Cape lion in 2009. Characteristics The type specimen of the Cape lion was described as very large with black-edged ears and a black mane extending beyond the shoulders and under the belly. Skulls of two lion specimen in the British Natural History Museum from the Orange River basin were described as a little shorter in the occipital regions than other lions in South Africa and with a tendency to develop the second lower premolar. American zoologist Edmund Heller described the Cape lion's skull as longer than those of equatorial lions, by at least on average, despite being comparatively narrow. He considered the Cape lion to have been 'distinctly' bigger than other lions in Africa. Lions approaching were shot south of the Vaal River. 19th century authors claimed that the Cape lion was bigger than the Asiatic lion. Results of a long-term study indicate that the colour of lion manes is influenced by climatic variables and varies between individuals. Manes are darker and longer in cool seasons. Distribution and habitat In the early 19th century, lions still occurred in the Karoo plains and in the Northern Cape. In 1844, lions were sighted south of the Riet River. The last lions south of the Orange River were sighted between 1850 and 1858. In the northern Orange Free State, lions may have survived into the 1860s. In 2003, six lions from Kgalagadi Transfrontier Park were relocated to Addo Elephant National Park in the Eastern Cape Province. In captivity In 2000, specimens asserted to be descendants of the Cape lion were found in captivity in Russia, and two of them were brought to South Africa. South African zoo director John Spence reportedly was long fascinated by stories of these grand lions scaling the walls of Jan van Riebeeck's Fort de Goede Hoop in the 17th century. He studied van Riebeeck's journals to discern the Cape lion's features, which included a long black mane, black in their ears, and reportedly a larger size. He believed that some Cape lions might have been taken to Europe and interbred with other lions. His 30-year search led to his discovery of black-maned lions with features of the Cape lion at the Novosibirsk Zoo in Siberia, in 2000. Besides having a black mane, the specimen that attracted Spence had a "wide face and sturdy legs". Novosibirsk Zoo's population, which had 40 cubs over a 30-year period, continues, and Spence,