Panthera pardus spelaea

Panthera pardus spelaea, also known as the European Ice Age leopard or the cave leopard, is a fossil leopard subspecies which roamed Europe in the Late Pleistocene and possibly the Holocene.

Taxonomy
The subspecies was first described as Felis pardus spelaea by Emil Bächler in 1936.

Several fossil bones from the Early, Middle and Late Pleistocene were described and proposed as different leopard subspecies:


 * P. p. antiqua
 * P. p. begoueni
 * P. p. sickenbergi
 * P. p. vraonensis

These are now considered junior synonyms of P. p. spelaea.

Mitochondrial genomes from Late Pleistocene European leopard specimens found in Germany suggests that they represent a distinct group of extinct leopards that diveged from the common ancestor of Asian leopards (including those from the Caucasus) around 500,000 years ago.

Description
The European Ice Age leopard's skull was medium-long, and its characteristics are closest to the Panthera pardus tulliana subspecies. An apparent depiction of this leopard in the Chauvet Cave shows a coat pattern similar to that of modern leopards but with a unspotted belly, presumably white. Like other mammals, leopards from the cold glacial periods of the Late Pleistocene are usually larger than those from the warm interglacial phases. As in modern leopards, there was a strong sexual dimorphism, with males being larger than females.

Distribution
The timing of arrival of leopards in Europe is disputed. Some authors have posited that they arrived in Europe during the late Early Pleistocene around 1.2-1.1 million years ago. while others have suggested that they arrived during the early Middle Pleistocene, around 600,000 years ago. While initially rare, leopards become much more common and widely distributed from the late Middle Pleistocene onwards, following the extinction of the "European jaguar" ''Panthera gombaszoegensis. ''

During the Last Glacial Maximum, leopards persisted in relatively temperate glacial refugia in the Iberian, Italian and Balkan Peninsulas. Bone fragments of P. p. spelaea were excavated in Switzerland, Italy, Spain, Germany, Great Britain, Poland and Greece. Leopard fossils dating to ~43,000 BP were found in the Radochowska Cave in Poland. The most complete skeleton of P. p. spelaea is known from Vjetrenica Cave in southern Bosnia and Herzegovina, where four leopard fossils were found. These are dated to the end of the Late Pleistocene, about 29,000–37,000 years ago. A cave painting of a leopard in the Chauvet Cave in southern France is dated to about 25,000–37,500 years old. The last European Ice Age leopards vanished from most parts of Europe about 24,000 years ago, just before the Last Glacial Maximum. In Germany, the leopard survived at least into the early Weichselian glaciation.

The site of Equi in northwestern Italy represents the richest concentration of leopard remains from Pleistocene Europe, with some 200 leopards having been excavated. The youngest reliable records for leopards outside of eastern Europe are from the Iberian Peninsula, around 17-11,000 years ago, with records in the Iberian Peninsula possibly extending into the early Holocene, during the Mesolithic. Modern (Asian-type) leopards are still found on the fringes of Europe in the North Caucasus.

Palaeobiology
Fossils of European Ice Age leopards in Europe are sometimes found in caves, where they apparently sought shelter or hid their prey. They generally preferred smaller caves, most likely because larger caves were usually occupied by larger predators such as cave bears, cave lions (P. spelaea), or humans. In European Ice Age caves, leopard bones are far rarer than those of lions, and all currently known fossils belong to adults, suggesting that they rarely, if ever, raised their cubs in caves. Where leopard remains are found in larger caves, they are often found in the cave's deeper recesses, as in Baumann's and Zoolithen Cave in Germany. It is not precisely known which prey species these leopards hunted, although they may have been similar to modern snow leopards, which prey on ibex, deer and wild boar. It is likely that leopards scavenged or occasionally killed cave bears during hibernation in their dens. During the cold phases, European Ice Age leopards occurred mainly in mountain or alpine boreal forests or in mountains above the treeline, and were not usually found in the lowland mammoth steppes.