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==Evolutionary history== ===Fossil record=== Fossils of [[tarsiiform]] primates have been found in Asia, Europe, and North America (with disputed fossils from Northern Africa), but extant tarsiers are restricted to several Southeast Asian islands. The fossil record indicates that their [[dentition]] has not changed much, except in size, over the past 45 million years. Within the family Tarsiidae, there are two extinct genera—''[[Xanthorhysis]]'' and ''[[Afrotarsius]]''; however, the placement of ''Afrotarsius'' is not certain,<ref name=Hartwig1>{{cite book | last1 = Gunnell | first1 = G. | last2 = Rose | first2 = K. | editor1-last = Hartwig | editor1-first = W.C. | title = The Primate Fossil Record | publisher = Cambridge University Press | year = 2002 | isbn = 978-0-521-66315-1 | chapter = Tarsiiformes: Evolutionary History and Adaptation}}</ref> and it is sometimes listed in its own family, [[Afrotarsiidae]], within the infraorder Tarsiiformes,<ref name=McKenna_Bell>McKenna, M.C., and Bell, S.K. 1997. ''Classification of Mammals Above the Species Level.'' Columbia University Press, New York, 337–340 pp. {{ISBN|0-231-11013-8}}</ref> or considered an [[Simian|anthropoid]] primate.<ref name=sirindhornae/> So far, four fossil species of tarsiers are known from the fossil record: * ''[[Tarsius eocaenus]]'' is known from the [[Middle Eocene]] in [[China]].<ref name=Hartwig1/><ref name=2006Rossie>{{cite journal | last1 = Rossie | first1 = J.B. | last2 = Ni | first2 = X. | last3 = Beard | first3 = K.C. | title = Cranial remains of an Eocene tarsier | year = 2006 | url = http://www.pnas.org/content/103/12/4381.full.pdf | journal = PNAS | volume = 103| issue = 12 | pages = 4381–4385 | pmid = 16537385|doi=10.1073/pnas.0509424103 | pmc=1450180| doi-access = free }}</ref> * ''[[Hesperotarsius|Hesperotarsius thailandicus]]'' lived during the [[Early Miocene]] in northwestern [[Thailand]].<ref name=Hartwig1/><ref name=Nowak>{{cite book | last = Nowak | first = R.M. | publisher = Johns Hopkins University Press | title = Walker's Mammals of the World | url = https://archive.org/details/walkersmammalsof0002nowa | url-access = registration | edition = 6th | year = 1999 | isbn = 978-0-8018-5789-8 | pages = 94–97}}</ref> * ''[[Hesperotarsius|Hesperotarsius sindhensis]]'' lived during the [[Miocene]] in [[Pakistan]].<ref>{{cite journal | last1 = Zijlstra | first1 = Jelle S. | last2 = Flynn | first2 = Lawrence J. | last3 = Wessels | first3 = Wilma | year = 2013 | title = The westernmost tarsier: A new genus and species from the Miocene of Pakistan | journal = Journal of Human Evolution | volume = 65 | issue = 5| pages = 544–550 | doi = 10.1016/j.jhevol.2013.06.015 | pmid=23928350| bibcode = 2013JHumE..65..544Z }}</ref> * ''[[Tarsius sirindhornae]]'' lived during the [[Middle Miocene]] in northern Thailand.<ref name=sirindhornae>{{cite journal | author = Chiamanee, Y., Lebrun, R., Yamee, C., and Jaeger, J.-J. | year = 2010 | title = A new Middle Miocene tarsier from Thailand and the reconstruction of its orbital morphology using a geometric–morphometric method | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 278| issue = 1714| pages = 1956–1963| doi = 10.1098/rspb.2010.2062 | pmid=21123264 | pmc=3107645}}</ref> The genus ''Tarsius'' has a longer fossil record than any other primate genus, but the assignment of the Eocene and Miocene fossils to the genus is dubious.