Editing Primate (section)

==Behavior==

===Social systems===
[[Richard Wrangham]] stated that [[Social structure|social systems]] of primates are best classified by the amount of movement by females occurring between groups.<ref name="social">{{cite book |author=Wrangham, R. W. |author-link=Richard Wrangham |chapter=Mutualism, kinship and social evolution |year=1982 |title=Current Problems in Sociobiology |publisher=Cambridge University Press |pages=269–89 |isbn=0-521-24203-7}}</ref> He proposed four categories:

* Female transfer systems&nbsp;– females move away from the group in which they were born. Females of a group will not be closely related whereas males will have remained with their natal groups, and this close association may be influential in social behavior. The groups formed are generally quite small.<ref name="social"/> This organization can be seen in [[chimpanzee]]s, where the males, who are typically related, will cooperate in defense of the group's territory.<ref>{{cite journal |author=Goldberg, T. L. |author2=Wrangham, R. W. | title=Genetic correlates of social behavior in wild chimpanzees: evidence from mitochondrial DNA | journal=Animal Behaviour | volume=54 | issue=3 | pages=559–70 | date=September 1997 | pmid=9299041 | doi=10.1006/anbe.1996.0450|s2cid=18223362 }}</ref> Evidence of this social system (called [[patrilocal residence]] when used by the [[Anthropology]]) has also been found among [[Neanderthal]] remains in [[Spain]]<ref>{{cite news | last = Bowdler | first = Neil | title = Neanderthal family found cannibalised in cave in Spain | url = https://www.bbc.co.uk/news/science-environment-12049854 | work = [[BBC News]] | date = 21 December 2010}}</ref> and in remains of ''[[Australopithecus]]'' and ''[[Paranthropus robustus]]'' groups in southern Africa.<ref name="CavewomenBBC2011">{{cite news | first=Neil | last=Bowdler | url=https://www.bbc.co.uk/news/science-environment-13609260 | title=Ancient cave women 'left childhood homes' | date=2 June 2011 | work=[[BBC News]] | access-date=2 June 2011}}</ref><ref name="Copeland2011">{{cite journal | last1 = Copeland | first1 = Sandi R.  |display-authors=etal|title=Strontium isotope evidence for landscape use by early hominins  |volume=474 | issue = 7349 | date=1 June 2011 |pages=76–78 |journal=Nature |doi=10.1038/nature10149 |pmid=21637256| s2cid = 205225222 }}</ref> Among New World Monkeys, [[spider monkey]]s and [[muriqui]]s use this system.<ref>{{cite book|title=Primates in Perspective|author1=Fiore, A. D. |author2=Campbell, C. J. |name-list-style=amp |chapter=The Atelines|year=2007|page=175|publisher=Oxford University Press|editor=Campbell, C. J. |editor2=Fuentes, A. |editor3=MacKinnon, K. C. |editor4=Panger, M. |editor5=Bearder, S. K.|isbn=978-0-19-517133-4}}</ref>
[[File:Ringstaartmakis - Ring-tailed Lemur.jpg|thumb|right|A social huddle of [[ring-tailed lemur]]s. The two individuals on the right exposing their white ventral surface are sunning themselves.]]
* Male transfer systems&nbsp;– while the females remain in their natal groups, the males will emigrate as adolescents. Group sizes are usually larger.<ref name="social"/> This system is common among the [[ring-tailed lemur]], [[capuchin monkey]]s and [[Cercopithecinae|cercopithecine monkeys]].<ref name="Strier2007" />
* Monogamous species&nbsp;– a male–female bond, sometimes accompanied by a juvenile offspring. There is shared responsibility of parental care and territorial defense. The offspring leaves the parents' territory during adolescence.<ref name="social"/> [[Indri]], [[lariang tarsier]]s, [[Callitrichidae]] monkeys and gibbons use this system, although "monogamy" in this context does not necessarily mean absolute sexual fidelity.<ref>{{cite book|title=Primates in Perspective|author=Bartlett, T. Q.|chapter=The Hylobatidae|year=2007|editor=Campbell, C. J. |editor2=Fuentes, A. |editor3=MacKinnon, K. C. |editor4=Panger, M. |editor5=Bearder, S. K.|publisher=Oxford University Press|isbn=978-0-19-517133-4|page=283}}</ref><ref name=Dixon/> These species do not live in larger groups.
* Solitary species&nbsp;– males and females live in overlapping home ranges.<ref name="social"/><ref name=Dixon/> This type of organization is found in lorises, galagos, [[mouse lemur]]s, aye-ayes and orangutans.<ref name=Dixon/>

Other systems are known to occur as well. For example, with [[howler monkey]]s and [[gorilla]]s both the males and females typically transfer from their natal group on reaching sexual maturity, resulting in groups in which neither the males nor females are typically related.<ref name="Sussman2003" /><ref name="Watts 1996">{{Cite book|author=Watts D. P.|year=1996|contribution=Comparative socio-ecology of gorillas|editor1=McGrew W. C. |editor2=Marchant L. F. |editor3=Nishida, T.|title=Great Ape Societies|url=https://archive.org/details/greatapesocietie00mcgr|location=Cambridge (England|publisher=Cambridge Univ Press|pages=[https://archive.org/details/greatapesocietie00mcgr/page/16 16]–28|isbn=978-0521555364}}</ref> Some prosimians, [[Colobinae|colobine]] monkeys and [[Callitrichinae|callitrichid]] monkeys also use this system.<ref name="Strier2007" />

