that in recent historical time, i.e., Pleistocene, the geographic ranges of D. melanogaster and D. simulans were completely disjunct. Hence, the two cryptic species are assumed to have been allopatric species in the Afrotropical region, and indeed to some extent they remain so. Drosophila erectaDrosophila erecta has been recorded from the south and mid Ivory Coast: Yale at the bottom (400 m) of Mt. Nimba at the border of Guinea,

The finding of new melanogaster sister species may help us in understanding more about how the emergence of genetic novelties, particularly in insular habitats, can result in speciation. Here we report on the discovery of Drosophila santomea, which is the first melanogaster sibling found off West-equatorial Africa, on São Tomé, one of the Gulf of Guinea islands. Although the eight other melanogaster sister species are remarkably conservative in their morphology except for their terminalia, the new find has a morphological trait distinguishing it from all of these: a pure yellow body coloration of both sexes without the normal black abdominal banding. Evidence from the terminalia, polytene and mitotic chromosomes, period gene and allozymes are provided indicating that it is nonetheless the nearest relative of Drosophila yakuba with which it coexists on the island. The new find is a clear-cut taxon as shown by the production of sterile male hybrids, eventually with developmental defects, in both directions of cross with yakuba and by the existence of an altitudinal divide accompanied by a hybrid zone at mid-elevation on the island. Molecular and karyotypic data further support this conclusion. In contrast to the significant divergence of their nuclear DNAs, an intriguing similarity in their cytochrome b sequences was observed indicating a recent coalescence common to santomea, yakuba and also teissieri cytoplasms. These were shown to harbour the same Wolbachia endosymbiotic bacteria which could possibly be responsible for mitochondrial DNA hitchhiking across the species barrier.

Drosophila yakuba is widely distributed in sub-Saharan Africa, while D. santomea is endemic to the volcanic island of São Tomé in the Atlantic Ocean, 280 km west of Gabon. On São Tomé, D. yakuba is found mainly in open lowland forests, and D. santomea is restricted to the wet misty forests at higher elevations. At intermediate elevations, the species form a hybrid zone where hybrids occur at a frequency of 1%. To determine the extent of gene flow between these species we studied polymorphism and divergence patterns in 29 regions distributed throughout the genome, including mtDNA and three genes on the Y chromosome. This multilocus approach, together with the comparison to the two allopatric species D. mauritiana and D. sechellia, allowed us to distinguish between forces that should affect all genes and forces that should act on some genes (e.g., introgression). Our results show that D. yakuba mtDNA has replaced that of D. santomea and that there is also significant introgression for two nuclear genes, yellow and salr. The majority of genes, however, has remained distinct. These two species therefore do not form a ''hybrid swarm'' in which much of the genome shows substantial introgression while disruptive selection maintains distinctness for only a few traits (e.g., pigmentation and male genitalia).

In this study, the phylogenetic relationships of 164 species of the family Drosophilidae are discussed, using the Amyrel gene, a member of the a-amylase multigene family. This study focuses on numerous species groups in the subgenera Sophophora and Drosophila of the genus Drosophila but also includes other closely related genera. Nucleotide data were analysed by several methods: maximum parsimony, neighbour joining, maximum likelihood and Bayesian inference. Heterogeneity of base composition (mainly low GC contents in the species groups willistoni and saltans) has been addressed. In all analyses, the genus Drosophila appeared paraphyletic. The subgenus Sophophora clearly appeared to be a monophyletic group, showing well-resolved clades, with the Neotropical groups arising in a basal position. Here, it is proposed to raise the species subgroups ananassae and montium to the rank of species group, and to restrict the melanogaster species group to the melanogaster subgroup plus the ÔOrientalÕ subgroups, among which the suzukii subgroup is polyphyletic. Some related genera such as Zaprionus, Liodrosophila, Scaptomyza and Hirtodrosophila are clustered with, or inside the subgenus Drosophila, which is therefore paraphyletic and should be reviewed.