<ref>{{cite book | last = Simons | first = E.L. | year = 2003 | title = Tarsiers: past, present, and future | editor1-last = Wright | editor1-first = P.C. | editor2-last = Simons | editor2-first = E.L. | editor3-last = Gursky | editor3-first = S. | chapter = The Fossil Record of Tarsier Evolution | publisher = Rutgers University Press | isbn = 978-0-8135-3236-3}}</ref> ===Classification=== {{see also|List of tarsiiformes}} The [[phylogenetic]] position of extant tarsiers within the order Primates has been debated for much of the 20th century, and tarsiers have alternately been classified with [[strepsirrhine]] primates in the suborder Prosimii, or as the sister group to the [[simian]]s (Anthropoidea) in the infraorder [[Haplorrhini|Haplorhini]]. Analysis of [[Retrotransposon#SINEs|SINE]] insertions, a type of macromutation to the DNA, is argued to offer very persuasive evidence for the [[monophyly]] of Haplorhini, where other lines of evidence, such as [[DNA sequence]] data, remain ambiguous. Thus, some systematists argue the debate is conclusively settled in favor of a monophyletic Haplorrhini. In common with simians, tarsiers have a mutation in the [[L-gulonolactone oxidase]] (GULO) gene, which prevents their bodies from synthesizing vitamin C so they must find it in the diet. Since the strepsirrhines do not have this mutation and have retained the ability to make vitamin C, the genetic trait that confers the need for it in the diet would tend to place tarsiers with haplorhines.<ref>{{cite journal | title = Vitamin C biosynthesis in prosimians: Evidence for the anthropoid affinity of ''Tarsius'' | author1 = Pollock, J. I. | author2 = Mullin, R. J. | name-list-style = amp | journal = American Journal of Physical Anthropology | year = 1986 | volume = 73 | issue = 1 | pages = 65–70 | doi = 10.1002/ajpa.1330730106 | url = http://www3.interscience.wiley.com/journal/110488482/abstract?CRETRY=1&SRETRY=0 | archive-url = https://archive.today/20120628232930/http://www3.interscience.wiley.com/journal/110488482/abstract?CRETRY=1&SRETRY=0 | url-status = dead | archive-date = 28 June 2012 | pmid = 3113259 | access-date = 16 March 2010 | url-access = subscription }}</ref> [[File:Bohol Tarsier.jpg|thumb|right|[[Philippine tarsier]] (''Carlito syrichta''), one of the smallest primates]] At a lower phylogenetic level, the tarsiers have, until recently, all been placed in the genus ''Tarsius'',<ref name=MSW3/> while it was debated whether the species should be placed in two (a [[Sulawesi]] and a [[Philippine]]-western group) or three separate genera (Sulawesi, Philippine and western groups).<ref name="Asian">{{cite journal |last=Brandon-Jones |first=D. |year=2004 |title=Asian primate classification |journal=International Journal of Primatology |volume=25 |issue=1 |pages=97–164 |doi=10.1023/B:IJOP.0000014647.18720.32 |s2cid=29045930 |display-authors=etal}}</ref> Species level [[Taxonomy (biology)|taxonomy]] is complex, with [[Morphology (biology)|morphology]] often being of limited use compared to vocalizations.{{Citation needed|date=March 2012}} Further confusion existed over the validity of certain names. Among others, the widely used ''T. dianae'' has been shown to be a [[junior synonym]] of ''T. dentatus'', and comparably, ''T. spectrum'' is now considered a junior synonym of ''T. tarsier''.<ref name=MSW3/> <!--Anything to save in here? Text is uncited (although a ref is given for part). The recent taxonomy changes address these issues Several "vocal morphs" may represent undescribed [[taxa]], taxonomically separable from ''T. tarsier'' (=''spectrum'') (such as [[Minahasa]] and a tarsier from the [[Togian Islands]]), and many others from Sulawesi and surrounding islands (Shekelle & Leksono 2004). This may also be the case for a number of poorly known, isolated Philippine populations (such as the [[Basilan]], [[Leyte (island)|Leyte]] and [[Dinagat Islands|Dinagat]] populations of the ''T. syrichta'' group). --> In 2010, [[Colin Groves]] and Myron Shekelle suggested splitting the genus ''Tarsius'' into three genera, the Philippine tarsiers (genus ''[[Carlito (genus)|Carlito]]''), the western tarsiers (genus ''[[Cephalopachus]]''), and the eastern tarsiers (genus ''Tarsius''). This was based on differences in [[dentition]], eye size, limb and hand length, tail tufts, tail sitting pads, the number of [[Mammary gland|mammae]], chromosome count, [[socioecology]], vocalizations, and distribution. The senior taxon of the species, ''T. tarsier'' was restricted to the population of a [[Selayar Islands|Selayar]] island, which then required the resurrection of the defunct taxon ''T. fuscus''.