The transfer of females or males from their native group is likely an adaptation for avoiding inbreeding.<ref name=Charpentier>{{cite journal |vauthors=Charpentier MJ, Widdig A, Alberts SC |title=Inbreeding depression in non-human primates: a historical review of methods used and empirical data |journal=American Journal of Primatology |volume=69 |issue=12 |pages=1370–86 |date=December 2007 |pmid=17486606 |doi=10.1002/ajp.20445 |s2cid=46626761 |url=https://hal.archives-ouvertes.fr/hal-02079454 }}</ref> An analysis of breeding records of captive primate colonies representing numerous different species indicates that the infant mortality of inbred young is generally higher than that of non-inbred young.<ref name=Charpentier /><ref>{{cite journal | vauthors = Ralls K, Ballou J | year = 1982 | title = Effect of inbreeding on infant mortality in captive primates | doi = 10.1007/BF02693747 | journal = International Journal of Primatology | volume = 3 | issue = 4 | pages = 491–505 | s2cid = 10954608 | url = http://si-pddr.si.edu/dspace/bitstream/10088/6162/1/8133A167-A994-450A-ADF4-DAD3F4EAA6F6.pdf }}{{Dead link|date=October 2023 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> This effect of inbreeding on infant mortality is probably largely a result of increased expression of deleterious recessive alleles (see [[Inbreeding depression]]).
[[File:Three chimpanzees with apple.jpg|thumb|right|[[Chimpanzee]]s are social great apes.]]
Primatologist [[Jane Goodall]], who studied in the [[Gombe Stream National Park]], noted [[fission-fusion society|fission-fusion societies]] in chimpanzees.<ref>{{cite journal |last1=Constable |first1=J. L. |last2=Ashley |first2=M. V. |last3=Goodall |first3=J. |last4=Pusey |first4=A. E. |title=Noninvasive paternity assignment in Gombe chimpanzees |url=https://archive.org/details/sim_molecular-ecology_2001-05_10_5/page/1279 |journal=Molecular Ecology |volume=10 |issue=5 |pages=1279–300 | date=May 2001 |pmid=11380884 |doi=10.1046/j.1365-294X.2001.01262.x|bibcode=2001MolEc..10.1279C |s2cid=46604532 }}</ref> There is ''fission'' when the main group splits up to forage during the day, then ''fusion'' when the group returns at night to sleep as a group. This social structure can also be observed in the [[hamadryas baboon]],<ref name="Rowe1996">{{cite book|title=The Pictorial Guide to the Living Primates|url=https://archive.org/details/pictorialguideto0000rowe|url-access=registration|author=Rowe, N.|year=1996|publisher=Pogonias Press|isbn=0-9648825-0-7|pages=[https://archive.org/details/pictorialguideto0000rowe/page/n15 4], 139, 143, 15 185, 223}}</ref> [[spider monkey]]s<ref name="Sussman2003" /> and the [[bonobo]].<ref name="Rowe1996" /> The [[gelada]] has a similar social structure in which many smaller groups come together to form temporary herds of up to 600 monkeys.<ref name="Rowe1996" /> [[Human]]s also form fission-fusion societies. In hunter-gatherer societies, humans form groups which are made up of several individuals that may split up to obtain different resources.<ref>{{Cite journal|last1=Couzin|first1=Iain D.|last2=Laidre|first2=Mark E.|date=August 2009|title=Fission–fusion populations|journal=Current Biology|volume=19|issue=15|pages=R633–R635|doi=10.1016/j.cub.2009.05.034|pmid=19674541|s2cid=13549970|issn=0960-9822|doi-access=free|bibcode=2009CBio...19.R633C }}</ref>

These social systems are affected by three main ecological factors: distribution of resources, [[Group size measures|group size]], and [[predation]].<ref name="vertlife">{{cite book |last1=Pough |first1=F. W. |last2=Janis |first2=C. M. |last3=Heiser |first3=J. B. |title=Vertebrate Life |url=https://archive.org/details/vertebratelife0000poug |chapter=Primate Societies |year=2005 |orig-year=1979 |edition=7th |publisher= Pearson |pages=[https://archive.org/details/vertebratelife0000poug/page/621 621]–623 |isbn=0-13-127836-3}}</ref> Within a social group there is a balance between cooperation and competition. Cooperative behaviors in many primates species include [[social grooming]] (removing [[Parasitism|skin parasites]] and cleaning wounds), food sharing, and collective defense against predators or of a territory. Aggressive behaviors often signal competition for food, sleeping sites or mates. Aggression is also used in establishing [[dominance hierarchy|dominance hierarchies]].<ref name="vertlife" /><ref>[[Barbara Smuts|Smuts, B.B.]], Cheney, D.L. Seyfarth, R.M., Wrangham, R.W., & Struhsaker, T.T. (Eds.) (1987). ''Primate Societies''. Chicago: University of Chicago Press for articles on the structure and function of various primate societies.</ref>

In November 2023, scientists reported, for the first time, evidence that groups of primates, particularly [[bonobo]]s, are capable of cooperating with each other.<ref name="NYT-20231116">{{cite news |last=Zimmer |first=Carl |authorlink=Carl Zimmer |title=Scientists Find First Evidence That Groups of Apes Cooperate - Some bonobos are challenging the notion that humans are the only primates capable of group-to-group alliances. |url=https://www.nytimes.com/2023/11/16/science/bonobos-cooperation-study.html |date=16 November 2023 |work=[[The New York Times]] |url-status=live |archiveurl=https://archive.today/20231116194259/https://www.nytimes.com/2023/11/16/science/bonobos-cooperation-study.html |archivedate=16 November 2023 |accessdate=17 November 2023 }}</ref><ref name="SCI-20231116">{{cite journal |author=Samuni, Liran |display-authors=et al. |title=Cooperation across social borders in bonobos
|url=https://www.science.org/doi/10.1126/science.adg0844 |date=16 November 2023 |journal=[[Science (journal)|Science]] |volume=382 |issue=6672 |pages=805–809 |doi=10.1126/science.adg0844 |pmid=37972165 |bibcode=2023Sci...382..805S |url-status=live |archiveurl=https://archive.today/20231117125744/https://www.science.org/doi/10.1126/science.adg0844 |archivedate=17 November 2023 |accessdate=17 November 2023 }}</ref>

===Interspecific associations===
Several species of primates are known to associate in the wild. Some of these associations have been extensively studied. In the [[Taï National Park|Tai Forest]] of Africa, several species coordinate anti-predator behavior. These include the [[Diana monkey]], [[Campbell's mona monkey]], [[lesser spot-nosed monkey]], [[western red colobus]], [[king colobus]] (western [[Black-and-white colobus|black and white colobus]]), and [[sooty mangabey]], which coordinate anti-predator alarm calls.<ref name="tai">{{cite book|title=Monkeys of Tai Forest, An African Primate Community|chapter=Interactions between African Crowned Eagles and Their Prey Community|author1=Shultz, S. |author2=Thomsett, S. |name-list-style=amp |editor=McGraw, W. |editor2=Zuberbuhler, K. |editor3=Noe, R.|year=2007|publisher=Cambridge University Press|isbn=978-0-521-81633-5|pages=181}}</ref> Among the predators of these monkeys is the [[common chimpanzee]].<ref name="tai2007">{{cite book|title=Monkeys of Tai Forest, An African Primate Community|chapter=Interactions between Red Colobus Monkeys and Chimpanzees|author=Bshary, R.|editor=McGraw, W. |editor2=Zuberbuhler, K. |editor3=Noe, R.|year=2007|publisher=Cambridge University Press|isbn=978-0-521-81633-5|pages=155–170}}</ref>

The [[red-tailed monkey]] associates with several species, including the western red colobus, [[blue monkey]], [[Wolf's mona monkey]], [[mantled guereza]], [[black crested mangabey]] and [[Allen's swamp monkey]].<ref name="Rowe1996" /> Several of these species are preyed upon by the common chimpanzee.<ref name="chimpcolobus">{{cite book|title=Chimpanzee and Red Colobus : the ecology of predator and prey|url=https://archive.org/details/chimpanzeeredcol0000stan|author=Stanford, C.|year=1998|publisher=Harvard University Press|isbn=0-674-00722-0|pages=[https://archive.org/details/chimpanzeeredcol0000stan/page/130 130]–138, 233}}</ref>

