African populations of Drosophila simulans are thought to be ancestral in this model species and are increasingly used for testing general hypotheses in evolutionary genetics. It is often assumed that African populations are more likely to be at a neutral mutation drift equilibrium than other populations. Here we examine population structuring and the demographic profile in nine populations of D. simulans. We surveyed sequence variation in four X-linked genes (runt, sevenless, Sex-lethal, and vermilion) that have been used in a parallel study in the closely related species D. melanogaster. We found that an eastern group of populations from continental Africa and Indian Ocean islands (Kenya, Tanzania, Madagascar, and Mayotte Island) is widespread, shows little differentiation, and has probably undergone demographic expansion. The other two African populations surveyed (Cameroon and Zimbabwe) show no evidence of population expansion and are markedly differentiated from each other as well as from the populations from the eastern group. Two other populations, Europe and Antilles, are probably recent invaders to these areas. The Antilles population is probably derived from Europe through a substantial bottleneck. The history of these populations should be taken into account when drawing general conclusions from variation patterns.
DROSOPHILA melanogaster and D. simulans are two cosmopolitan sibling species widely used in evolutionary studies. Despite their recent common ancestry, they show surprising differences in genomic parameters. Sequence variation parameters differ across chromosomes, across species, and between African and European populations. A number of hypotheses involving differences in history, in effective population size, in life history traits, and in chromosome inversion frequencies have been alleged for explaining these contrasts (see, e.g., Begun 1996; Moriyama and Powell 1996;Irvin et al. 1998;Andolfatto and Przeworski 2000;Andolfatto 2001;Charlesworth 2001;Capy and Gibert 2004;Gravot et al. 2004). A long and established history of studies has made these two species excellent models for evolutionary analyses. They have independently undergone a similar evolution and are thus considered useful systems for studying the critical factors involved in genetic evolution. Despite these strengths as model systems, population histories can add unique, contingent factors to the patterns of variation that can obscure patterns of change if they are not accounted for in data interpretation.Biometrical studies conducted in D. melanogaster by Teissier (1957) showed long ago that this species is geographically differentiated. More recent studies based on molecular markers have confirmed that African D. melanogaster populations are genetically differentiated (Bénassi and Veuille 1995;Michalakis and Veuille 1996;Veuille et al. 1998;Baudry et al. 2004). In D. simulans, Hamblin and Veuille (1999) studied two genes, vermilion (v) and Glucose-6 phosphate dehydrogenase (G6pd), in several populations from the Old ...