Editing Primate (section)

===Evolutionary history===
{{Further|Evolution of primates}}
The primate lineage is thought to go back at least near the [[Cretaceous–Paleogene boundary]] or around 74–63 ([[mya (unit)|mya]]).<ref name="2010Williams">{{cite journal | last1 = Williams | first1 = B.A. | last2 = Kay | first2 = R.F. | last3 = Kirk | first3 = E.C. | title = New perspectives on anthropoid origins | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 107 | pages = 4797–4804 | doi = 10.1073/pnas.0908320107 | issue = 11 | pmid = 20212104 | year = 2010 | pmc = 2841917 | bibcode=2010PNAS..107.4797W| doi-access = free }}</ref><ref name="StanyonSpringer2012">{{cite journal |last1=Stanyon |first1=Roscoe |last2=Springer |first2=Mark S.|last3=Meredith|first3=Robert W. |last4=Gatesy |first4=John|last5=Emerling|first5=Christopher A. |last6=Park |first6=Jong|last7=Rabosky|first7=Daniel L. |last8=Stadler |first8=Tanja |last9=Steiner |first9=Cynthia|last10=Ryder|first10=Oliver A.|last11=Janečka|first11=Jan E. |last12=Fisher |first12=Colleen A. |last13=Murphy |first13=William J. |title=Macroevolutionary Dynamics and Historical Biogeography of Primate Diversification Inferred from a Species Supermatrix |journal=PLOS ONE |volume=7 |issue=11|year=2012 |pages=e49521 |issn=1932-6203 |doi=10.1371/journal.pone.0049521 |pmid=23166696 |pmc=3500307|bibcode=2012PLoSO...749521S|doi-access=free }}</ref><ref name="JamesonHou2011">{{cite journal|last1=Jameson|first1=Natalie M. |last2=Hou |first2=Zhuo-Cheng |last3=Sterner |first3=Kirstin N. |last4=Weckle |first4=Amy |last5=Goodman |first5=Morris |last6=Steiper |first6=Michael E. |last7=Wildman |first7=Derek E. |title=Genomic data reject the hypothesis of a prosimian primate clade |journal=Journal of Human Evolution |volume=61 |issue=3|date=September 2011|pages=295–305 |issn=0047-2484 |doi=10.1016/j.jhevol.2011.04.004 |pmid=21620437|bibcode=2011JHumE..61..295J }}</ref><ref name="PozziHodgson2014">{{cite journal |last1=Pozzi |first1=Luca |last2=Hodgson|first2=Jason A. |last3=Burrell |first3=Andrew S. |last4=Sterner |first4=Kirstin N.|last5=Raaum |first5=Ryan L.|last6=Disotell|first6=Todd R. |title=Primate phylogenetic relationships and divergence dates inferred from complete mitochondrial genomes|journal=Molecular Phylogenetics and Evolution |volume=75 |date=June 2014|pages=165–183|issn=1055-7903|doi=10.1016/j.ympev.2014.02.023 |pmid=24583291 |pmc=4059600|bibcode=2014MolPE..75..165P }}</ref><ref name="StanyonFinstermeier2013">{{cite journal |last1=Stanyon |first1=Roscoe |last2=Finstermeier |first2=Knut|last3=Zinner |first3=Dietmar|last4=Brameier|first4=Markus |last5=Meyer |first5=Matthias |last6=Kreuz |first6=Eva |last7=Hofreiter|first7=Michael|last8=Roos|first8=Christian|title=A Mitogenomic Phylogeny of Living Primates |journal=PLOS ONE |volume=8|issue=7|date=16 July 2013 |pages=e69504 |issn=1932-6203 |doi=10.1371/journal.pone.0069504 |pmid=23874967 |pmc=3713065 |bibcode=2013PLoSO...869504F|doi-access=free }}</ref> The earliest possible primate/proto-primate may be ''[[Purgatorius]]'', which dates back to [[Danian|Early Paleocene]] of North America ~66mya.<ref name="placental_radiation" /><ref>{{Cite journal |last1=Wilson Mantilla |first1=Gregory P. |last2=Chester |first2=Stephen G. B. |last3=Clemens |first3=William A. |last4=Moore |first4=Jason R. |last5=Sprain |first5=Courtney J. |last6=Hovatter |first6=Brody T. |last7=Mitchell |first7=William S. |last8=Mans |first8=Wade W. |last9=Mundil |first9=Roland |last10=Renne |first10=Paul R. |title=Earliest Palaeocene purgatoriids and the initial radiation of stem primates |journal=Royal Society Open Science |year=2021 |volume=8 |issue=2 |pages=210050 |doi=10.1098/rsos.210050 |pmc=8074693 |pmid=33972886|bibcode=2021RSOS....810050W }}</ref> The oldest known primates from the fossil record date to the Late Paleocene of Africa, c.57 mya (''[[Altiatlasius]]'')<ref name="BAWilliams_2010">{{Cite journal | last1 = Williams | first1 = B. A. | last2 = Kay | first2 = R. F. | last3 = Kirk | first3 = E. C. | title = New perspectives on anthropoid origins | doi = 10.1073/pnas.0908320107 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 107 | issue = 11| pages = 4797–4804 | year = 2010 | pmid = 20212104 | pmc = 2841917|bibcode = 2010PNAS..107.4797W| doi-access = free }}</ref> or the Paleocene-Eocene transition in the northern continents, c. 55 mya (''[[Cantius]]'', ''[[Donrussellia]]'', ''[[Altanius]]'', ''[[Plesiadapis]]'' and ''[[Teilhardina]]'').<ref name = "Miller_2005">{{Cite journal | last1 = Miller | first1 = E. R. | last2 = Gunnell | first2 = G. F. | last3 = Martin | first3 = R. D. | title = Deep Time and the Search for Anthropoid Origins | doi = 10.1002/ajpa.20352 | journal = American Journal of Physical Anthropology | volume = 128 | pages = 60–95 | year = 2005 | pmid = 16369958 | url = http://www.paleontology.lsa.umich.edu/Accomplishments/deeptime.ajpa2005.pdf | archive-date = 2012-04-18 | access-date = 2015-09-04 | archive-url = https://web.archive.org/web/20120418022859/http://www.paleontology.lsa.umich.edu/Accomplishments/deeptime.ajpa2005.pdf | url-status = dead }}</ref><ref name=ChatterjeeEtal2009>{{Cite journal|date=27 October 2009 |first1=Helen J |last1=Chatterjee |first2=Simon Y.W. |last2=Ho |first3=Ian |last3=Barnes |first4=Colin |last4=Groves |author-link4=Colin Groves |title=Estimating the phylogeny and divergence times of primates using a supermatrix approach |journal=BMC Evolutionary Biology |volume= 9 |doi=10.1186/1471-2148-9-259|pmid=19860891|pages=259|pmc=2774700 |issue=1 |doi-access=free |bibcode=2009BMCEE...9..259C }}</ref><ref name=placental_radiation>{{cite journal|author=O'Leary, M. A.|date=8 February 2013|title= The placental mammal ancestor and the post–K-Pg radiation of placentals|journal=Science|volume=339|issue=6120|pages=662–667|doi=10.1126/science.1229237|pmid=23393258|display-authors=etal |bibcode=2013Sci...339..662O|hdl=11336/7302 |s2cid=206544776|hdl-access=free}}</ref> Other studies, including molecular clock studies, have estimated the origin of the primate branch to have been in the mid-Cretaceous period, around 85 mya.<ref>{{cite journal|title=Molecular Clock Calibrations and Metazoan Divergence Dates|url=https://archive.org/details/sim_journal-of-molecular-evolution_1999-09_49_3/page/385|author=Lee, M.|journal=Journal of Molecular Evolution|volume=49|issue=3|date=September 1999 |pages=385–391|doi=10.1007/PL00006562|pmid=10473780|bibcode=1999JMolE..49..385L|s2cid=1629316}}</ref><ref>{{cite web|title=Scientists Push Back Primate Origins From 65&nbsp;Million To 85&nbsp;Million Years Ago|url=https://www.sciencedaily.com/releases/2002/04/020418073440.htm|website=Science Daily|access-date=2008-10-24}}</ref><ref>{{cite journal|title=Using the fossil record to estimate the age of the last common ancestor of extant primates|url=https://archive.org/details/sim_nature-uk_2002-04-18_416_6882/page/726|author1=Tavaré, S. |author2=Marshall, C. R. |author3=Will, O. |author4=Soligo, C. |author5=Martin R.D.|journal=Nature|date=April 18, 2002|pages=726–729 |volume=416|issue=6882 |doi=10.1038/416726a |pmid=11961552|bibcode = 2002Natur.416..726T|s2cid=4368374 }}</ref>

