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'''''Oreopithecus''''' (from the Greek {{lang|grc|[[wikt:ὄρος|ὄρος]]}}, {{transltransliteration|grc|oros}} and {{lang|grc|[[wikt:πίθηκος|πίθηκος]]}}, {{transltransliteration|grc|pithekos}}, meaning "hill-ape") is an [[extinction|extinct]] genus of [[hominoid]] [[primate]] from the [[Miocene]] epoch whose [[fossil]]s have been found in today's [[Tuscany]] and [[Sardinia]] in [[Italy]].<ref name="NSW-20191223">{{cite news |last=Osbourne |first=Hannah |title=Strange swamp-dwelling prehistroic ape that counldn't walk on two legs or climb trees poses evolutionary puzzle |url=https://www.newsweek.com/strange-swamp-dwelling-prehistoric-ape-that-couldnt-walk-two-legs-climb-trees-poses-evolutionary-1478852 |date=December 23, 2019 |magazine=[[Newsweek]] |access-date=December 23, 2019 |df=dmy-all}}</ref> It existed nine to seven million years ago in the Tusco-Sardinian area when this region was an isolated island in a chain of islands stretching from central Europe to northern Africa in what was becoming the [[Mediterranean Sea]].{{efn|In what remained of the [[Tethys Sea]], or what was becoming the [[Mediterranean Sea]]; see [[Mediterranean Basin#Geology and paleoclimatology|Geology and paleoclimatology of the Mediterranean Basin]]; see also [[Messinian salinity crisis]].}}
 
''Oreopithecus'' was one of many European immigrants that settled this area in the [[Vallesian]]–[[Turolian]] transition and one of few hominoids, together with ''[[Sivapithecus]]'' in Asia, to survive the so-called [[Vallesian|Vallesian Crisis]].<ref name="Agustí">{{harvnb|Agustí|Antón|2002| pp=Prefix ''ix'', 174–175, 193, 197–199}}</ref> To date, dozens of individuals have been discovered at the [[Tuscany|Tuscan]] localities of <!-- ref for localities -->[[Montebamboli]], [[Montemassi]], Casteani, [[Ribolla]], and, most notably, in the fossil-rich lignite mine in the [[Baccinello|Baccinello Basin]],<ref name="Agustí" /> making it one of the best-represented fossil [[ape]]s.
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Known as the "enigmatic [[hominoid]]", ''Oreopithecus'' can dramatically rewrite the palaeontological map depending on whether it is a descendant of the European ape ''[[Dryopithecus]]'' or an African [[Anthropoidea|anthropoid]].<ref name="Agustí"/> Some have suggested the unique locomotory behavior of ''Oreopithecus'' requires a revision of the current consensus on the timing of bipedality in human developmental history, but there is limited agreement on this point among [[paleontologist]]s.
 
