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{{For|an explanation of very similar terms|Plesiadapiformes}}{{About|the type of animal}}{{pp-move-indef}}{{short description|An order of mammals}}{{automatic taxobox| name = Primates
Paleocene–Holocene>Present, {{Fossil range0}}| image = File:Primates - some families.jpg|300pxrect 0 0 188 142 Aye-ayerect 0 250 188 142 Ring-tailed lemurrect 0 250 188 380 Capuchin monkeyrect 0 380 188 515 Spider monkeyrect 0 515 188 820 Gibbonrect 376 0 188 142 Tarsierrect 376 250 188 142 Red slender lorisrect 376 250 188 380 Lion tamarinrect 376 380 188 515 Hamadryas baboonrect 376 515 188 820 Common chimpanzee| image_caption = Some primate families, from top to bottom: Daubentoniidae, Tarsiidae, Lemuridae, Lorisidae, Cebidae, Callitrichidae, Atelidae, Cercopithecidae, Hylobatidae, Hominidae| taxon = PrimatesCarl Linnaeus>Linnaeus, 1758{{MSW3 Grovesid=12100001}}| subdivision_ranks = Suborders, Infraorders, Parvorders, Superfamilies, and Families (16)Primate#Classification of living primates>classification| range_map = Range of Non-human Primates.png| range_map_caption = Range of the nonhuman primates (green)}}A primate ({{IPAc-en|audio=En-us-primate.ogg|ˈ|p|r|aɪ|m|eɪ|t}} {{respell|PRY|mayt}}) (from Latin primat-, from primus: "prime, first rank") is a eutherian mammal constituting the taxonomic order Primates. Primates arose 85–55 million years ago from small terrestrial mammals (Primatomorpha), which adapted to living in the trees of tropical forests: many primate characteristics represent adaptations to life in this challenging environment, including large brains, visual acuity, color vision, altered shoulder girdle, and dexterous hands. Primates range in size from Madame Berthe's mouse lemur, which weighs {{convert|30|g|oz|0|abbr=on}}, to the eastern gorilla, weighing over {{convert|200|kg|lb|abbr=on}}. There are 190–448 species of living primates, depending on which classification is used. New primate species continue to be discovered: over 25 species were described in the first decade of the 2000s, and eleven since 2010.Primates are divided into two distinct suborders (see diagram under History of terminology). The first is the strepsirrhines (from Greek 'wet-nosed') – lemurs, galagos, and lorisids. (The colloquial names ending in -nosed actually refer to the rhinarium of the primate.) The second is haplorhines - the "dry-nosed" primates (from Greek 'simple-nosed') – tarsier, monkey, and ape clades, the last of these including humans. Simians (infraorder Simiiformes, from Greek simos 'flat-nosed') are monkeys and apes, cladistically including: the catarrhines (from Latin 'narrow nosed') consisting of the Old World monkeys and apes; and the platyrrhines (from Latin 'flat-nosed'): this division occurred about 60 million years ago. Forty million years ago, simians from Africa migrated to South America, presumably by drifting on debris, and gave rise to the New World monkeys. Twenty five million years ago the remaining Old World simians (catarrhines) split into apes and Old World monkeys. Common names for the simians are the (Old World) baboons, macaques, gibbons, and great apes; and the (New World) capuchins, howlers and squirrel monkeys.Primates have large brains (relative to body size) compared to other mammals, as well as an increased reliance on visual acuity at the expense of the sense of smell, which is the dominant sensory system in most mammals. These features are more developed in monkeys and apes, and noticeably less so in lorises and lemurs. Some primates are trichromats, with three independent channels for conveying color information. Except for apes, primates have tails. Most primates also have opposable thumbs. Many species are sexually dimorphic; differences may include muscle mass, fat distribution, pelvic width, canine tooth size, hair distribution, and coloration. Primates have slower rates of development than other similarly sized mammals, reach maturity later, and have longer lifespans. Depending on the species, adults may live in solitude, in mated pairs, or in groups of up to hundreds of members. Some primates, including gorillas, humans and baboons, are primarily terrestrial rather than arboreal, but all species have adaptations for climbing trees. Arboreal locomotion techniques used include leaping from tree to tree and swinging between branches of trees (brachiation); terrestrial locomotion techniques include walking on two limbs (bipedalism) and modified walking on four limbs (knuckle-walking).Primates are among the most social of animals, forming pairs or family groups, uni-male harems, and multi-male/multi-female groups. Non-human primates have at least four types of social systems, many defined by the amount of movement by adolescent females between groups. Most primate species remain at least partly arboreal: the exceptions are some great apes, baboons, and humans, who left the trees for the ground and now inhabit every continent.Close interactions between humans and non-human primates (NHPs) can create opportunities for the transmission of zoonotic diseases, especially virus diseases, including herpes, measles, ebola, rabies, and hepatitis. Thousands of non-human primates are used in research around the world because of their psychological and physiological similarity to humans. About 60% of primate species are threatened with extinction. Common threats include deforestation, forest fragmentation, monkey drives, and primate hunting for use in medicines, as pets, and for food. Large-scale tropical forest clearing for agriculture most threatens primates.

