BirdIslandGenomic is three years project funded by ANR .
Abstract of the Project
In this project, we propose to investigate island evolution from the genomic point-of-view. Understanding the influence of population size variation on molecular evolution is currently a major topic in the field. Since island species have evolved in isolated and small populations, they provide an unique opportunity to study the impact of non-adaptive forces on biological evolution. Here we propose to gather a population genomic dataset composed of two island/mainland pairs of species, using endemic birds of Réunion Island and their mainland south-Africa relative as model organisms. Indeed, we are convinced that mainland/island comparison provide one of the best empirical set-up to study the impact of non-adaptive forces on genome evolution in natural population. Four completely sequenced genomes, eight transcriptomes and a genome-wide polymorphism composed of forty individuals will be used to unravel the influence of population size on the accumulation of deleterious mutations in the genome. Our goal will be to establish if different regions of the genome are differentially affected by the reduction of population size. We will also aim to evaluate the evolution of genome architecture (e.g., on genome size) will be compared between the mainland and the island species. According to recent theory, the evolution of genome architecture could also be influence by non-adaptive force such as genetic drift. So far, however, most of the empirical evidences come from comparison between distantly related species. We argue that the mainland/island comparison provide an unique opportunity to study the micro-evolution of genome architecture and to test the importance of genetic drift in the emergence of genomic complexity. Finally, the very large and comprehensive datasets generate during this project will also be used to tackle two fundamental topics of evolutionary biology: i) the evolution of sex chromosomes and ii) the genomic of speciation.
The Mascarene white-eyes (Zosterops) as model species
In our project, we propose to focus on the small oceanic island of Réunion. This island is situated near the Tropic of Capricorn, 700km east of Madagascar. It is a relatively small volcanic island with an area of ~2,500km (three and a half times smaller than Corsica) forming a loose archipelago, called the Mascarene archipelago, with Mauritius (at ~200km) and Rodrigues (at ~800km). Réunion and Mauritius are volcanic Islands that have never been connected to any large land-mass. Their great isolation is confirmed by the high degree of endemism found in the archipelago (e.g., 85% of the terrestrial birds are endemic) (Thébaud et al. 2009). Among the seven extant terrestrial (near-)endemic bird species, four represent good candidates for this project. These are the Réunion Stonechat (Saxicola tectes), the Réunion Bulbul (Hypsipetes borbonicus), the Mascarene White-eye (Zosterops borbonicus) and the Réunion Olive White-eye (Z. olivaceus). Among the endemic birds of the Réunion Island, the White-eyes stands because i) its phylogeny is well documented (Figure 1), ii) the closest mainland relatives are common birds that have large repartition ranges (suggesting a large population size) in Africa ( the African Yellow White-eye (Z. senegalensis) and the Cape White-eye (Z. pallidus)) and iii) the Mascarene White-eye is currently studied by a French team leaded by Pr. Christophe Thébaut from the University of Toulouse (Milá et al. 2010).
From left to right: brown, gray-headed brown, and gray forms of the Reunion gray white-eye. (c) Photos: Borja Milá
According to molecular dating, the colonization of white-eyes in the Mascarene archipelago occurred around 1.5 to 2 millions of years (My) ago from an African or an Asian ancestor (Warren et al. 2006, Moyle et al. 2009). Interestingly, the two species endemic from the Réunion island have a sister (sub-)species on Mauritius. The two pairs are called the “Mascarene grey white-eyes” (Z. b. borbonicus and Z. b. mauritianus) and the “Mascarene olive white-eyes” (Z. o. olivaceus and Z. o. chloronothos). All the four taxa form a distinct monophyletic clade (Figure 1). This phylogeny suggests either a single colonization event followed by a species exchange between islands or an independent colonization of the two species followed by an inter-island dispersal (Warren et al. 2006). Either ways, their history guarantees that the divergence within the olive and the grey species has occurred on an island with small population size. Warren et al. (2006) dated this divergence around 0.5 My whereas the divergence between the grey and the olive white-eyes is dated to be 1.2 My old. Therefore, our candidate species have evolved on small islands since at least half a million years and maybe as long as 1 My. Moreover, the distance between the island and the nearest continent (700 km) probably prevented gene flow to occur after colonization. This isolation is important because gene flow could mitigates the effect of genetic drift by increasing genetic variance and bringing new beneficial mutations.
You can learn more about the Réunion Zosterops at zoboscience web page.
Figure 1 : Phylogeny and repartition range of the candidate species. The phylogeny is based on ATP6 sequences obtain from Oatley et al. (2012) for the southern African species and Warren et al. (2006) for the islands species. Stars indicate bootstrap support >80%. Repartition range was obtained from IUCN/red list map (http://www.iucnredlist.org).
Collaborators
References
- Milá B, Warren BH, Heeb P, Thébaud C. 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 Evol. Biol. 10:158.
- Oatley G, Voelker G, Crowe TM, Bowie RC. 2012. A multi-locus phylogeny reveals a complex pattern of diversification related to climate and habitat heterogeneity in southern African white-eyes. Mol. Phylogenet. Evol. 64:633–644.
- Warren BH, Bermingham E, Prys-Jones RP, Thébaud C. 2006. Immigration, species radiation and extinction in a highly diverse songbird lineage: white-eyes on Indian Ocean islands. Mol. Ecol. 15:3769–3786.
- Moyle RG, Filardi CE, Smith CE, Diamond J. 2009. Explosive Pleistocene diversification and hemispheric expansion of a “great speciator.” Proc. Natl. Acad. Sci. 106:1863–1868