Simons (1960) considered ''Oreopithecus'' closely related to the early Oligocene ''[[Apidium]]'', a small arboreal anthropoid that lived nearly 34&nbsp;million years ago in Egypt.<ref name="Simons-1960">{{harvnb|Simons|1960}}</ref> ''Oreopithecus'' shows strong links to modern apes in its [[postcranium]] and, in this respect, it is the most modern Miocene ape below the neck, with closest similarities to the postcranial elements of ''[[Dryopithecus]]'', but its dentition is adapted to a leafy diet and a close link is uncertain. Others claim it to be either the sister taxon to [[Cercopithecoidea]] or an even direct human ancestor, but it is usually placed in its own subfamily within [[Hominidae]]. It could instead be added to the same subfamily as ''Dryopithecus'', perhaps as a distinct tribe (Oreopithecini).<ref name="Delson-2000">{{harvnb|Delson|Tattersall|Van Couvering|2000| p=465}}</ref> A cladistic analysis of ''Nyanzapithecus alesi'' recovers ''Oreopithecus'' as a member of the proconsulid subfamily [[Nyanzapithecinae]].<ref>{{cite journal |first1=Isaiah |last1=Nengo |first2=Paul |last2=Tafforeau |first3=Christopher C. |last3=Gilbert |first4=John G. |last4=Fleagle |first5=Ellen R. |last5=Miller |first6=Craig |last6=Feibel |first7=David L. |last7=Fox |first8=Josh |last8=Feinberg |first9=Kelsey D. |last9=Pugh |first10=Camille |last10=Berruyer |first11=Sara |last11=Mana |first12=Zachary |last12=Engle |first13=Fred |last13=Spoor |display-authors=6 |year=2017 |title=New infant cranium from the African Miocene sheds light on ape evolution |journal=Nature |volume=548 |issue=7666 |pages=169–174 |doi=10.1038/nature23456|pmid=28796200 |bibcode=2017Natur.548..169N |s2cid=4397839 |url=https://discovery.ucl.ac.uk/id/eprint/1570349/1/Spoor_Nengo-et-al_text.pdf }}</ref> A 2023 phylogenetic analysis suggested found a close relationship with [[Gibbon|gibbonsgibbon]]s, though the author suggested that this was likely due to having a similar climbing lifestyle and retained plesiomorphies, rather than a real close relationsip, but suggested that it was unlikely that ''Oreopithecus'' was a member of Hominidae.<ref>{{Cite journal |last=Pugh |first=Kelsey D. |date=April 2022-04 |title=Phylogenetic analysis of Middle-Late Miocene apes |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248421001925 |journal=Journal of Human Evolution |language=en |volume=165 |pages=103140 |doi=10.1016/j.jhevol.2021.103140}}</ref>
 
==Physical characteristics==
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Its foot has been described as chimp-like, but is different from those of extant primates. The habitual line of leverage of the primate foot is parallel to the third [[metatarsal bone]]. In ''Oreopithecus'', the [[Anatomical terms of ___location#Left and right (lateral), and medial|lateral]] metatarsals are permanently [[Abduction (kinesiology)|abducted]] so that this line falls between the first and second metatarsals instead. Furthermore, the shape of the [[tarsus (skeleton)|tarsus]] indicate loads on the foot were transmitted to the medial side of the foot instead of the lateral, like in other primates.<ref name="Köhler-1997">{{harvnb|Köhler|Moyà-Solà|1997}}</ref> The metatarsals are short and straight, but have a lateral orientation increase. Its foot proportions are close to the unusual proportions of ''Gorilla'' and ''Homo'' but are distinct from those found in specialized climbers. The lack of predators and the limitation of space and resources in ''Oreopithecus''{{'}} insular environment favored a locomotor system optimized for low energy expenditure rather than speed and mobility.<ref name="Köhler-1997"/>
 
''Oreopithecus'' has been claimed to exhibit features that are adaptations to upright walking, such as the presence of a [[lumbar]] curve, in distinction to otherwise similar species known from the same period. Since the fossils have been dated to about 8 [[million years ago]], this would represent an unusually early appearance of upright posture.<ref name="Köhler-1997"/> However, a reevaluation of the spine from a skeleton of ''Oreopithecus'' has led to the conclusion that it lacked adaptations for habitual [[biped]]ality.<ref name="Ghose_2013">{{cite web |last=Ghose |first=Tia |title=Strange ancient ape walked on all fours |website=LiveScience.Com |publisher=TechMedia Network |date=2013-08-05 |url=http://www.livescience.com/38643-ancient-ape-walked-on-all-fours.html |access-date=2013-08-07 |df=dmy-all}}</ref><ref name="RussoShapiro2013">{{cite journal |last1=Russo |first1=G.A. |last2=Shapiro |first2=L.J. |title=Reevaluation of the lumbosacral region of ''Oreopithecus bambolii'' |journal=Journal of Human Evolution |date=2013-07-23 |df=dmy-all |doi=10.1016/j.jhevol.2013.05.004 |volume=65 |issue=3 |pages=253–265 |pmid=23891006}}</ref>
 
====Semicircular canals====