By modern [[Cladistics|cladistic]] reckoning, the order Primates is [[monophyletic]]. The suborder [[Strepsirrhini]], the "[[rhinarium|wet-nosed]]" primates, is generally thought to have split off from the primitive primate line about 63&nbsp;mya,<ref>{{cite journal |title=Molecular Remodeling of Members of the Relaxin Family During Primate Evolution |journal=Molecular Biology and Evolution|volume=18 |pages=393–403 |author1=Klonisch, T. |author2=Froehlich, C. |author3=Tetens, F. |author4=Fischer, B. |author5=Hombach-Klonisch, S. |year=2001 |pmid=11230540|issue=3 |doi=10.1093/oxfordjournals.molbev.a003815|doi-access=free }}</ref> although earlier dates are also supported.<ref name="genome">{{cite journal |title=Development and Application of a Phylogenomic Toolkit: Resolving the Evolutionary History of Madagascar's Lemurs |journal=Genome Research|volume=18 |pages=489–499 |year=2008 |author=Horvath, J.|doi=10.1101/gr.7265208 |pmid=18245770 |issue=3 |pmc=2259113 |display-authors=1 |last2=Weisrock|first2=D. W.|last3=Embry|first3=S. L.|last4=Fiorentino|first4=I. |last5=Balhoff |first5=J. P.|last6=Kappeler|first6=P.|last7=Wray|first7=G. A.|last8=Willard|first8=H. F.|last9=Yoder |first9=A. D.}}</ref> The seven strepsirrhine families are the five related [[lemur]] families and the two remaining families that include the [[Lorisidae|lorisids]] and the [[galago]]s.<ref name="MSW3" /><ref name="Mittermeier2008">{{cite journal |title=Lemur Diversity in Madagascar |author-link1=Russell Mittermeier |author1=Mittermeier, R. |author2=Ganzhorn, J. |author3=Konstant, W. |author4=Glander, K. |author5=Tattersall, I. |author6=Groves, C. |author7=Rylands, A. |author8=Hapke, A. |author9=Ratsimbazafy, J. |author10=Mayor, M. |author11=Louis, E. |author12=Rumpler, Y. |author13=Schwitzer, C. |author14=Rasoloarison, R. |journal=International Journal of Primatology |doi=10.1007/s10764-008-9317-y |pages=1607–1656 |volume=29 |issue=6 |date=December 2008 |s2cid=17614597 |url=https://dukespace.lib.duke.edu/dspace/bitstream/10161/6237/1/08%20lemur%20diversity.pdf |access-date=2019-09-24 |archive-date=2021-02-15 |archive-url=https://web.archive.org/web/20210215163911/https://dukespace.lib.duke.edu/dspace/bitstream/handle/10161/6237/08 |url-status=dead }}</ref> Older classification schemes wrap [[Sportive lemur|Lepilemuridae]] into [[Lemuridae]] and [[Galago|Galagidae]] into [[Lorisidae]], yielding a four-one family distribution instead of five-two as presented here.<ref name="MSW3" /> During the [[Eocene]], most of the northern continents were dominated by two groups, the [[adapiform]]s and the [[omomyid]]s.<ref name="Sellers" /><ref name="evolution">{{cite book|title=Primates in Perspective|author=Hartwig, W.|chapter=Primate Evolution|editor=Campbell, C. |editor2=Fuentes, A. |editor3=MacKinnon, K. |editor4=Panger, M. |editor5=Bearder, S.|year=2007|publisher=Oxford University Press|isbn=978-0-19-517133-4|pages=13–17}}</ref> The former are considered members of Strepsirrhini, but did not have a [[toothcomb]] like modern lemurs; recent analysis has demonstrated that ''[[Darwinius|Darwinius masillae]]'' fits into this grouping.<ref name="Williams2010">{{Cite journal | last1 = Williams | first1 = B. A. | last2 = Kay | first2 = R. F. | last3 = Christopher Kirk | first3 = E. | last4 = Ross | first4 = C. F. | title = ''Darwinius masillae'' is a strepsirrhine—a reply to Franzen ''et al.'' (2009) | doi = 10.1016/j.jhevol.2010.01.003 | journal = Journal of Human Evolution | volume = 59 | issue = 5 | pages = 567–573; discussion 573–9 | year = 2010 | pmid = 20188396 | bibcode = 2010JHumE..59..567W | url = http://strainlab.uchicago.edu/publications/Williams%20et%20al%202010.pdf | archive-url = https://web.archive.org/web/20130517155919/http://strainlab.uchicago.edu/publications/Williams%20et%20al%202010.pdf | archive-date = 2013-05-17 | accessdate = 2015-09-04 | url-status = dead }}</ref> The latter was closely related to tarsiers, monkeys, and apes. How these two groups relate to extant primates is unclear. Omomyids perished about 30&nbsp;mya,<ref name="evolution" /> while adapiforms survived until about 10&nbsp;mya.<ref>{{cite book|title=Primate Evolution and Human Origins|author1=Ciochon, R. |author2=Fleagle, J. |name-list-style=amp |location=Menlo Park, California|publisher=Benjamin/Cummings|year=1987|isbn= 978-0-202-01175-2|pages=72}}</ref>

