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===Communication=== {{Further|Great ape language}} {{listen | filename = Roep Indri Indri.ogg | title = Indri lemur wailing | description = Indri lemur wailing, [[ogg]]/[[Vorbis]] format. | format = [[Vorbis]] | filename2 = Howler monkey.ogg | title2 = Howler monkey roaring | description2 = Howler monkey roaring, [[ogg]]/[[Vorbis]] format. | format2 = [[Vorbis]] | filename3 = Vervet_Monkey_(Chlorocebus_pygerythrus)_(W_CERCOPITHECUS_AETHIOPS_R2_C2).ogg | title3 = Vervet monkey alarm call | description3 = Vervet monkey alarm call, [[ogg]]/[[Vorbis]] format. | format3 = [[Vorbis]] | filename4 = Symphalangus syndactylus 20170623 DouΓ©-la-Fontaine.ogg | title4 = Siamang singing | description4 = Siamang singing, [[ogg]]/[[Vorbis]] format. | format4 = [[Vorbis]] }} [[File:howler monkey.jpg|thumb|A pair of [[Alouatta caraya|black howler monkeys]] vocalizing]] [[Lemur]]s, [[loris]]es, [[tarsier]]s, and New World monkeys rely on [[Olfaction|olfactory signals]] for many aspects of social and reproductive behavior.<ref name="britannica" /> Specialized glands are used to [[mark territories]] with [[pheromone]]s, which are detected by the [[vomeronasal organ]]; this process forms a large part of the communication behavior of these primates.<ref name="britannica" /> In Old World monkeys and apes this ability is mostly [[Vestigiality|vestigial]], having regressed as [[Trichromacy|trichromatic]] eyes evolved to become the main sensory organ.<ref>{{cite journal |title=Relaxed selective pressure on an essential component of pheromone transduction in primate evolution |last1=Liman |first1=E. R. |last2=Innan |first2=H. |journal=Proceedings of the National Academy of Sciences of the United States of America |year=2003 |volume=100 |issue=6 |pages=3328β3332 |url=http://www.pnas.org/cgi/reprint/100/6/3328.pdf |access-date=2008-07-23 |doi=10.1073/pnas.0636123100 |pmid=12631698 |pmc=152292|bibcode = 2003PNAS..100.3328L|doi-access=free }}</ref> Primates also use vocalizations, gestures, and facial expressions to convey psychological state.<ref>{{cite book |last1=Egnor |first1=R. |last2=Miller |first2=C. |last3=Hauser |first3=M.D. |year=2004 |chapter=Nonhuman Primate Communication |title=Encyclopedia of Language and Linguistics |edition=2nd |publisher=Elsevier |isbn=0-08-044299-4 |chapter-url=http://www.wjh.harvard.edu/~mnkylab/publications/animalcommunication/PrimateComm_ElsevierEncy.pdf |url-status=dead |archive-url=https://web.archive.org/web/20080910043811/http://www.wjh.harvard.edu/~mnkylab/publications/animalcommunication/PrimateComm_ElsevierEncy.pdf |archive-date=2008-09-10}}</ref><ref name="Pollick"/> Facial musculature is very developed in primates, particularly in monkeys and apes, allowing for complex facial communication. Like humans, chimpanzees can distinguish the faces of familiar and unfamiliar individuals.<ref>{{cite journal|author=Burrows, A. M.|year=2008|title=The facial expression musculature in primates and its evolutionary significance|journal=BioEssays|volume=30|issue=3|pages=212β225|doi=10.1002/bies.20719|pmid=18293360|s2cid=205478149|url=https://www.researchgate.net/publication/5559596}}</ref> Hand and arm gestures are also important forms of communication for great apes and a single gesture can have multiple functions.<ref name="Pollick">{{cite journal|last1=Pollick |first1=A. S. |last2=de Waal |first2=F. B. M.