Reinforcement in hybrids among once isolated semispecies ofDanaus chrysippus(L.) and evidence for sex chromosome evolution

1. The aim of this paper is to investigate mechanisms of reinforcement between two semi-isolated semispecies of the African savannah butterfly Danaus chrysippus. The biogeography of colour genes suggests that four semispecies evolved in once isolated refugia. They expanded their ranges in response to Holocene climatic changes to form a hybrid zone in central-east Africa. 2. Danaus chrysippus is a superspecies within which cycles of alternating cladogenesis and reticulation among semispecies have probably operated over some 4 Myr. Semispecies are inter-fertile but show Haldane rule effects in crosses; gene flow is massive but subject to isolation by distance. 3. One semispecies shows linkage disequilibrium, vis- a-vis others, for haplotype, karyotype (W-linkage of colour genes which function as reproductive isolating barriers) and all-female broods caused by a male-killer endosymbiont. Introgression of colour genes between D. c. dorippus and D. c. chrysippus is constrained by sex linkage and male killing. 4. Reinforcement in hybrid zones comprises allochronic migration, assortative mating, (assumed) sex chromosome incompatibility and sex-ratio distortion. Gene introgression from D. c. dorippus to other semispecies is maintained by a high frequency of backcrossing between hybrid males and females of the latter.