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Barbary macaque
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== Taxonomy and phylogeny == [[File:Barbarymaqaqueskull&brain.png|thumb|right|Skull and brain, as illustrated in [[Paul Gervais|Gervais]]' ''Histoire naturelle des mammifères'']] [[File:Macaca sylvanus 3d scan Natural History Museum University of Pisa C 1600.stl|thumb|3d model of skeleton]] The Barbary macaque is first described in scientific literature by [[Aristotle]] in the fourth century BCE work ''[[History of Animals]]''. He writes of an ape with "arms like a man, only covered with hair", "feet [which] are exceptional in kind ... like large hands", and "a tail as small as small can be, just a sort of indication of a tail". It is likely that [[Galen]] (129–c.216) dissected the Barbary macaque in the second century CE, presuming the internal structure to be the same as a human. Such was the authority of his work, some mistakes he made were not corrected until [[Andreas Vesalius]] (1514–1564) proved otherwise over a thousand years later.<ref name="McDermott1938"/> The Barbary macaque was included in the grouping ''Simia'' by [[Conrad Gessner]] in his 1551 work ''[[Historia animalium (Gessner book)|Historia Animalium]]'',<ref name="McDermott1938"/> a name which he claimed was already in use by the Greeks.<ref name="StilesOrleman1927"/> Gessner's ''[[Simia]]'' was subsequently used as one of [[Carl Linnaeus]]' four primate [[Genus|genera]] when he published ''[[Systema Naturae]]'' in 1758. Linnaeus proposed the [[scientific name]] ''Simia sylvanus'' for the Barbary macaque.<ref name=Linnaeus1758/> During the next 150 years primate taxonomy was subject to great changes and the Barbary macaque was placed in over thirty different [[taxa]].<ref name="Fooden2007"/> The confusion over the use of ''Simia'' became so great that the [[International Commission on Zoological Nomenclature]] (ICZN) [[nomen suppressum|suppressed]] its use in 1929.<ref name="StilesOrleman1927"/><ref name="Fooden2007"/> This meant the Barbary macaque was placed in the next oldest genus assigned to it, ''[[Macaca]]'', described by [[Bernard Germain de Lacépède]] in 1799.<ref name="Fooden2007"/> === Phylogeny === The Barbary macaque is the most [[Basal (phylogenetics)|basal]] macaque species.<ref name="Li_etal2009"/><ref name="HodgesCortes2006"/> [[Phylogenetic]] and [[Molecular biology|molecular]] analyses show it is a [[sister group]] to all Asian macaque species. The results of a phylogenetic analysis show that the chromosomes of Barbary macaque resemble those of the [[rhesus macaque]] with the exception of chromosomes 1, 4, 9, and 16. It was also discovered that chromosome 18 in the Barbary macaque is [[Sequence homology|homologous]] to [[chromosome 13]] in humans.<ref name="Fooden2007" /> [[Polymerase chain reaction]] studies have found [[Alu element|''Alu'' element'']] insertions, small pieces of genetic code in [[genome]]s, can infer primate phylogenetic relationships. Using this method the phylogenetic relationship of ten species within the genus ''Macaca'' has been resolved, showing the Barbary macaque to be a sister group to all other macaques.