In South America, [[squirrel monkey]]s associate with [[capuchin monkey]]s.<ref name="move">{{cite book|title=On the Move : how and why animals travel in groups|chapter=Social Manipulation Within and Between Troops Mediates Primate Group Movement|author=Boinski, S.|editor=Boinski, S. |editor2=Garber, P.|year=2000|publisher=University of Chicago Press|isbn=0-226-06340-2|pages=447–448}}</ref> This may have more to do with foraging benefits to the squirrel monkeys than anti-predation benefits.<ref name="move" />

===Mating systems===
{{Multiple image |align= right |direction=vertical |image1=SAGUI-DE-TUFOS-BRANCOS.jpg|caption1=Two [[common marmoset]]s, the species lives in monogamous pairs|image2=Gelada group.jpg|caption2=[[Gelada]] harem: one male and multiple females}}
The [[mating system]]s of primates vary between [[Monogamy in animals|monogamy]], [[Polyandry in animals|polyandry]], [[Polygyny in animals|polygyny]] and [[polygynandry]]. In monogamous species, adult males and females form long-lasting [[pair bond]]s. Compared to other systems, there is little competition for mating rights and males and females tend to be similar in size. Polyandry, which involves groups consisting of single females mating with multiple males, may arise as a secondary mating system in monogamous species. In the [[brown-mantled tamarin]], a female may breeding with one or two males. Polyandry may have developed due to the high frequency of twin births, which require more help in raising.<ref name=Dixon>{{cite book|author=Dixon, Alan F|year=2012|title=Primate Sexuality|publisher=Oxford University Press|pages=32–61|isbn=9780199544646}}</ref> 

Polygynous species include gorillas, [[Northern plains gray langur|Hanuman langur]]s, geladas, hamadryas baboons, [[proboscis monkey]]s, and [[golden snub-nosed monkey]]s, and consists of one male mating with multiple females within a [[Harem (zoology)|harem]] or one-male unit. Sexual dimorphism tends to be higher in these species and males may also develop prominent [[secondary sex characteristic]]s. In the patriarchal hamadryas baboon, the males aggressively herd females into their groups and violently discipline those that wander. By contrast, in gelada society, which is based on female kinship, a male is dependent on the support of the females in his unit and cannot impose on them. Polygynous males must defend their harems from rivals, who may try to take over.<ref name=Dixon/>

In some species, such as ring-tailed lemurs, [[sifaka]]s, [[macaque]]s, most [[baboon]]s, [[mangabey]]s, [[squirrel monkey]]s, [[woolly monkey]]s, spider monkeys, [[woolly spider monkey]]s, chimpanzees and bonobos, both males and females mate with multiple partners. Polygynandry occurs in [[Multi-male group|multimale-multifemale groups]], and since females mate many times before conception, males have large testicles for [[sperm competition]]. Males may exist in a [[dominance hierarchy]] and those at the top will try to monopolize access to the females. Consortships may occur in some species but these are short-term. In solitary-living species, males and females mate with partners whose home ranges they overlap with. This is known as a 'dispersed' mating system.<ref name=Dixon/>

Genetic evidence indicates that humans were predominantly [[Polygyny|polygynous]] for most of their existence as a species, but that this began to shift during the Neolithic, when [[monogamy]] started becoming widespread concomitantly with the transition from nomadic to sedentary societies.<ref>{{Cite journal |last1=Dupanloup |first1=Isabelle |last2=Pereira |first2=Luisa |last3=Bertorelle |first3=Giorgio |last4=Calafell |first4=Francesc |last5=Prata |first5=Maria João |last6=Amorim |first6=Antonio |last7=Barbujani |first7=Guido |date=1 July 2003 |title=A Recent Shift from Polygyny to Monogamy in Humans Is Suggested by the Analysis of Worldwide Y-Chromosome Diversity |url=http://link.springer.com/10.1007/s00239-003-2458-x |journal=[[Journal of Molecular Evolution]] |volume=57 |issue=1 |pages=85–97 |doi=10.1007/s00239-003-2458-x |pmid=12962309 |bibcode=2003JMolE..57...85D |issn=0022-2844 |access-date=13 July 2024 |via=Springer Link}}</ref> Most modern human societies consist of monogamous [[marriage]]s, but allow for polygyny, particularly for those of a high status.<ref name=Dixon/>

===Sexual behavior=== 
[[File:Bonobo sexual behavior 1.jpg|thumb|[[Bonobo]]s mating, [[Jacksonville Zoo and Gardens]].]]
Female primates may signal to the male their receptiveness though various displays including eye-contact, tongue-clicking and presenting of the rump. Female lemurs, lorises and galagos will position themselves in the [[Lordosis behavior|lordosis]] pose while female chimpanzees, bonobos and some Old World monkeys develop [[sexual swelling]]s on the rump. Copulation in primates typically involves the males mounting the females from behind, as with most mammals. Belly-to-belly copulation has been recorded in apes, both gibbons and the great apes. Human [[sex position]]s are modifications of these two positions.<ref name=Dixon2/> 

Primates may engage in sexual activity as part of social bonding; including [[Homosexual behavior in animals|homosexual behaviour]].<ref name=Dixon2>{{cite book|author=Dixon, Alan F|year=2012|title=Primate Sexuality|publisher=Oxford University Press|pages=130–131, 135, 149–156, 200–202|isbn=9780199544646}}</ref> Such behavior play an important role in bonobo society in particular. female bonobos engage in mutual genital-rubbing behavior, possibly to bond socially with each other, thus forming a female nucleus of bonobo society. The bonding among females enables them to dominate most of the males.<ref name="deWaal1995">{{cite journal | vauthors = de Waal FB | title = Bonobo sex and society | journal = Scientific American | volume = 272 | issue = 3 | pages = 82–8 | date = March 1995 | pmid = 7871411 | doi = 10.1038/scientificamerican0395-82 | url = http://www.biozentrum.uni-wuerzburg.de/uploads/media/Bonobo_sex_01.pdf | access-date = 21 December 2011 | url-status = dead | bibcode = 1995SciAm.272c..82W | archive-url = https://web.archive.org/web/20120127051545/http://www.biozentrum.uni-wuerzburg.de/uploads/media/Bonobo_sex_01.pdf | archive-date = 27 January 2012 | author-link = Frans de Waal }}</ref>

