Dryopithecus is a genus of extinct great apes from the middle–late Miocene boundary of Europe 12.5 to 11.1 million years ago (mya). Since its discovery in 1856, the genus has been subject to taxonomic turmoil, with numerous new species being described from single remains based on minute differences amongst each other, and the fragmentary nature of the holotype specimen makes differentiating remains difficult. There is currently only one uncontested species, the type species D. fontani, though there may be more. The genus is placed into the tribe Dryopithecini, which is either an offshoot of orangutans, African apes, or is its own separate branch. A male specimen was estimated to have weighed in life. Dryopithecus likely predominantly ate ripe fruit from trees, suggesting a degree of suspensory behaviour to reach them, though the anatomy of a humerus and femur suggest a greater reliance on walking on all fours (quadrupedalism). The face was similar to gorillas, and males had longer canines than females, which is typically correlated with high levels of aggression. They lived in a seasonal, paratropical climate, and may have built up fat reserves for winter. European great apes likely went extinct during a drying and cooling trend in the Late Miocene which caused the retreat of warm-climate forests. Etymology The genus name Dryopithecus comes from Ancient Greek drus "oak tree" and pithekos "ape" because the authority believed it inhabited an oak or pine forest in an environment similar to modern day Europe. The species D. fontani was named in honour of its discoverer, local collector Monsieur Alfred Fontan. Taxonomy The first Dryopithecus fossils were described from the French Pyrenees by French paleontologist Édouard Lartet in 1856, three years before Charles Darwin published his On the Origin of Species. Subsequent authors noted similarities to modern African great apes. In his The Descent of Man, Darwin briefly noted that Dryopithecus casts doubt on the African origin of apes: Dryopithecus taxonomy has been the subject of much turmoil, with new specimens being the basis of a new species or genus based on minute differences, resulting in several now-defunct species. By the 1960s, all non-human apes were classified into the now-obsolete family Pongidae, and extinct apes into Dryopithecidae. In 1965, English palaeoanthropologist David Pilbeam and American palaeontologist Elwyn L. Simons separated the genus–which included specimens from across the Old World at the time–into three subgenera: Dryopithecus in Europe, Sivapithecus in Asia, and Proconsul