<ref name="Li_etal2009"/> {| class="wikitable" style="margin: 1em auto 1em auto" border="1" width="400pt" |- ! Phylogeny of ten species of ''Macaca''<ref name="Li_etal2009"/> |- | {{Clade|style=font-size:90%; line-height:110%; border: 0px solid darkgray; margin: 1em auto 1em auto; |label1=''[[Macaca]]'' |1={{Clade |1='''''Macaca sylvanus'' (Barbary macaque)''' |2={{Clade |1={{Clade |2={{Clade |1={{Clade |1=''[[Lion-tailed macaque|M. silenus]]'' (lion-tailed macaque) |2=''[[Southern pig-tailed macaque|M. nemestrina]]'' (southern pig-tailed macaque) }} }} |1={{Clade |1={{Clade |1=''[[Celebes crested macaque|M. nigra]]'' (Celebes crested macaque) }} }} }} |2={{Clade |1={{Clade |2={{Clade |1={{Clade |1=''[[Tibetan macaque|M. thibetana]]'' (Tibetan macaque) |2=''[[Stump-tailed macaque|M. arctoides]]'' (stump-tailed macaque) }} }} |1=''[[Bonnet macaque|M. radiata]]'' (bonnet macaque) }} |2={{Clade |2={{Clade |1={{Clade |1=''[[Japanese macaque|M. fuscata]]'' (Japanese macaque) |2=''[[Rhesus macaque|M. mulatta]]'' (rhesus macaque) }} }} |1=''[[Crab-eating macaque|M. fascicularis]]'' (crab-eating macaque) }} }} }} }} }} |} === Fossil record === Barbary macaque [[fossil]]s have been found across Europe, from the Atlantic Ocean to the Black Sea, dating from the Early [[Pliocene]] (early [[Zanclean]] {{mya|5.3|3.6}} <ref>{{Cite journal |last=Alba |first=D. M. |last2=Delson |first2=E. |last3=Morales |first3=J. |last4=Montoya |first4=P. |last5=Romero |first5=G. |date=2018 |title=Macaque remains from the early Pliocene of the Iberian Peninsula |journal=[[Journal of Human Evolution]] |volume=123 |pages=141–147 |doi=10.1016/j.jhevol.2018.07.005}}</ref>)to the [[Late Pleistocene]], assigned to various subspecies including ''M. s. sylvanus'', ''M. s. pliocena'' and ''M. s. florentina''.<ref name="EltonO'Regan2014" /> The [[Insular dwarfism|insular dwarf]] ''[[Macaca majori|M. majori]]'' endemic to Sardinia-Corsica during the [[Early Pleistocene]], usually considered to have derived from Barbary macaque, is generally considered a distinct species. Remains from [[Norfolk]], England, dating to the [[Middle Pleistocene]], at [[53rd parallel north|53 degrees latitude]], are amongst the northernmost records of non-human primates.<ref name="EltonO'Regan2014"/> [[Archaic humans]] and Barbary macaque remains were found co-occurring at numerous sites.<ref>{{Cite journal |last1=Castaños |first1=P. |last2=Murelaga |first2=X. |last3=Arrizabalaga |first3=A. |last4=Iriarte |first4=M.-J. |date=2011 |title=First evidence of ''Macaca sylvanus'' (Primates, Cercopithecidae) from the Late Pleistocene of Lezetxiki II cave (Basque Country, Spain) |journal=Journal of Human Evolution |volume=60 |issue=6 |pages=816–820 |doi=10.1016/j.jhevol.2011.01.011 |pmid=21481922 |bibcode=2011JHumE..60..816C}}</ref> It is thought possible that humans consumed Barbary macaques.<ref>{{Cite journal |last1=Konidaris |first1=G. E. |last2=Athanassiou |first2=A. |last3=Panagopoulou |first3=E. |last4=Harvati |first4=K. |date=2022 |title=First record of ''Macaca'' (Cercopithecidae, Primates) in the Middle Pleistocene of Greece |journal=Journal of Human Evolution |volume=162 |pages=103104 |doi=10.1016/j.jhevol.2021.103104 |pmc=8788689 |pmid=34883260 |bibcode=2022JHumE.16203104K}}</ref><ref>{{Cite journal |last1=Mecozzi |first1=B. |last2=Iannucci |first2=A. |last3=Sardella |first3=R. |last4=Curci |first4=A. |last5=Daujeard |first5=C. |last6=Moncel |first6=M.