<ref name=2010_Groves_Shekelle>{{cite journal | last1 = Groves | first1 = C. | last2 = Shekelle | first2 = M. | title = The Genera and Species of Tarsiidae | journal = International Journal of Primatology | volume = 31 | issue = 6 | pages = 1071–1082 | year = 2010 | doi = 10.1007/s10764-010-9443-1| s2cid = 21220811 }}</ref> In 2014, scientists published the results of a genetic study from across the range of the [[Philippine tarsier]], revealing previously unrecognised genetic diversity. Three subspecies are recognised in the established taxonomy: ''Carlito syrichta syrichta'' from [[Leyte]] and [[Samar]], ''C. syrichta fraterculus'' from [[Bohol]], and ''C. syrichta carbonarius'' from [[Mindanao]]. Their analysis of mitochondrial and nuclear DNA sequences suggested that ssp. ''syrichta'' and ''fraterculus'' may represent a single lineage, whereas ssp. ''carbonarius'' may represent two lineages – one occupies the majority of Mindanao while the other is in northeastern Mindanao and the nearby [[Dinagat Island]], which the authors termed the 'Dinagat-Caraga tarsier'. More detailed studies that integrate morphological data will be needed to review the taxonomy of tarsiers in the Philippines.<ref>{{Cite journal |last1=Brown |first1=Rafe M. |last2=Weghorst |first2=Jennifer A. |last3=Olson |first3=Karen V. |last4=Duya |first4=Mariano R. M. |last5=Barley |first5=Anthony J. |last6=Duya |first6=Melizar V. |last7=Shekelle |first7=Myron |last8=Neri-Arboleda |first8=Irene |last9=Esselstyn |first9=Jacob A. |last10=Dominy |first10=Nathaniel J. |last11=Ong |first11=Perry S. |last12=Moritz |first12=Gillian L. |last13=Luczon |first13=Adrian |last14=Diesmos |first14=Mae Lowe L. |last15=Diesmos |first15=Arvin C. |date=19 August 2014 |title=Conservation Genetics of the Philippine Tarsier: Cryptic Genetic Variation Restructures Conservation Priorities for an Island Archipelago Primate |journal=PLOS ONE |language=en |volume=9 |issue=8 |pages=e104340 |doi=10.1371/journal.pone.0104340 |doi-access=free |issn=1932-6203 |pmc=4138104 |pmid=25136854 |bibcode=2014PLoSO...9j4340B }}</ref> * Infraorder Tarsiiformes **'''Family Tarsiidae''': tarsiers ***'''Genus ''[[Carlito (genus)|Carlito]]''''' **** [[Philippine tarsier]], ''Carlito syrichta'' ***** ''C. s. syrichta'' ***** ''C. s. fraterculus'' (to be combined into ''C. s. syrichta''?) ***** ''C. s. carbonarius'' *** '''Genus ''[[Cephalopachus]]''''' **** [[Horsfield's tarsier]], ''Cephalopachus bancanus'' ***** ''C. b. bancanus'' ***** ''C. b. natunensis'' ***** ''C. b. boreanus'' ***** ''C. b. saltator'' *** '''Genus ''[[Tarsius]]''''' ****[[Dian's tarsier]], ''T. dentatus'' ****[[Makassar tarsier]] ''T. fuscus'' ****[[Lariang tarsier]], ''T. lariang'' ****[[Niemitz's tarsier]], ''T. niemitzi'' ****[[Peleng tarsier]], ''T. pelengensis'' ****[[Sangihe tarsier]], ''T. sangirensis'' ****[[Gursky's spectral tarsier]], ''T. spectrumgurskyae''<ref name="Shekelle">{{cite journal | title=Two New Tarsier Species (Tarsiidae, Primates) and the Biogeography of Sulawesi, Indonesia | first1=Myron |last1=Shekelle |first2=Colin P |last2=Groves |author-link2=Colin Groves |first3=Ibnu |last3=Maryanto |first4=Russell A |last4=Mittermeier |author-link4=Russell A Mittermeier |date=May 2017 |journal=Primate Conservation |volume=31 |issue=1 |pages=1–9 |url=https://www.researchgate.net/publication/316714884}}</ref> ****[[Jatna's tarsier]], ''T. supriatnai''<ref name=Shekelle/> ****[[Spectral tarsier]], ''T. tarsier'' ****[[Siau Island tarsier]], ''T. tumpara'' ****[[Pygmy tarsier]], ''T. pumilus'' ****[[Wallace's tarsier]], ''T. wallacei''
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