|year=2007|title=Ape gestures and language evolution|journal=Proceedings of the National Academy of Sciences|volume=104|issue=19|pages=8184β8189|doi=10.1073/pnas.0702624104|pmid=17470779 |pmc=1876592|bibcode=2007PNAS..104.8184P|doi-access=free }}</ref> Chest-beating in male gorillas is a form of visual and non-vocal sound communication that serves to show fitness to both rivals and females.<ref>{{Cite journal |last1=Wright |first1=E. |last2=Grawunder|first2=S|last3=Ndayishimiye|first3=E|last4=Galbany|first4=J|last5=McFarlin|first5=S. C.|last6=Stoinski|first6=T. S.|last7=Robbins|first7=M. M. |year=2021 |title=Chest beats as an honest signal of body size in male mountain gorillas (''Gorilla beringei beringei'') |journal=Scientific Reports |volume=11 |issue=1 |page=6879 |doi=10.1038/s41598-021-86261-8 |pmid=33833252 |pmc=8032651 |bibcode=2021NatSR..11.6879W |doi-access=free}}</ref> Primates are a particularly vocal group of mammals.<ref name="Ankel-Simons2010"/> [[Indri]]s and [[black-and-white ruffed lemur]]s make distinctive, loud songs and choruses which maintain territories and act as [[alarm call]]s.<ref>{{cite journal|url=http://gibbons.de/main/non-gibbon/pdf_files/2006indri_varecia.pdf |author1=Geissmann, Thomas |author2=Mutschler, Thomas |name-list-style=amp |title=Diurnal Distribution of Loud Calls in Sympatric Wild Indris (''Indri indri'') and Ruffed Lemurs (''Varecia variegata''): Implications for Call Functions|pmid=16736264|year=2006|volume=47|issue=4|pages=393β6|doi=10.1007/s10329-006-0189-5|journal=Primates; Journal of Primatology|s2cid=1586657 }}</ref> The [[Philippine tarsier]], has a high-frequency limit of auditory sensitivity of approximately 91 kHz with a dominant frequency of 70 kHz, among the highest recorded for any terrestrial mammal. For Philippine tarsiers, these ultrasonic vocalizations might represent a private channel of communication that subverts detection by predators, prey and competitors, enhances energetic efficiency, or improves detection against low-frequency background noise.<ref name="Ramsier etal., (2012)">{{cite journal |last1=Ramsier |first1=M.A. |last2=Cunningham |first2=A.J. |last3=Moritz |first3=G.L. |last4=Finneran |first4=J.J. |last5=Williams |first5=C.V. |last6=Ong |first6=P.S. |last7=Gursky-Doyen |first7=S.L. |last8=Dominy |first8=N.J. | year = 2012 | title = Primate communication in the pure ultrasound| journal = Biology Letters | volume = 8 | issue = 4| pages = 508β511 | doi = 10.1098/rsbl.2011.1149 | pmid = 22319094 | pmc=3391437}}</ref> Male [[howler monkey]]s are among the loudest land mammals as their roars can be heard up to {{Convert|4.8|km|mi|abbr=on}}, and relate to intergroup spacing, territorial protection and possibly mate-guarding.<ref>{{cite journal |last1=da Cunha |first1=R. G. T. |last2=Byrne |first2=R.|year=2006|title=Roars of Black Howler Monkeys (Alouatta caraya): Evidence for a Function in Inter-Group Spacing|journal=Behaviour|volume=143|issue=10|pages=1169β1199|jstor=4536401|doi=10.1163/156853906778691568}}</ref><ref>{{cite web|title=Black howler monkey|date=25 April 2016|publisher=Smithsonian's National Zoo & Conservation Biology Institute|url=https://nationalzoo.si.edu/animals/black-howler-monkey|access-date=2016-07-10}}</ref> Male and female [[siamang]]s both possess inflatable pouches in the throat with which pair -bonds use to sing "duets" to each other.<ref>{{cite journal|last1=Haimoff|first1=E. H.|year=1983|title=Brief report: Occurrence of anti-resonance in the song of the siamang (''Hylobates syndactylus'')|journal=American Journal of Primatology|volume=5|issue=3|pages=249β256|doi=10.1002/ajp.1350050309|pmid=31986856 |s2cid=85262432 }}</ref> The [[vervet monkey]] gives a distinct alarm call for each of at least four different predators, and the reactions of other monkeys vary according to the call.