===Life history===
[[File:Bonnet macaque nursing, Bangalore.jpg|A [[crab-eating macaque]] breastfeeding her baby|thumb]]
Primates have slower rates of development than other mammals. All primate infants are [[breastfeeding|breastfed]] by their mothers (with the exception of some human cultures and various zoo raised primates which are fed formula) and rely on them for grooming and transportation. In some species, infants are protected and transported by males in the group, particularly males who may be their fathers. Other relatives of the infant, such as siblings and aunts, may participate in its care as well. Most primate mothers cease [[ovulation]] while breastfeeding an infant; once the infant is [[Weaning|weaned]] the mother can reproduce again. This often leads to weaning conflict with infants who attempt to continue breastfeeding.<ref name="Strier2007" />

[[Infanticide in primates|Infanticide]] is common in polygynous species such as [[gray langur]]s and gorillas. Adult males may kill dependent offspring that are not theirs so the female will return to estrus and thus they can sire offspring of their own. Social monogamy in some species may have evolved to combat this behavior.<ref>{{cite journal|author1=Opie, Christopher |author2=Atkinson, Quentin D. |author3=Dunbarc, Robin I. M. |author4=Shultz, Susanne |year=2013|title=Male infanticide leads to social monogamy in primates|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=110|issue=33|pages=13328–13332|doi=10.1073/pnas.1307903110|pmid=23898180 |pmc=3746880|bibcode=2013PNAS..11013328O|doi-access=free }}</ref> Polygynandry may also lessen the risk of infanticide since paternity becomes uncertain.<ref>{{cite journal|author1=De Ruiter, Jan R.|author2= Van Hooff, Jan A. R. A. M.|author-link2=Jan van Hooff|author3=Scheffrahn, Wolfgang |name-list-style=amp |year=1994|title=Social and genetic aspects of paternity in wild long-tailed macaques (''Macaca fascicularis'')|journal=Behaviour|volume=129|issue=3–4|pages=203–24|jstor=4535195|doi=10.1163/156853994x00613}}</ref>

Primates have a longer [[Juvenile (organism)|juvenile]] period between weaning and sexual maturity than other mammals of similar size.<ref name="Strier2007" /> Some primates such as [[galago]]s and [[New World monkey]]s use tree-holes for [[Nest-building in primates|nesting]], and park juveniles in leafy patches while foraging. Other primates follow a strategy of "riding", i.e. carrying individuals on the body while feeding. Adults may construct or use nesting sites, sometimes accompanied by juveniles, for the purpose of resting, a behavior which has developed secondarily in the great apes.<ref name="Kappeler1998">{{cite journal |last1=Kappeler |first1=Peter M. |year=1998 |title=Nests, Tree Holes, and the Evolution of Primate Life Histories |journal=American Journal of Primatology |issue=1 |pages=7–33 |pmid=9730211 |doi=10.1002/(SICI)1098-2345(1998)46:1<7::AID-AJP3>3.0.CO;2-# |volume=46|s2cid=196589387 }}</ref><ref name="Ross2001">{{cite journal |last1=Ross |first1= Caroline|year=1991 |title=Park or ride? Evolution of infant carrying in primates. |journal=International Journal of Primatology |publisher=Kluwer Academic Publishing |volume=22 |issue=5 |pages=749–771 |doi=10.1023/A:1012065332758|s2cid= 25301078}}</ref> During the juvenile period, primates are more susceptible than adults to [[predation]] and [[starvation]]; they gain experience in feeding and avoiding predators during this time.<ref name="Strier2007" /> They learn social and fighting skills, often through playing.<ref name="Strier2007" /> Primates, especially females, have longer lifespans than other similarly sized mammals,<ref name="Strier2007" /> this may be partially due to their slower metabolisms.<ref>{{cite web |url=http://www.ibtimes.com/humans-primates-burn-50-percent-fewer-calories-each-day-other-mammals-1539866 |title=Humans And Primates Burn 50 Percent Fewer Calories Each Day Than Other Mammals |last1=Mintz |first1=Zoe |date=14 January 2014 |website=www.ibtimes.com |publisher=IBT Media Inc. |access-date=2014-01-14}}</ref> Late in life, female catarrhine primates appear to undergo a cessation of reproductive function known as [[menopause]]; other groups are less studied.<ref name="MLWalker2">{{cite journal | author = Walker ML, Herndon JG | title = Menopause in nonhuman primates? | journal = Biology of Reproduction | volume = 79 | pages = 398–406 | year = 2008 | pmid = 18495681 | doi = 10.1095/biolreprod.108.068536 | issue = 3 | pmc=2553520 | last2 = Herndon}}</ref>

===Diet and feeding===
{{Multiple image |align= right |direction=vertical |image1=Colubusmonkey.JPG|caption1=Leaf eating [[mantled guereza]]|image2=Microcebus murinus -Artis Zoo, Amsterdam, Netherlands-8c.jpg|thumb|caption2=A [[mouse lemur]] eating fruit}}
Primates exploit a variety of food sources. It has been said that many characteristics of modern primates, including humans, derive from an early ancestor's practice of taking most of its food from the tropical canopy.<ref>{{cite magazine |last1 = Milton |first1 = K. |year=1993 |title = Diet and Primate Evolution |url=http://nature.berkeley.edu/miltonlab/pdfs/diet_primate_evolution.pdf |magazine=Scientific American |volume=269 |issue=2 |pages=86–93 |pmid=8351513 |doi=10.1038/scientificamerican0893-86|bibcode=1993SciAm.269b..86M}}</ref> Most primates include fruit in their diets to obtain easily digested nutrients including [[carbohydrate]]s and [[lipid]]s for energy.<ref name="Strier2007" /> Primates in the suborder [[Strepsirrhini]] (non-tarsier prosimians) are able to synthesize [[vitamin C]], like most other mammals, while primates of the suborder [[Haplorhini]] (tarsiers, monkeys and apes) have lost this ability, and require the vitamin in their diet.<ref>{{cite journal | title = Vitamin C biosynthesis in prosimians: Evidence for the anthropoid affinity of Tarsius | last1 = Pollock | first1 = J. I. | last2 = Mullin | first2 = R. J. | 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 = 2012-06-28 | pmid = 3113259 | accessdate = 2010-03-16 }}</ref>

