The evolutionary history of the lion Panthera leo began in Pliocene east Africa, as open habitats expanded towards the end of the Cenozoic. During the middle–late Pleistocene, lions spread to most parts of Eurasia, North America, and may have eventually reached as far south as Peru. Lions probably evolved group-living behaviour before they expanded out of Africa, and this trait is likely to have prevailed in subsequent populations. The first lions were presumed to have been maneless, and maneless forms seem to have persisted in Europe, and possibly the New World, until around 10 000 years ago. The maned form may have appeared c. 320 000–190 000 years ago, and may have had a selective advantage that enabled it to expand to replace the range of earlier maneless forms throughout Africa and western Eurasia by historic times: ‘latest wave hypothesis’.