According to genetic studies, the lemurs of Madagascar diverged from the lorisoids approximately 75 mya.<ref name="genome" /> These studies, as well as chromosomal and molecular evidence, also show that lemurs are more closely related to each other than to other strepsirrhine primates.<ref name="genome" /><ref name="Garbutt">{{cite book | last = Garbutt | first = N. | title = Mammals of Madagascar, A Complete Guide | publisher = A&C Black Publishers | year = 2007 | isbn = 978-0-300-12550-4 | pages = 85–86}}</ref> However, Madagascar split from Africa 160 mya and from India 90 mya.<ref name="Lemurs">{{cite book | last1 = Mittermeier | first1 = R.A. | author-link = Russell Mittermeier | title = Lemurs of Madagascar | edition = 2nd | publisher = Conservation International | year = 2006 | isbn=1-881173-88-7 | pages = 23–26|display-authors=etal}}</ref> To account for these facts, a founding lemur population of a few individuals is thought to have reached Madagascar from Africa via a single [[rafting event]] between 50 and 80 mya.<ref name="genome" /><ref name="Garbutt" /><ref name="Lemurs" /> Other colonization options have been suggested, such as multiple colonizations from Africa and India,<ref name="Sellers">{{cite web|url=http://homepage.mac.com/wis/Personal/lectures/human-origins/PrimateEvolution.pdf |title=Primate Evolution |access-date=2008-10-23 |last=Sellers |first=Bill |date=2000-10-20 |publisher=University of Edinburgh |pages=13–17 |url-status=dead |archive-url=https://web.archive.org/web/20081029184647/http://homepage.mac.com/wis/Personal/lectures/human-origins/PrimateEvolution.pdf |archive-date=2008-10-29}}</ref> but none are supported by the genetic and molecular evidence.<ref name="genome" />
[[File:Brown Lemur in Andasibe.jpg|right|thumb|[[Common brown lemur]], a [[Strepsirrhini|strepsirrhine]] primate]]
Until recently, the [[aye-aye]] has been difficult to place within Strepsirrhini.<ref name="MSW3" /> Theories had been proposed that its family, Daubentoniidae, was either a lemuriform primate (meaning its ancestors split from the lemur line more recently than lemurs and lorises split) or a sister group to all the other strepsirrhines. In 2008, the aye-aye family was confirmed to be most closely related to the other Malagasy lemurs, likely having descended from the same ancestral population that colonized the island.<ref name="genome" />

Suborder [[Haplorhini]], the simple-nosed or "dry-nosed" primates, is composed of two sister clades.<ref name="MSW3" /> [[Prosimian]] tarsiers in the family Tarsiidae (monotypic in its own infraorder Tarsiiformes), represent the most [[Basal (phylogenetics)|basal]] division, originating about 58 mya.<ref>{{cite conference |title=Evolutionary Biology of Tarsiers |author=Shekelle, M. |year=2005 |url=http://rmbr.nus.edu.sg/bejc/ |access-date=2008-08-22 |url-status=dead |archive-url=https://web.archive.org/web/20080907073204/http://rmbr.nus.edu.sg/bejc/ |archive-date=2008-09-07 }}</ref><ref>{{cite journal |title=Rapid electrostatic evolution at the binding site for cytochrome c on cytochrome c oxidase in anthropoid primates |author=Schmidt, T. |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=102 |issue=18|pages=6379–6384 |doi=10.1073/pnas.0409714102 |date=3 May 2005 |pmid=15851671 |pmc=1088365 |display-authors=1 |last2=Wildman |first2=DE |last3=Uddin |first3=M |last4=Opazo |first4=JC |last5=Goodman |first5=M |last6=Grossman |first6=LI |bibcode = 2005PNAS..102.6379S|doi-access=free }}</ref> The