In Drosophila miranda, a chromosome fusion between the Y chromosome and the
autosome corresponding to Muller's element Chas created a new sex chromoso
me system. The chromosome attached to the ancestral Y chromosome is transmi
tted paternally and hence is not exposed to crossing over. This chromosome.
conventionally called the neo-Y, and the homologous neo-X chromosome displ
ay many properties of evolving sex chromosomes, We report here the transpos
ition of the exuperantial (exu1) locus from a neo-sex chromosome to the anc
estral X chromosome of D, miranda. Exu1 is known to have several critical d
evelopmental functions, including a male-specific role in spermatogenesis.
The ancestral location of exu1 is conserved in the sibling species of D. mi
randa, as well as in a more distantly related species. The transposition of
exu1 can be interpreted as an adaptive fixation, driven by a selective adv
antage conferred by its effect oil dosage compensation. This explanation is
supported by the pattern of within-species sequence variation at exu1 and
the nearby exu2 locus. The implications of this phenomenon for genome evolu
tion are discussed.