SEX-CHROMOSOME MEIOTIC DRIVE IN STALK-EYED FLIES

Meiotically driven sex chromosomes can quickly spread to fixation and cause population extinction unless balanced by selection or suppressed by genetic modifiers. We report results of genetic analyses that demo nstrate that extreme female-biased sex ratios in two sister species of stalk-eyed flies, Cyrtodiopsis dalmanni and C. whitei, are due to a m eiotic drive element on the X chromosome (X-d). Relatively high freque ncies of X-d in C. dalmanni and C. whitei (13-17% and 29%, respectivel y) cause female-biased sex ratios in natural populations of both speci es. Sex ratio distortion is associated with spermatid degeneration in male carriers of X-d. Variation in sex ratios is caused by Y-linked an d autosomal factors that decrease the intensity of meiotic drive. Y-li nked polymorphism for resistance to drive exists in C. dalmanni in whi ch a resistant Y chromosome reduces the intensity and reverses the dir ection of meiotic drive. When paired with X-d, modifying Y chromosomes (Y-m) cause the transmission of predominantly Y-bearing sperm, and on average, production of 63% male progeny. The absence of sex ratio dis tortion in closely related monomorphic outgroup species suggests that this meiotic drive system may predate the origin of C. whitei and C. d almanni. We discuss factors likely to be involved in the persistence o f these sex linked polymorphisms and consider the impact of X-d on the operational sex ratio and the intensity of sexual selection in these extremely sexually dimorphic flies.