earliest known haplorhine skeleton, that of 55 MA old tarsier-like ''[[Archicebus]]'', was found in central China,<ref name="SCN-20130605">{{cite web|last=Wade |first=Lizzie |title=Early Primate Weighed Less Than an Ounce |url=http://news.sciencemag.org/sciencenow/2013/06/crucial-link-in-primate-evolutio.html |publisher=[[ScienceNow]] |date=June 5, 2013 |access-date=2013-06-07 |url-status=dead |archive-url=https://web.archive.org/web/20130608230639/https://news.sciencemag.org/sciencenow/2013/06/crucial-link-in-primate-evolutio.html |archive-date=2013-06-08}}</ref> supporting an already suspected Asian origin for the group.<ref name="Kay2012">{{cite journal|last1=Kay |first1=R. F. |title=Evidence for an Asian origin of stem anthropoid s|journal=Proceedings of the National Academy of Sciences of the United States of America |volume= 109|issue= 26 |date= 2012 |pages= 10132–10133 |doi=10.1073/pnas.1207933109 |bibcode = 2012PNAS..10910132K |pmid=22699505 |pmc=3387095|doi-access=free }}</ref> The infraorder [[Simian|Simiiformes]] (simian primates, consisting of monkeys and apes) emerged about 40&nbsp;mya,<ref name="evolution" /> possibly also in Asia; if so, they [[Oceanic dispersal|dispersed]] across the [[Tethys Ocean|Tethys Sea]] from Asia to Africa soon afterwards.<ref name = "Chaimanee">{{Cite journal | last1 = Chaimanee | first1 = Y. | last2 = Chavasseau | first2 = O. | last3 = Beard | first3 = K. C. | last4 = Kyaw | first4 = A. A. | last5 = Soe | first5 = A. N. | last6 = Sein | first6 = C. | last7 = Lazzari | first7 = V. | last8 = Marivaux | first8 = L. | last9 = Marandat | first9 = B. | last10 = Swe | first10 = M. | last11 = Rugbumrung | first11 = M. | last12 = Lwin | first12 = T. | last13 = Valentin | first13 = X. | last14 = Zin-Maung-Maung-Thein | last15 = Jaeger | first15 = J. -J. | title = Late Middle Eocene primate from Myanmar and the initial anthropoid colonization of Africa | doi = 10.1073/pnas.1200644109 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 109 | issue = 26 | pages = 10293–10297 | year = 2012 | pmid = 22665790| pmc = 3387043| bibcode = 2012PNAS..10910293C| doi-access = free }}</ref> There are two simian clades, both [[wikt:parvorder|parvorder]]s: [[Catarrhini]], which developed in Africa, consisting of [[Old World monkey]]s, humans and the other apes, and Platyrrhini, which developed in South America, consisting of [[New World monkey]]s.<ref name="MSW3" /> A third clade, which included the [[eosimiids]], developed in Asia, but became extinct millions of years ago.<ref>{{cite journal |author=Marivaux, L. |title=Anthropoid primates from the Oligocene of Pakistan (Bugti Hills): Data on early anthropoid evolution and biogeography|journal=Proceedings of the National Academy of Sciences of the United States of America |date=2005-06-14 |volume=102 |issue=24 |pages=8436–8441 |doi=10.1073/pnas.0503469102 |pmid= 15937103|pmc=1150860|display-authors=1|last2= Antoine |first2=PO |last3= Baqri |first3=SR|last4= Benammi |first4=M|last5= Chaimanee |first5=Y |last6=Crochet |first6=JY |last7=De Franceschi|first7= D|last8=Iqbal |first8= N|last9=Jaeger|first9= JJ|bibcode = 2005PNAS..102.8436M|doi-access=free}}</ref>

As in the case of lemurs, the origin of New World monkeys is unclear. Molecular studies of concatenated nuclear sequences have yielded a widely varying estimated date of divergence between platyrrhines and catarrhines, ranging from 33 to 70 mya, while studies based on mitochondrial sequences produce a narrower range of 35 to 43 mya.