-H. |date=2021 |title=''Macaca ulna'' from new excavations at the Notarchirico Acheulean site (Middle Pleistocene, Venosa, southern Italy) |journal=Journal of Human Evolution |volume=153 |page=102946 |doi=10.1016/j.jhevol.2020.102946 |pmid=33639588 |bibcode=2021JHumE.15302946M |url=https://hal.archives-ouvertes.fr/hal-03153089/file/Mecozzi%20et%20al%20JHE%20Macaca%20Notarchirico.pdf}}</ref> The youngest known remains of a Barbary macaque in Europe were discovered at Hunas in Bavaria, Germany, dated to 85,000–40,000 years ago. The distribution of the Barbary macaque in Europe was likely strongly influenced by climate, only extending into Northern Europe during [[interglacial]] intervals.<ref name="EltonO'Regan2014"/><ref>{{Cite journal |last=Alba |first=D. M. |last2=Carlos Calero |first2=J. A. |last3=Mancheño |first3=M. Á. |last4=Montoya |first4=P. |last5=Morales |first5=J. |last6=Rook |first6=L. |date=2011 |title=Fossil remains of ''Macaca sylvanus florentina'' (Cocchi, 1872) (Primates, Cercopithecidae) from the Early Pleistocene of Quibas (Murcia, Spain) |journal=Journal of Human Evolution |volume=61 |issue=6 |pages=703–718 |doi=10.1016/j.jhevol.2011.09.003}}</ref> It was restricted to more southerly regions during colder glacial phases.<ref>{{Cite journal |last1=Fidalgo |first1=D. |last2=Rosas |first2=A. |last3=Estalrrich |first3=A. |last4=García-Tabernero |first4=A. |last5=Pastor |first5=F. J. |last6=Pineda |first6=A. |last7=Huguet |first7=R. |last8=Cáceres |first8=I. |last9=Ollé |first9=A. |last10=Vallverdú |first10=J. |last11=Saladie |first11=P. |date=2023 |title=First presence of ''Macaca sylvanus'' at the late Early Pleistocene of Barranc de la Boella (La Mina locality, Francolí Basin, NE Iberia) |journal=Journal of Human Evolution |volume=179 |pages=103368 |doi=10.1016/j.jhevol.2023.103368 |pmid=37094515 |bibcode=2023JHumE.17903368F |hdl=10261/308760 |hdl-access=free}}</ref> The disappearance of the Barbary macaque in Europe may have been caused by humans.<ref>{{Cite journal |last1=Meloro |first1=C. |last2=Elton |first2=S. |date=2013 |title=The Evolutionary History and Palaeo-Ecology of Primate Predation: ''Macaca sylvanus'' from Plio-Pleistocene Europe as a Case Study |journal=[[Folia Primatologica]] |volume=83 |issue=3–6 |pages=216–235 |doi=10.1159/000343494 |pmid=23363585 |url=https://dro.dur.ac.uk/37222/}}</ref> Fossils of the Barbary macaque are known from the Guefaït-4.2 site in Morocco, dating to around the Pliocene-Pleistocene boundary approximately 2.6 million years ago where measurements of [[Δ13C|carbon isotopes]] and [[Δ18O|oxygen isotopes]] indicate that the macaque consumed primarily the fresh fruits and leaves of [[C3 carbon fixation|C<sub>3</sub>]] plants.<ref>{{Cite journal |last1=Ramírez-Pedraza |first1=I. |last2=Martínez |first2=L. M. |last3=Aouraghe |first3=H. |last4=Rivals |first4=F. |last5=Tornero |first5=C. |last6=Haddoumi |first6=H. |last7=Estebaranz-Sánchez |first7=F. |last8=Rodríguez-Hidalgo |first8=A. |last9=van der Made |first9=J. |last10=Oujaa |first10=A. |last11=Ibáñez |first11=J. J. |last12=Mhamdi |first12=H. |last13=Souhir |first13=M. |last14=Aissa |first14=A. M. |last15=Chacón |first15=M. G. |last16=Sala-Ramos |first16=R. |date=2023 |title=Multiproxy approach to reconstruct fossil primate feeding behavior: Case study for macaque from the Plio-Pleistocene site Guefaït-4.2 (eastern Morocco) |journal=Frontiers in Ecology and Evolution |volume=11 |doi=10.3389/fevo.2023.1011208 |doi-access=free |hdl=10261/307158 |hdl-access=free}}</ref>
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