It is still unclear whether the genus evolved in
Africa or
Asia, although the oldest known fossils are from Africa and dated to about 3.8 mya. The earliest remains from Asia currently attributed to the genus is
Crocuta honanensis from the Early Pleistocene of China dating to around 2.5-2.2 million years ago, but its relationship to the living spotted hyena is ambiguous.
Crocuta first appears in Europe around 800,000 years ago at the beginning of the
Middle Pleistocene, replacing the giant hyena
Pachycrocuta. The Eurasian "
cave hyenas" (
Crocuta spelaea,
Crocuta ultima and others) have either been considered subspecies of the living spotted hyena, Genetic analysis of cave hyenas have found them to be strongly genetically divergent from living African spotted hyenas, albeit with some evidence of limited interbreeding between the two populations. Two
extinct species are known to have coexisted with each other in eastern Africa during the
Pliocene;
Crocuta eturono and
Crocuta dietrichi, each one probably occupying a different
niche in regards to their preference for
scavenging or
hunting. In
Ahl al Oughlam, a Pliocene site in
Morocco, lived the species
Crocuta dbaa. In
China there was a Pliocene species,
Crocuta honanensis, The holotype of
Crocuta sivalensis from the Indian subcontinent has been determined to be a specimen of
Pliocrocuta and unrelated to
Crocuta proper and thus this species is invalid, though
Crocuta proper is indeed recognised from the Early Pleistocene
Swalik Hills of the northern Indian subcontinent. A 2022 review of the genus recognised 7 extinct species in addition to living spotted hyena, with African species including
C.eturono in the late Pliocene,
C. venustula in the Pliocene and Early Pleistocene (with the authors regarding
C. dietrichi and
C. dbaa as synonyms of this species), and
C. ultra in the Early-Middle Pleistocene, with two species,
C. honanensis (Early Pleistocene of China and the Indian continent) and
C. ultima (Middle-Late Pleistocene) recognised in Asia, and
C. intermedia (Middle Pleistocene) and
C. spelaea (Middle-Late Pleistocene) recognised in Europe. ==References==