In Europe, the leopard occurred at least since the Pleistocene. Fossil bones and teeth dating from the Pliocene were excavated in Perrier in France, northeast of London, and in Valdarno, Italy. Similar fossils dating back to the Pleistocene were excavated mostly in loess and caves at 40 sites in the continent – from near Lisbon, near Gibraltar, and Santander Province in northern Spain to several sites in France, Switzerland, Italy, Austria, Germany, in the north up to Derby in England, in the east to Přerov in the Czech Republic, and the Baranya in southern Hungary, and in Biśnik Cave in south-central Poland. Four subspecies of European Pleistocene leopards were proposed. P. p. begoueni known from the beginning of the early Pleistocene was replaced about 0. 6 million years ago by P. p. sickenbergi, which in turn was replaced by P. p. antiqua around 0. 3 million years ago. The most recent P. p. spelaea appeared at the beginning of the Late Pleistocene and survived until about 24,000 years ago in several parts of Europe.
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