Many primates have anatomical specializations that enable them to exploit particular foods, such as fruit, leaves, gum or [[insect]]s.<ref name="Strier2007" /> For example, leaf eaters such as howler monkeys, [[black-and-white colobus]]es and [[sportive lemur]]s have extended digestive tracts which enable them to absorb nutrients from leaves that can be difficult to digest.<ref name="Strier2007" /> [[Marmoset]]s, which are gum eaters, have strong [[incisor]] teeth, enabling them to open tree bark to get to the gum, and claws rather than nails, enabling them to cling to trees while feeding.<ref name="Strier2007" /> The [[aye-aye]] combines rodent-like teeth with a long, thin middle finger to fill the same ecological niche as a woodpecker. It taps on trees to find insect larvae, then gnaws holes in the wood and inserts its elongated middle finger to pull the larvae out.<ref>{{cite journal |last1=Milliken |first1=G. W. |last2=Ward |first2=J. P. |last3=Erickson |first3=C. J. |year=1991 |title=Independent digit control in foraging by the aye-aye (''Daubentonia madagascariensis'') |journal=Folia Primatologica |volume=56 |issue=4 |pages=219–224 |doi=10.1159/000156551 |pmid=1937286}}</ref> Some species have additional specializations. For example, the [[grey-cheeked mangabey]] has thick [[tooth enamel|enamel]] on its teeth, enabling it to open hard fruits and seeds that other monkeys cannot.<ref name="Strier2007" /> The [[gelada]] is the only primate species that feeds primarily on grass.<ref name="Hiller">{{cite web |last=Hiller|first=C.|year=2000|url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Theropithecus_gelada.html|title = ''Theropithecus gelada''|access-date=2008-08-08|work=Animal Diversity Web}}</ref>

====Hunting====
[[File:COLLECTIE TROPENMUSEUM Portret van een Dajak jager op Borneo met een gevangen zwijn over de schouder TMnr 60043389.jpg|thumb|upright|alt=Portrait of a Dayak hunter in Borneo with a boar over his shoulder|Humans have traditionally hunted prey for subsistence.]]
[[Tarsier]]s are the only [[Extant taxon|extant]] [[obligate carnivore|obligate carnivorous]] primates, exclusively eating insects, crustaceans, small vertebrates and snakes (including [[Venomous snake|venomous]] species).<ref>{{cite book|title=Tarsiers Past, Present and Future|chapter=Introduction|last1=Wright |first1=P. |last2=Simmons |first2=E. |last3=Gursky |first3=S.|editor=Wright, P. |editor2=Simmons, E. |editor3=Gursky, S.|year=2003|publisher=Rutgers University Press|isbn=0-8135-3236-1|pages=1}}</ref> [[Capuchin monkey]]s can exploit many different types of plant matter, including fruit, leaves, flowers, buds, nectar and seeds, but also eat insects and other [[invertebrate]]s, bird eggs, and small vertebrates such as birds, [[lizard]]s, [[squirrel]]s and [[bat]]s.<ref name="Sussman2003" />

The [[common chimpanzee]] eats an [[omnivorous]] [[frugivorous]] diet. It prefers fruit above all other food items and even seeks out and eats them when they are not abundant. It also eats leaves and leaf buds, seeds, blossoms, stems, pith, bark and resin. Insects and meat make up a small proportion of their diet, estimated as 2%.<ref name=Goodall1986>{{cite book | last = Goodall | first = Jane | author-link = Jane Goodall | year = 1986 | title = The Chimpanzees of Gombe: Patterns of Behavior | publisher = Belknap Press of Harvard University Press | isbn = 0-674-11649-6 | url = https://archive.org/details/chimpanzeesofgom00good}}</ref><ref>{{cite web|last=Guernsey|first=Paul|title=WHAT DO CHIMPS EAT?|url=http://www.allaboutwildlife.com/what-do-chimps-eat|work=All About Wildlife|access-date=2013-04-22|archive-date=2019-11-18|archive-url=https://web.archive.org/web/20191118084847/http://www.allaboutwildlife.com/what-do-chimps-eat|url-status=dead}}</ref> The meat consumption includes predation on other primate species, such as the [[western red colobus]] monkey.<ref name="tai2007"/> The [[bonobo]] is an [[omnivorous]] [[frugivore]] – the majority of its diet is fruit, but it supplements this with leaves, meat from small [[vertebrate]]s, such as [[anomalure]]s, [[flying squirrel]]s and [[duiker]]s,<ref>{{cite journal |author=Ihobe H |title=Observations on the meat-eating behavior of wild bonobos (''Pan paniscus'') at Wamba, Republic of Zaire |journal=Primates |volume=33 |issue=2 |pages=247–250|year=1992 |doi=10.1007/BF02382754|s2cid=10063791 }}</ref> and [[invertebrate]]s.<ref>{{cite book |last1=Rafert |first1=J. |first2=E.O. |last2=Vineberg |year=1997 |chapter=Bonobo Nutrition – relation of captive diet to wild diet |chapter-url=http://www.nagonline.net/HUSBANDRY/Diets%20pdf/Bonobo%20Nutrition.pdf |url-status=dead |archive-url=https://web.archive.org/web/20120425232556/http://www.nagonline.net/HUSBANDRY/Diets%20pdf/Bonobo%20Nutrition.pdf |archive-date=2012-04-25 |title=Bonobo Husbandry Manual |publisher=American Association of Zoos and Aquariums}}</ref> In some instances, bonobos have been shown to consume lower-order primates.<ref>{{cite journal |last1=Surbeck |first1=M |last2=Fowler |first2=A |last3=Deimel |first3=C |last4=Hohmann |first4=G |title=Evidence for the consumption of arboreal, diurnal primates by bonobos (''Pan paniscus'') |journal=American Journal of Primatology |volume=71 |issue=2 |pages=171–4 |year=2008 |doi=10.1002/ajp.20634 |pmid=19058132|s2cid=32622605 }}</ref><ref name=Surbeck>{{cite journal |author=Surbeck M, Hohmann G |title=Primate hunting by bonobos at LuiKotale, Salonga National Park |journal=Current Biology |volume=18 |issue=19 |pages=R906–7 |date=14 October 2008 |doi=10.1016/j.cub.2008.08.040 |pmid=18957233 |last2=Hohmann|s2cid=6708310 |doi-access=free |bibcode=2008CBio...18.R906S }}</ref>

