Mammalian sexual differentiation: lessons from the spotted hyena.

Female spotted hyenas (Crocuta crocuta) are the only female mammals that lack an external vaginal opening. Mating and birth take place through a urogenital canal that exits at the tip of a hypertrophied clitoris. This 'masculine' phenotype spurred a search for an alternate source of fetal androgens. Although androstenedione from the maternal ovary is readily metabolized to testosterone by the hyena placenta, formation of the penile clitoris and scrotum appear to be largely androgen independent. However, secretions from the fetal testes underlie sex differences in the genitalia and central nervous system that are essential for male reproduction. Naturally circulating androgens, acting prenatally, reduce reproductive success in adult female spotted hyenas. Effects on aggression and dominance might offset these reproductive 'costs' of female androgenization in utero.