<ref name=ChatterjeeEtal2009/><ref name="Schrago">{{cite journal |author1=Schrago, C.G. |author2=Russo, C.A.M. |name-list-style=amp | title = Timing the Origin of New World Monkeys | journal = Molecular Biology and Evolution | volume = 20 | issue = 10 | year = 2003 | pages = 1620–1625 | format = PDF Reprint | doi = 10.1093/molbev/msg172 | pmid = 12832653| doi-access = free }}</ref> The anthropoid primates possibly traversed the Atlantic Ocean from Africa to South America during the [[Eocene]] by [[island hopping]], facilitated by [[Mid-Atlantic Ridge|Atlantic Ocean ridges]] and a lowered sea level.<ref name="Sellers" /> Alternatively, a single [[Oceanic dispersal|rafting event]] may explain this transoceanic colonization. Due to [[continental drift]], the Atlantic Ocean was not nearly as wide at the time as it is today.<ref name="Sellers" /> Research suggests that a small {{convert|1|kg|lb|abbr=on}} primate could have survived 13 days on a raft of vegetation.<ref name="Houle">{{cite journal | author = Houle, A. | title = The origin of platyrrhines: An evaluation of the Antarctic scenario and the floating island model | url = https://archive.org/details/sim_american-journal-of-physical-anthropology_1999-08_109_4/page/541 | journal = [[American Journal of Physical Anthropology]] | volume = 109 | issue = 4 | year = 1999 | pages = 541–559 | doi = 10.1002/(SICI)1096-8644(199908)109:4<541::AID-AJPA9>3.0.CO;2-N | pmid = 10423268}}</ref> Given estimated current and wind speeds, this would have provided enough time to make the voyage between the continents.
[[File:Tamarin portrait 2 edit3.jpg|thumb|[[Emperor tamarin]], a [[New World monkey]]]]
Apes and monkeys spread from Africa into Europe and Asia starting in the [[Miocene]].<ref name="Andrews">{{cite journal |author1=Andrews, P. |author2=Kelley, J. |name-list-style=amp | title = Middle Miocene Dispersals of Apes |journal=Folia Primatologica | volume = 78 | year = 2007 | pages = 328–343 | doi = 10.1159/000105148 | pmid = 17855786 | issue = 5–6|s2cid=19293586 }}</ref> Soon after, the lorises and tarsiers made the same journey. The first hominin fossils were discovered in northern Africa and date back 5–8 mya.<ref name="evolution" /> Old World monkeys disappeared from Europe about 1.8 mya.<ref name="Strier2007">{{cite book|title=Primate Behavioral Ecology |author=Strier, K. |publisher = [[Allyn & Bacon]] | edition = 3rd|year=2007|isbn=978-0-205-44432-8|pages=7, 64, 71, 77, 182–185, 273–280, 284, 287–298}}</ref> Molecular and fossil studies generally show that modern humans originated in Africa 100,000–200,000 years ago.<ref>{{cite book |author1=Pough, F. W. |author2=Janis, C. M. |author3=Heiser, J. B. |title=Vertebrate Life |url=https://archive.org/details/vertebratelife0000poug |chapter=Primate Evolution and the Emergence of Humans |year=2005 |orig-year=1979 |edition=7th |publisher= Pearson |pages=[https://archive.org/details/vertebratelife0000poug/page/650 650] |isbn=0-13-127836-3}}</ref>

Although primates are well studied in comparison to other animal groups, several new species have [[Primates discovered in the 2000s|been discovered recently]], and genetic tests have revealed previously unrecognised species in known populations. ''Primate Taxonomy'' listed about 350 species of primates in 2001;<ref name="PT">{{cite book |title=Primate Taxonomy |publisher=Smithsonian Institution Press |year=2001 |author=Groves, C. P. |isbn=1-56098-872-X}}</ref> the author, [[Colin Groves]], increased that number to 376 for his contribution to the third edition of ''[[Mammal Species of the World]]'' (MSW3).<ref name=MSW3/> However, publications since the taxonomy in MSW3 was compiled in 2003 have pushed the number to 522 species, or 708 including subspecies.<ref name="IUCN_SSC">{{cite web |title=Primate diversity by region |author=IUCN/SSC Primate Specialist Group |date=1 March 2021 |publisher=International Union for the Conservation of Nature |url=http://www.primate-sg.org/primate_diversity_by_region/}}</ref>