Until the development of agriculture approximately 10,000 years ago, ''Homo sapiens'' employed a hunter-gatherer method as their sole means of food collection. This involved combining stationary food sources (such as fruits, grains, tubers, and mushrooms, insect larvae and aquatic mollusks) with [[Game (food)|wild game]], which must be hunted and killed in order to be consumed.<ref>{{cite journal |author=Cordain L |title=Origins and evolution of the Western diet: health implications for the 21st century |journal=Am. J. Clin. Nutr. |volume=81 |issue=2 |pages=341–54 |date=February 2005 |pmid=15699220 |name-list-style=vanc |author2=Eaton SB |author3=Sebastian A |display-authors=3 |last4=Mann |first4=N |last5=Lindeberg |first5=S |last6=Watkins |first6=BA |last7=O'Keefe |first7=JH |last8=Brand-Miller |first8=J |doi=10.1093/ajcn.81.2.341|doi-access=free }}</ref> It has been proposed that humans have used fire to prepare and [[cooking|cook]] food since the time of ''[[Homo erectus]]''.<ref>{{cite journal |author=Ulijaszek SJ |title=Human eating behaviour in an evolutionary ecological context |journal=Proc Nutr Soc |volume=61 |issue=4 |pages=517–26 |date=November 2002 |pmid=12691181 |doi=10.1079/PNS2002180|doi-access=free }}</ref> Around ten thousand years ago, [[History of agriculture|humans developed agriculture]],<ref>[http://www.archaeology.org/9707/newsbriefs/squash.html Earliest agriculture in the Americas] {{webarchive|url=https://web.archive.org/web/20100603232246/http://www.archaeology.org/9707/newsbriefs/squash.html |date=3 June 2010 }}
[http://sciencenow.sciencemag.org/cgi/content/full/2007/213/2 Earliest cultivation of barley]  {{webarchive|url=https://web.archive.org/web/20070216093200/http://sciencenow.sciencemag.org/cgi/content/full/2007/213/2 |date=16 February 2007 }}
[http://news.bbc.co.uk/2/hi/science/nature/5038116.stm Earliest cultivation of figs] {{webarchive|url=https://web.archive.org/web/20060602081110/http://news.bbc.co.uk/2/hi/science/nature/5038116.stm |date=2 June 2006 }}, retrieved 19 February 2007</ref> which substantially altered their diet. This change in diet may also have altered human biology; with the spread of [[dairy farming]] providing a new and rich source of food, leading to the evolution of the ability to digest [[lactose]] in some adults.<ref>{{cite journal |author=Krebs JR |title=The gourmet ape: evolution and human food preferences |journal=Am. J. Clin. Nutr. |volume=90 |issue=3 |pages=707S–11S |date=September 2009 |pmid=19656837 |doi=10.3945/ajcn.2009.27462B|doi-access=free }}</ref><ref>{{cite journal |vauthors=Holden C, Mace R |title=Phylogenetic analysis of the evolution of lactose digestion in adults |url=https://archive.org/details/sim_human-biology_1997-10_69_5/page/605 |journal=Hum. Biol. |volume=69 |issue=5 |pages=605–28 |date=October 1997 |pmid=9299882}}</ref>

===As prey===
Predators of primates include various species of [[carnivora]]ns, [[birds of prey]], [[reptile]]s, and other primates. Even gorillas have been recorded as prey. Predators of primates have diverse hunting strategies and as such, primates have evolved several different [[antipredator adaptation]]s including [[crypsis]], [[Alarm signal|alarm calls]] and [[Mobbing (animal behavior)|mobbing]]. Several species have separate alarm calls for different predators such as air-borne or ground-dwelling predators. Predation may have shaped group size in primates as species exposed to higher predation pressures appear to live in larger groups.<ref>{{cite book|author=Fichtel, Claudia|contribution=Predation|year=2012|title=The Evolution of Primate Societies|editor1=Mitani, John C. |editor2=Call, Josep |editor3=Kappeler, Peter M. |editor4=Palombit, Ryne A. |editor5=Silk, Joan B. |editor5-link=Joan Silk |publisher=University of Chicago Press|pages=169–84|isbn=978-0-226-53172-4}}</ref>

===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 Doué-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=3328–3332 |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=212–225|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=8184–8189|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=393–6|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&nbsp;kHz with a dominant frequency of 70&nbsp;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 = 508–511 | 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=1169–1199|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=249–256|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=1070–1094 |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=97–105 |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=131–139 |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&nbsp;million years ago) from ''[[Pan (genus)|Pan]]'' (5 to 6&nbsp;million years ago) to the emergence of full [[behavioral modernity]] some 50,000–150,000 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 = 205–23 |date=Aug 2000 | doi = 10.1006/jhev.2000.0415 | pmid = 10968929 | s2cid = 7403772 | doi-access = free | bibcode = 2000JHumE..39..205A }}</ref>

