlie heteromorphic chromosome evolution? Heredity 106:531-532
PDF | doi: 10.1038/hdy.2010.106


Wu GC, M Joron & CD Jiggins (2010). Signatures of selection in loci governing major colour patterns in Heliconius butterflies and related species. BMC Evoutionary Biology 10:368, « Highly accessed »
PDF | doi: 10.1186/1471-2148-10-368

Baxter SW, NJ Nadeau, L Maroja, PA Wilkinson, BA Counterman, A Dawson, M Beltrán, S Pérez-Espona, N Chamberlain, L Ferguson, R Clark, C Davidson, R Glithero, J Mallet, WO McMillan, MR Kronforst, M Joron, R H. ffrench-Constant & CD Jiggins (2010). Genomic hotspots for adaptation: the population genetics of Müllerian mimicry in the Heliconius melpomene clade. PLoS Genetics 6: e1000794.
PDF | doi: 10.1371/journal.pgen.1000794 | PLoS Primer by Arnaud Martin et al.

Counterman BA, F Araujo-Pérez, HM Hines, SW Baxter, CM Morrison, DP Lindstrom, R Papa, L Ferguson, M Joron, RH ffrench-Constant, C Smith, D Nielsen, R Chen, CD Jiggins, RD Reed, G Halder, J Mallet, and WO McMillan (2010). Genomic hotspots for adaptation: the population genetics of Müllerian mimicry in Heliconius erato. PLoS Genetics 6: e1000796.
PDF | doi: 10.1371/journal.pgen.1000796 | PLoS Primer by Arnaud Martin et al.

Ferguson L, SF Lee, N Chamberlain, N Nadeau, M Joron, SW Baxter, P Wilkinson, A Papanicolaou, S Kumar, TJ Kee, R Clark, C Davidson, H Beasley, A Surridge, G Wu, H Vogel, RH ffrench-Constant & CD Jiggins (2010). Characterization of a hotspot for mimicry: Assembly of a butterfly wing transcriptome to genomic sequence at the HmYb/Sb locus. Molecular Ecology 19: 240-254.
PDF | doi: 10.1111/j.1365-294X.2009.04475.x | Special Issue: Next Generation Molecular Ecology


Elias M, M Joron, K Willmott, V Kaiser, K Silva-Brandão, C Arias, LM Gómez Piñeres, S Uribe, AVZ Brower, AVL Freitas & CD Jiggins (2009). Out of the Andes: patterns of diversification in clearwing butterflies. Molecular Ecology 18: 1716-1729.


Joron M (2008). Batesian mimicry: Can a leopard change its spots -and get them back? Current Biology 18 (11): r476-478.

Baxter SW, R Papa, N Chamberlain, SJ Humphray, M Joron, RH ffrench-Constant, WO McMillan & CD Jiggins (2008). Parallel evolution in the genetic basis of Müllerian mimicry in Heliconius butterflies. Genetics 180: 1567-1577.

Joron M, R Papa, J Mallet, et al. (2007) [Meeting Abstract]. Conserved but flexible: Genetic control of mimicry in Heliconius butterfly wing patterns. Journal of Insect Science 7: 24.

Elias M, M Joron, K Willmott, et al. (2007) [Meeting Abstract]. Phylogenetic hypothesis, pattern of speciation and evolution of wing pattern in neotropical Napeogenes butterflies (Lepidoptera : Nymphalidae). Journal of Insect Science 7: 13-14.


Joron M, R Papa, M Beltrán, N Chamberlain, J Mavárez, S Baxter, EB Bermingham, S Humphray, J Rogers, H Beasley, K, Barlow, RH ffrench-Constant, J Mallet, WO McMillan & CD Jiggins (2006). A conserved supergene locus controls wing pattern diversity in Heliconius butterflies. PLoS Biology 4(10): e303.
PDF (3.5 Mb) | doi: 10.1371/journal.pbio.0040303 | PLoS Primer by Axel Meyer | Media coverage

Joron M, CD Jiggins, A Papanicolaou & WO McMillan (2006). Heliconius wing patterns: an evo-devo model for understanding phenotypic diversity. Heredity. 97: 157 – 167 (Evo-devo Special issue).
PDF | Heredity podcast


Whinnett A, M Zimmermann, KR Willmott, N Herrera, R Mallarino, F Simpson, M Joron, G Lamas & J Mallet (2005). Strikingly variable divergence times inferred across an Amazonian butterfly ‘suture zone’. Proceedings of the Royal Society B 272: 2525-2533.

