Speciation Research on the Reunion Gray White-Eye (Zosterops borbonicus)


Welcome to our site on Reunion Island's Gray White-eye (Zosterops borbonicus), an excellent model species to study the evolutionary process of species formation.

The Gray White-eye presents a striking, geographically structured plumage color polymorphism on the island of Reunion, and we are using genetic and phenotypic analyses to investigate the evolutionary mechanisms driving lineage divergence.

We are a group of researchers at the Laboratoire Evolution et Diversité Biologique (EDB) at the University of Toulouse (France) and the National Museum of Natural Sciences of the Spanish Research Council (Spain).

Below you will find an introduction to our study system, contact information and a list of publications to find out more.


From left to right: brown, gray-headed brown, and gray forms of the Reunion gray white-eye. (c) Photos: Borja Milá


Watch this short video about the project!




Plumage forms within the gray white-eye and their distribution on the islands of Reunion and Mauritius. A. Gray form (highlands); B. Gray headed brown form (eastern lowlands); C. Brown naped form (southern lowlands); D. Brown form (divided into two different forms, highland brown and lowland brown). On Mauritius there is only one gray-colored form.






The island of Reunion has a rough topography and a striking diversity of habitat types for an island of its size.












Some habitat types on Reunion Island:







Scientific publications (updated 2017-05-30):



Bourgeois, Y.X.C., B. Delahaie, M. Gautier, E. Lhuillier, P.-J. G. Malé, J. A.M. Bertrand, J. Cornuault, K. Wakamatsu, O. Bouchez, C. Mould, J. Bruxaux, H. Holota, B. Milá*, C. Thébaud*. 2017. A novel locus on chromosome 1 underlies the evolution of a melanic plumage polymorphism in a wild songbird. Royal Society Open Science, 4(2):160805. *: senior co-authors.


Bourgeois, Y.X.C, J. A.M. Bertrand, B. Delahaie, J. Cornuault, T. Duval, B. Milá*, C. Thébaud*. 2016. Candidate gene analysis suggests untapped genetic complexity in melanin-based pigmentation in birds. Journal of Heredity, 2016:327-335.  *: senior co-authors


Bertrand, J. A. M., B. Delahaie, Y. X. C. Bourgeois, T. Duval, R. García-Jiménez, J. Cornuault, B. Pujol, C. Thébaud, B. Milá. 2016. The role of selection and historical factors in driving population differentiation along an elevational gradient in an island passerine bird. Journal of Evolutionary Biology, 29:824-836.


Bertrand, J. A. M., Y. X. C. Bourgeois, C. Thébaud. 2016. Population density of the Reunion grey white-eye Zosterops borbonicus within the summit ecosystems of Reunion, Mascarene Islands. Ostrich, 87(1):85-88.


Cornuault, J., B. Delahaie, J.A.M. Bertrand, Y.X.C. Bourgeois, B. Milá, P. Heeb, C. Thébaud. 2015. Morphological and plumage colour variation in the Réunion grey white-eye (Aves: Zosterops borbonicus): assessing the role of selection. Biological Journal of the Linnean Society, 114(2):459-473.


Bertrand, J. A. M., Y. X. Bourgeois, B. Delahaie, T. Duval, R. Garcia-Jiménez, J. Cornuault, P. Heeb, B. Milá, B. Pujol, C. Thébaud. 2014. Extremely reduced dispersal and gene flow in an island bird. Heredity, 112:190-196.


Cornuault, J., J. A. M. Bertrand, B. Warren, B. Milá, C. Thébaud, P. Heeb. 2013. Timing and number of colonizations but not diversification rates affect diversity patterns in hemosporidian lineages on a remote oceanic archipelago. American Naturalist, 182:820-833.


Bourgeois, Y. X. C., E. Lhuillier, T. Cézard, J. A. M. Bertrand, B. Delahaie, J. Cornuault, T. Duval, O. Bouchez, B. Milá, C. Thébaud. 2013. Mass production of SNP markers in a nonmodel passerine bird through RAD sequencing and contig mapping to the zebra finch genome. Molecular Ecology Resources, 13(5):899-907.


Cornuault, J., A. Khimoun, R. J. Harrigan, Y. Bourgeois, B. Milá, C. Thébaud, P. Heeb. 2013. The role of ecology in the geographical separation of blood parasites infecting an insular bird. Journal of Biogeography, 40(7):1313-1323.


Bourgeois, Y., J. A.M. Bertrand, C. Thébaud, B. Milá. 2012. Investigating the role of the Melanocortin-1 receptor gene in an extreme case of microgeographical variation in the pattern of melanin-based plumage pigmentation. PLOS ONE, 7(12):e50906.


Cornuault, J., A. Bataillard, B. Warren, A. Lootvoet, P. Mirleau, T. Duval, B. Milá, C. Thébaud, P. Heeb. 2012. The role of immigration and in-situ radiation in explaining blood parasite assemblages in an island bird clade. Molecular Ecology, 21:1438-1452.


Bertrand, J. A. M., R. García-Jiménez, Y. Bourgeois, T. Duval, P. Heeb, C. Thébaud, B. Milá. 2012. Isolation and characterization of twelve polymorphic microsatellite loci for investigating an extreme case of microgeographical variation in an island bird (Zosterops borbonicus). Conservation Genetics Resources, 4(2):323-326.


Milá, B., B. H. Warren, P. Heeb, C. Thébaud. 2010. The geographic scale of diversification on islands: genetic and morphological divergence at a very small spatial scale in the Mascarene grey white-eye (Aves: Zosterops borbonicus). BMC Evolutionary Biology, 10:158.



Outreach publications:


Bertrand, J. A. M., B. Delahaie, Y. X. C. Bourgeois, C. Thébaud. 2013. Comment naissent de nouvelles espèces?  L’Oiseau Magazine, No. 112. Magazine of the League pour la Protection des Oiseaux (League for the Protection of Birds), France (http://www.lpo.fr/)







Support provided by:

National Geographic Society, USA
Agence Nationale de la Recherche, France
Institut Français pour la Biodiversité, France
Consejo Superior de Investigaciones Científicas, Spain





For more information on the project contact:

Dr. Christophe Thébaud at thebaud.weebly.com/

Dr. Borja Milá at www.borjamila.com

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Some members of the team: Christophe Thébaud, Borja Milá, Joris Bertrand and Boris Delahaie.