===Intelligence and cognition===
{{Main|Primate cognition}}
Primates have advanced cognitive abilities: some make tools and use them to acquire food and for social displays;<ref>{{cite journal |last1=Boesch |first1=C. |last2=Boesch |first2=H. |title=Tool Use and Tool Making in Wild Chimpanzees |journal=[[Folia Primatologica]] |year=1990 |volume=54 |pages=86–99 |doi=10.1159/000156428 |pmid=2157651 |issue=1–2}}</ref><ref>{{cite journal |last1=Westergaard |year=1998 |title=Why some capuchin monkeys (''Cebus apella'') use probing tools (and others do not) |url=https://archive.org/details/sim_journal-of-comparative-psychology_1998-06_112_2/page/207 |journal=Journal of Comparative Psychology |volume=112 |issue=2 |pages=207–211 |doi=10.1037/0735-7036.112.2.207 |pmid=9642788 |first1=G. C. |last2=Lundquist|first2=A. L.|display-authors=2 |last3=Haynie |first3=Michael Katherine |last4=Kuhn |first4=Heather E. |last5=Suomi |first5=Stephen J.}}</ref> some can perform tasks requiring cooperation, influence and rank;<ref>{{cite journal |last1=de Waal |first1=F. B. M. |last2=Davis |first2=J. M. |year=2003 |title=Capuchin cognitive ecology: cooperation based on projected returns |journal=Neuropsychologia |volume=41 |pages=221–228 |doi=10.1016/S0028-3932(02)00152-5 |pmid=12459220 |issue=2|s2cid=8190458 }}</ref> they are status conscious, manipulative and capable of deception;<ref>{{cite journal |last1=Paar |first1=L. A. |last2=Winslow |first2=J. T. |last3=Hopkins |first3=W. D. |last4=de Waal |first4=F. B. M. |year=2000 |title=Recognizing facial cues: Individual discrimination by chimpanzees (''Pan troglodytes'') and rhesus monkeys (''Macaca mulatta'') |url=https://archive.org/details/sim_journal-of-comparative-psychology_2000-03_114_1/page/47 |journal=Journal of Comparative Psychology |volume=114 |issue=1 |pages=47–60 |doi=10.1037/0735-7036.114.1.47 |pmid=10739311 |pmc=2018744}}</ref><ref name="Byrne and Corp">{{cite journal |last=Byrne |first=Richard |author2=Corp, Nadia |title=Neocortex size predicts deception rate in primates |journal= Proceedings of the Royal Society of London. Series B: Biological Sciences|year=2004 |volume=271 |issue=1549 |pages=1693–1699 |doi=10.1098/rspb.2004.2780|pmid=15306289 |pmc=1691785}}</ref> they can recognise [[Kinship|kin]] and [[conspecific]]s;<ref>{{cite journal |last1=Paar |first1=L. A. |last2=de Waal |first2=F. B. M. |year=1999 |title=Visual kin recognition in chimpanzees |journal=Nature |volume=399 |pages=647–648 |doi=10.1038/21345 |pmid=10385114 |issue=6737|bibcode = 1999Natur.399..647P|s2cid=4424086 }}</ref><ref>{{cite journal |year=1997 |title=Discrimination of macaques by macaques: The case of sulawesi species |journal=Primates |volume=38 |issue=3 |pages=233–245 |last1=Fujita |first1=K. |last2=Watanabe |first2=K. |last3=Widarto |first3=T. H. |last4=Suryobroto |first4=B. |doi=10.1007/BF02381612|s2cid=21042762 }}</ref> and they can learn to use symbols and understand aspects of human language including some relational syntax and concepts of number and numerical sequence.<ref>{{cite journal |last=Call |first=J. |year=2001 |title=Object permanence in orangutans (''Pongo pygmaeus''), chimpanzees (''Pan troglodytes''), and children (''Homo sapiens'') |url=https://archive.org/details/sim_journal-of-comparative-psychology_2001-06_115_2/page/159 |journal=Journal of Comparative Psychology |volume=115 |issue=2 |pages=159–171 |doi=10.1037/0735-7036.115.2.159 |pmid=11459163}}</ref><ref>{{cite journal |last1=Itakura |first1=S. |last2=Tanaka |first2=M. |date=June 1998 |title=Use of experimenter-given cues during object-choice tasks by chimpanzees (''Pan troglodytes''), an orangutan (''Pongo pygmaeus''), and human infants (''Homo sapiens'') |url=https://archive.org/details/sim_journal-of-comparative-psychology_1998-06_112_2/page/119 |journal=Journal of Comparative Psychology |volume=112 |issue=2 |pages=119–126 |doi=10.1037/0735-7036.112.2.119 |pmid=9642782}}</ref><ref>{{cite journal |last1=Gouteux |first1=S. |last2=Thinus-Blanc |first2=C. |last3=Vauclair |first3=J. |year=2001 |title=Rhesus monkeys use geometric and nongeometric information during a reorientation task |journal=Journal of Experimental Psychology: General |volume=130 |issue=3 |pages=505–519 |doi=10.1037/0096-3445.130.3.505 |pmid=11561924 |url=http://cogprints.org/3590/1/Gouteux_et_al_JEPGEN_01.pdf}}</ref> Research in primate cognition explores problem solving, memory, social interaction, a [[Theory of mind#Non-human|theory of mind]], and numerical, spatial, and abstract concepts.<ref>{{cite book |author1=Tomasello, M. |author2=Call, J. |name-list-style=amp |title=Primate Cognition |publisher=Oxford University Press US |year=1997 |isbn=978-0-19-510624-4}}</ref> Comparative studies show a trend towards higher intelligence going from prosimians to New World monkeys to Old World monkeys, and significantly higher average cognitive abilities in the great apes.<ref name="Deaner">{{cite journal | last1 = Deaner | first1 = R. O. | last2 = van Schaik | first2 = C. P. | last3 = Johnson | first3 = V. E. | year = 2006 | title = Do some taxa have better domain-general cognition than others? A metaanalysis of nonhuman primate studies | journal = [[Evolutionary Psychology (journal)|Evolutionary Psychology]] | volume = 4 | pages = 149–196 | doi = 10.1177/147470490600400114| s2cid = 16702785 | doi-access = free }}</ref><ref name="Reader">{{Cite journal | last1 = Reader | first1 = S. M. | last2 = Hager | first2 = Y. | last3 = Laland | first3 = K. N. | title = The evolution of primate general and cultural intelligence | journal = [[Philosophical Transactions of the Royal Society B]] | volume = 366 | issue = 1567 | pages = 1017–1027 | date = 2011 | url = http://lalandlab.st-andrews.ac.uk/pdf/Publication163.pdf | doi = 10.1098/rstb.2010.0342 | pmid = 21357224 | access-date = 2011-07-04 | url-status=dead | archive-url = https://web.archive.org/web/20111003152030/http://lalandlab.st-andrews.ac.uk/pdf/Publication163.pdf | archive-date = 2011-10-03 | pmc = 3049098}}</ref> However, there is a great deal of variation in each group (e.g., among New World monkeys, both [[spider monkey|spider]]<ref name = "Deaner"/> and [[capuchin monkey]]s<ref name = "Reader"/> have scored highly by some measures), as well as in the results of different studies.<ref name = "Deaner"/><ref name = "Reader"/>

====Tool use and manufacture====
{{Main|Tool use by animals}}
[[File:Cultural-differences-in-ant-dipping-tool-length-between-neighbouring-chimpanzee-communities-at-srep12456-s2.ogv|thumb|right|[[Chimpanzee]]s using twigs to dip for ants]]
[[File:Macaca fascicularis aurea stone tools - journal.pone.0072872.g002.png|thumb|[[Crab-eating macaque]]s with [[stone tool]]s]]

In 1960, [[Jane Goodall]] observed a [[chimpanzee]] poking pieces of grass into a [[termite]] mound and then raising the grass to his mouth. After he left, Goodall approached the mound and repeated the behaviour because she was unsure what the chimpanzee was doing. She found that the termites bit onto the grass with their jaws. The chimpanzee had been using the grass as a tool to "fish" or "dip" for termites.<ref name="The Jane Goodall Institute1">{{cite web|url=https://www.janegoodall.org.uk/chimpanzees/chimpanzee-central/15-chimpanzees/chimpanzee-central/19-toolmaking|title=Toolmaking|publisher=The Jane Goodall Institute|access-date=2013-08-01}}</ref> There are more limited reports of the closely related [[bonobo]] using tools in the wild; it has been claimed they rarely use tools in the wild although they use tools as readily as chimpanzees when in captivity.<ref name="ApeTag">{{cite web |url=http://www.clemetzoo.com/apetag/Bonobos.html |title=Bonobos |publisher=ApeTag |year=2010 |access-date=2013-08-03 |url-status=dead |archive-url=https://web.archive.org/web/20131102050710/https://www.clemetzoo.com/apetag/Bonobos.html |archive-date=2013-11-02 }}</ref> It has been reported that females, both chimpanzee and bonobo, use tools more avidly than males.<ref>
{{cite journal|last1=Gruber |first1=T. |last2=Clay |first2=Z. |last3=Zuberbühler |first3=K.|year=2010|title= A comparison of bonobo and chimpanzee tool use: evidence for a female bias in the ''Pan'' lineage|journal= Animal Behaviour|volume= 80|issue=6|pages= 1023–1033|url=http://www.emory.edu/LIVING_LINKS/publications/articles/Gruber_etal_2010.pdf|doi=10.1016/j.anbehav.2010.09.005|s2cid=14923158 }}</ref> [[Orangutan]]s in [[Borneo]] scoop catfish out of small ponds. Over two years, anthropologist Anne Russon observed orangutans learning to jab sticks at catfish to scare them out of the ponds and in to their waiting hands.<ref>{{cite magazine|url=https://www.wired.com/wiredscience/2011/04/orangutan-tools-fishing/ |title=Orangutans use simple tools to catch fish |author=Bower, B. |magazine=[[Wired (website)|Wired]] |date=18 April 2011|access-date=2013-08-05}}</ref> There are few reports of [[gorilla]]s using tools in the wild. An adult female [[western lowland gorilla]] used a branch as a walking stick apparently to test water depth and to aid her in crossing a pool of water. Another adult female used a detached trunk from a small shrub as a stabilizer during food gathering, and another used a log as a bridge.<ref>{{cite journal|last1=Breuer |first1=T. |last2=Ndoundou-Hockemba |first2=M. |last3=Fishlock |first3=V.|year=2005|title= First observation of tool use in wild gorillas|journal= PLOS Biology |volume=3|issue=11|pages=e380 |doi=10.1371/journal.pbio.0030380|pmid=16187795|pmc=1236726 |doi-access=free }}</ref>