Joron M & Y Iwasa (2005). The evolution of a Müllerian mimic in a spatially-distributed community. Journal of Theoretical Biology 237: 87-103.
PDF | Selected as Must Read by Faculty of 1000 Biology (F1000 Factor 6.0)

Papanicolaou A, M Joron, W O McMillan, M L Blaxter & C D Jiggins (2005). Genomic tools and cDNA derived markers for butterflies. Molecular Ecology 14: 2883-2897.

Joron M (2005). Polymorphic mimicry, microhabitat use, and sex-specific behaviour. Journal of Evolutionary Biology 18: 547-556.

2003 and before

Joron M & PM Brakefield (2003). Captivity masks inbreeding effects on male mating success in butterflies. Nature 424: 191-194.
PDF | Supplementary information

Martin JL & M Joron (2003). Nest predation in forest birds: influence of predator type and predator’s habitat quality. Oikos 102(3): 641-653.

Joron M, IR Wynne, G Lamas & J Mallet (2001). Variable selection and the coexistence of multiple mimetic forms of the butterfly Heliconius numata. Evolutionary Ecology 13: 721-754 [dated 1999 by publisher].

Mallet J & M Joron (1999). Evolution of diversity in warning color and mimicry: polymorphisms, shifting balance, and speciation. Annual Review of Ecology and Systematics 30: 201-233.

Joron M & J Mallet (1998). Diversity in mimicry: paradox or paradigm? Trends in Ecology and Evolution 13: 461-466.

Book chapters

Joron M (in press) Se ressembler pour se protéger: le mimétisme (Ch. 5.7) In 101 secrets du vivant, l’ADN des écosystèmes (D. Faure, D Joly & S Salamitou, Eds), CNRS Editions, Paris.

Joly D & Joron M (in press) L’évolution des espèces décryptée par la lecture des chromosomes (Ch. 3.1) In 101 secrets du vivant, l’ADN des écosystèmes (D. Faure, D Joly & S Salamitou, Eds), CNRS Editions, Paris.

Joron M, X Vekemans, F. Depaulis, D. Eveillard, G. Farrant, L. Garczarek, S. Merlot, F. Partensky, H Roest-Crollius, C Smadja (2016). Evolution et adaptation des gènes et des génomes. In La Génomique Environnementale, la révolution du séquençage à haut débit (D Faure and D Joly, Eds), 192pp, ISTE Editions, London UK, 89-104

Elias M & Joron M (2015). Mimicry in Heliconius and Ithomiini butterflies: the profound consequences of an adaptation. In Origins – BIO Web of Conferences (M.-C. Maurel & P. Grandcolas, Eds). EDP sciences, Paris.
PDF | doi: 10.1051/bioconf/20150400008

Joron M & X Vekemans (2013) Étude de l’évolution adaptative des gènes et des génomes. In Cahier de Prospective en Génomique Environnementale, 100pp. CNRS Institut Ecologie et Environnement n°5

Brakefield PM & M Joron (2010). The flexibility of butterfly wing patterns and evolution in morphospace. In In Search of the Causes of Evolution : From Field Observations to Mechanisms (P.R. Grant and B.R. Grant, Eds), Princeton University Press, Princeton NJ, 93-114.

Joron M (2009). Mimicry. In Encyclopedia of Insects, 2nd Edition (R.T. Cardé & V.H. Resh, Eds), pp. 633-643. Academic Press, New York.

Joron M (2009). Aposematic coloration. In Encyclopedia of Insects, 2nd Edition (R.T. Cardé & V.H. Resh, Eds), pp. 33-38. Academic Press, New York.

Joron M (2003). Mimicry. In Encyclopedia of Insects (R.T. Cardé & V.H. Resh, Eds), pp.714-726. Academic Press, New York.

Joron M (2003). Aposematic coloration. In Encyclopedia of Insects (R.T. Cardé & V.H. Resh, Eds), pp.39-45. Academic Press, New York.


Charlesworth B, et al (1999). There is no new evidence that undermines evolution. Nature 444: 680-680.

Joron M & J Mallet (1999). Diversity in Mimicry: Reply. Trends in Ecology and Evolution 14: 151.


Joron M (2012). Déterminants génétiques et écologiques de la diversité adaptative, Habilitation a Diriger des Recherches, Université Paris-Sud, Orsay, France

Joron M (2000). Aposematic coloration and Müllerian mimicry: the diversification puzzle, Thèse de Doctorat, Université de Montpellier, Ecole Doctorale « Biologie de l’Evolution et Ecologie », Montpellier, France