A series of works by Sharleen T. Sakai’s group have correlated the proportions of the anterior endocranial region with social behaviour in hyenas. They found that in spotted hyenas (Crocuta crocuta), males have relatively larger anterior cerebrum than females. The relative volume of the anterior endocranial region is also significantly larger in this species when compared to other extant species of hyenas. The spotted hyenas are the most gregarious species, living in large clans, where females are dominant and philopatric, and males disperse and must adapt to the hierarchic system of a new clan. The anterior region of the hyena’s brain comprises mostly the frontal cortex, which mediates social behaviour. The authors hypothesize, in the light of the social brain hypothesis, that the development of the frontal region in this species, and particularly in males, might have been enhanced by the need for a larger behavioural flexibility in their complex social environment. More recently, Joan Madurell-Malapeira and his colleagues compared the endocasts of two extinct spotted hyenas (C. spelaea and C. ultima) with those of extant species. The fossil specimens have similar morphology to that of C. crocuta, but less developed anterior portion of their endocranium. The authors therefore propose this feature to be an autapomorphy of C. crocuta. Consequently, the social and foraging behaviour of these fossil species are presumably less specialized, and this might contradict some speculations about competition between hyenas and humans during Pleistocene.