The first direct observation of a non-ape primate using a tool in a wild environment occurred in 1988. Primatologist Sue Boinski watched an adult male white-faced capuchin beat a [[Bothrops asper|fer-de-lance]] snake to death with a dead branch.<ref name="Boinski1988">{{cite journal | title = Use of a club by a wild white-faced capuchin (''Cebus capucinus'') to attack a venomous snake (''Bothrops asper'')| year = 1988| last1 = Boinski| first1 = S.| journal = American Journal of Primatology| volume = 14| issue = 2| pages = 177–179| pmid = 31973450| s2cid = 84653622 |doi = 10.1002/ajp.1350140208}}</ref> The black-striped capuchin was the first non-ape primate for which routine tool use was documented in the wild; individuals were observed [[Primate archaeology|cracking nuts]] by placing them on a stone anvil and hitting them with another large stone.<ref name="Fragaszy">{{cite journal|last1=Fragaszy |first1=D. |last2=Izar |first2=P. |last3=Visalberghi |first3=E. |last4=Ottoni |first4=E.B. |last5=de Oliveira |first5=M.G.|year=2004|title= Wild capuchin monkeys (''Cebus libidinosus'') use anvils and stone pounding tools|journal= American Journal of Primatology|volume=64|issue=4|pages= 359–366|doi= 10.1002/ajp.20085|pmid=15580579|s2cid=16222308 }}</ref> In Thailand and Myanmar, [[crab-eating macaque]]s use stone tools to open nuts, oysters and other bivalves, and various types of sea snails.<ref>{{cite journal|last1=Gumert|first1=M.D.|last2=Kluck|first2=M.|last3=Malaivijitnond|first3=S.|title=The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the Andaman Sea region of Thailand|journal=American Journal of Primatology|year=2009|volume=71|issue=7|pages=594–608|doi=10.1002/ajp.20694|pmid=19405083|s2cid=22384150}}</ref> Chacma baboons use stones as weapons; stoning by these baboons is done from the rocky walls of the canyon where they sleep and retreat to when they are threatened. Stones are lifted with one hand and dropped over the side whereupon they tumble down the side of the cliff or fall directly to the canyon floor.<ref>{{cite journal |last1=Hamilton |first1=W.J. |last2=Buskirk |first2=R.E. |last3=Buskirk |first3=W.H.|year=1975|title= Defensive stoning by baboons|journal= Nature|volume= 256|pages= 488–489| doi=10.1038/256488a0|issue=5517|bibcode=1975Natur.256..488H|s2cid=4149862 }}</ref>

Although they have not been observed to use tools in the wild, lemurs in controlled settings have been shown to be capable of understanding the functional properties of the objects they had been trained to use as tools, performing as well as tool-using haplorhines.<ref>{{cite book | last1 = Fichtel | first1 = C. | last2 = Kappeler | first2 = P. M. | chapter = Chapter 19: Human universals and primate symplesiomorphies: Establishing the lemur baseline | editor1-last = Kappeler | editor1-first = P. M. | editor2-last = Silk | editor2-first = J. B. | title = Mind the Gap: Tracing the Origins of Human Universals | publisher = Springer | year = 2010 | isbn = 978-3-642-02724-6 | chapter-url = https://books.google.com/books?id=MFzxVH_OxjsC&pg=PA395}}</ref>

Soon after her initial discovery of tool use, Goodall observed other chimpanzees picking up leafy twigs, stripping off the leaves and using the stems to fish for insects. This change of a leafy twig into a tool was a major discovery. Prior to this, scientists thought that only humans manufactured and used tools, and that this ability was what separated humans from other animals.<ref name="The Jane Goodall Institute1" /> Chimpanzees have also been observed making "sponges" out of leaves and moss that suck up water.<ref>{{cite journal | author=Sugiyama, Y. | year=1995 | title=Drinking tools of wild chimpanzees at Bossou | journal=American Journal of Primatology | volume=37 | issue=1 | pages=263–269 | doi=10.1002/ajp.1350370308| pmid=31936951| s2cid=86473603 }}</ref> Sumatran orangutans have been observed making and using tools. They will break off a tree branch that is about 30&nbsp;cm long, snap off the twigs, fray one end and then use the stick to dig in tree holes for termites.<ref name="OrangutanIslands">{{cite web|url=http://orangutanislands.com/sumatra-orangutans.htm |title=Sumatran orangutans |publisher=OrangutanIslands.com |access-date=2013-08-02 |url-status=usurped |archive-url=https://web.archive.org/web/20131126074458/https://orangutanislands.com/sumatra-orangutans.htm |archive-date=2013-11-26}}</ref><ref>{{cite journal|last1=van Schaik |first1=C. |last2=Fox |first2=E. |last3=Sitompul |first3=A.|year=1996|title= Manufacture and use of tools in wild Sumatran orangutans|url=https://archive.org/details/sim_naturwissenschaften_1996-04_83_4/page/186 |journal= Naturwissenschaften|volume= 83|issue=4|pages= 186–188|doi=10.1007/BF01143062|pmid=8643126 |bibcode=1996NW.....83..186V|s2cid=27180148 }}</ref> In the wild, mandrills have been observed to clean their ears with modified tools. Scientists filmed a large male mandrill at [[Chester Zoo]] (UK) stripping down a twig, apparently to make it narrower, and then using the modified stick to scrape dirt from underneath its toenails.<ref name="Gill2011">{{cite web | url = http://www.bbc.co.uk/nature/14227783 | title = Mandrill monkey makes 'pedicuring' tool | first = Victoria | last = Gill | publisher = BBC | date = 22 July 2011 | access-date = 2013-08-11}}</ref> Captive gorillas have made a variety of tools.<ref name="Vancatova">{{cite web|url=http://www.rozhlas.cz/therevealed/comments/_zprava/488947|author=Vancatova, M.|title=Gorillas and Tools – Part I|year=2008|access-date=2013-08-04}}</ref>