<ref name=Seyfarth>{{cite journal| last1=Seyfarth| first1=R. M. |first2=D. L. |last2=Cheney |first3=Peter |last3=Marler |title=Vervet Monkey Alarm Calls: Semantic communication in a Free-Ranging Primate |journal=Animal Behaviour |volume=28 |issue=4 |pages=1070β1094 |year=1980 |doi=10.1016/S0003-3472(80)80097-2 | s2cid=53165940 |url=https://www.researchgate.net/publication/223576319}}</ref> Furthermore, many primate species including [[chimpanzee]]s,<ref>{{cite journal |last1=Leroux |first1=M. |last2=Schel |first2=A.M. |last3=Wilke |first3=C. |last4=Chandia |first4=B. |last5=Zuberbuhler |first5=K. |last6=Slocombe |first6=K.E. |last7=Townsend |first7=S. |title=Call combinations and compositional processing in wild chimpanzees |journal=Nature Communications |date=2023 |volume=14 |issue=1 |page=2225 |doi=10.1038/s41467-023-37816-y |pmid=37142584 |pmc=10160036 |bibcode=2023NatCo..14.2225L }}</ref> [[Campbell's mona monkey]]s<ref>{{cite journal |last1=Coye |first1=C. |last2=Ouattara |first2=K. |last3=Zuberbuhler |first3=K. |last4=Lemasson |first4=A. |title=Suffixation influences receivers' behaviour in non-human primates |journal=Proceedings of the Royal Society B |date=2015 |volume=282 |issue=1807 |page=20150265 |doi=10.1098/rspb.2015.0265 |doi-access=free|pmid=25925101 |pmc=4424650 }}</ref> or [[Diana monkey]]s<ref>{{cite journal |last1=Coye |first1=C. |last2=Zuberbuhler |first2=K. |last3=Lemasson |first3=A. |title=Morphologically structured vocalizations in female Diana monkeys |journal=Animal Behaviour |date=2016 |volume=115 |pages=97β105 |doi=10.1016/j.anbehav.2016.03.010|hdl=10023/10629 |hdl-access=free }}</ref> have been shown to combine vocalizations in sequences, suggesting [[syntax]] may not be uniquely humans as previously thought but rather evolutionary ancient, and its origins may be deeply rooted in the primate lineage.<ref>{{cite journal |last1=Leroux |first1=M. |last2=Townsend |first2=S. |title=Call Combinations in Great Apes and the Evolution of Syntax |journal=Animal Behavior and Cognition |date=2020 |volume=7 |issue=2 |pages=131β139 |doi=10.26451/abc.07.02.07.2020}}</ref> Consonant- and vowel-like sounds exist in some orangutan calls and they maintain their meaning over great distances.<ref>{{cite journal|last1=Lameira|first1=A. R.|display-authors=etal|year=2021|title=Orangutan information broadcast via consonant-like and vowel-like calls breaches mathematical models of linguistic evolution|journal=Biology Letters|volume=17|issue=9|doi=10.1098/rsbl.2021.0302|pmid=34582737 |pmc=8478518 }}</ref> The time range for the evolution of human language and/or its anatomical prerequisites extends, at least in principle, from the phylogenetic divergence of ''[[Homo]]'' (2.3 to 2.4 million years ago) from ''[[Pan (genus)|Pan]]'' (5 to 6 million years ago) to the emergence of full [[behavioral modernity]] some 50,000β150,000 years ago. Few dispute that ''[[Australopithecus]]'' probably lacked vocal communication significantly more sophisticated than that of [[great ape]]s in general.<ref name="Arcadi 2000">{{Cite journal | last1 = Arcadi | first1 = AC. | title = Vocal responsiveness in male wild chimpanzees: implications for the evolution of language | journal = J Hum Evol | volume = 39 | issue = 2 | pages = 205β23 |date=Aug 2000 | doi = 10.1006/jhev.2000.0415 | pmid = 10968929 | s2cid = 7403772 | doi-access = free | bibcode = 2000JHumE..39..205A }}</ref>
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