History of terminology

The English name "primates" is derived from Old French or French primat, from a noun use of Latin primat-, from primus ("prime, first rank").ENCYCLOPEDIA, Primate, Merriam-Webster Online Dictionary, Merriam-Webster,weblink 2008-07-21, The name was given by Carl Linnaeus because he thought this the "highest" order of animals.BOOK, The Book of Popular Science, 257, 1963,weblink The relationships among the different groups of primates were not clearly understood until relatively recently, so the commonly used terms are somewhat confused. For example, "ape" has been used either as an alternative for "monkey" or for any tailless, relatively human-like primate.{{Citation |last=Anon. |year=1911 |contribution=Ape |title=Encyclopædia Britannica |location=New York |publisher=Encyclopædia Britannica |url=https://archive.org/stream/encyclopaediabri02chisrich#page/160/mode/2up |accessdate=2011-07-10 |edition=11th |volume=XIX}}Sir Wilfrid Le Gros Clark was one of the primatologists who developed the idea of trends in primate evolution and the methodology of arranging the living members of an order into an "ascending series" leading to humans.{{Citation |last=Dixson |first=A.F. |year=1981 |title=The Natural History of the Gorilla |location=London |publisher=Weidenfeld & Nicolson |isbn=978-0-297-77895-0}} Commonly used names for groups of primates such as "prosimians", "monkeys", "lesser apes", and "great apes" reflect this methodology. According to our current understanding of the evolutionary history of the primates, several of these groups are paraphyletic: a paraphyletic group is one which does not include all the descendants of the group's common ancestor.Definitions of paraphyly vary; for the one used here see e.g. {{Citation|last=Stace |first=Clive A. |authorlink=Clive A. Stace |year=2010 |title=Classification by molecules: What's in it for field botanists? |journal=Watsonia |volume=28 |pages=103–122 |url=http://www.watsonia.org.uk/Wats28p103.pdf |accessdate=2010-02-07 |url-status=dead |archiveurl=https://web.archive.org/web/20110726104027weblink |archivedate=2011-07-26}}.In contrast with Clark's methodology, modern classifications typically identify (or name) only those groupings that are monophyletic; that is, such a named group includes all the descendants of the group's common ancestor.Definitions of monophyly vary; for the one used here see e.g. {{Citation |last=Mishler |first=Brent D |year=2009 |editor-last=Ayala |editor-first=F.J. |editor2-last=Arp |editor2-first=R. |contribution=Species are not Uniquely Real Biological Entities |title=Contemporary Debates in Philosophy of Biology |pages=110–122 |isbn=978-1-4443-1492-2 |doi=10.1002/9781444314922.ch6 |lastauthoramp=yes}}.The cladogram below shows one possible classification sequence of the living primates:BOOK, Cartmill, M., Smith, F. H., The Human Lineage, John Wiley & Sons, 2011, 978-1-118-21145-8,weblink groups that use common (traditional) names are shown on the right.{{Barlabel|style=font-size:90%;|labelwidth=6|size=9
color1=purple|label1=prosimianscolor2=crimson|label2=monkeyscolor3=darkgreen|label3=great apescolor5=green|label5=lesser apescolor4=darkred|label4=humans|cladogram={{Cladex
|label1=Primatomorpha
|1={{Clade
| 1= Dermoptera (File:Cynocephalus volans Brehm1883 (white background).jpg|50 px)|label2= Primates |2={{Cladex
|label1= Haplorhini 
|1={{Cladex
|label1= Simiiformes 
|1={{Cladex
|label1= Catarrhini 
|1={{Cladex
|label1= Hominoidea 
|1={{Cladex
|label1= Hominidae 
|1={{Cladex
|label1= Homininae 
|1={{Cladex
|label1= Hominini 
|1={{Cladex
|1=humans (genus Homo) (File:Bechuana of Distinction-1841 (white background).jpg|60 px)|barbegin1=darkred|barend1=darkred
|2=chimpanzees (genus Pan)(File:PanTroglodytesSmit (white background).jpg|50 px)|barbegin2=darkgreen
}}
|label2= Gorillini 
|2={{Cladex
|1=gorillas (genus Gorilla) (File:Gorila de llanura occidental. Gorilla gorilla - Blanca Martí de Ahumada (white background).jpg|70 px)|bar1=darkgreen
}}
}}
|2=orangutans (subfamily Ponginae) (File:Simia satyrus - 1837 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - White Background.jpg|60 px)|barend2=darkgreen
}}
|2=gibbons (family Hylobatidae) (File:Le gibbon (white background).jpg|50 px)|barbegin2=green|barend2=green
}}
|2=Old World monkeys (superfamily Cercopithecoidea) (File:Cynocephalus doguera - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).tiff|60 px)|barbegin2=crimson
}}
|2=New World monkeys (parvorder Platyrrhini) (File:Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen (Plate 8) (white background).jpg|50 px)|barend2=crimson
}}
|label2= Tarsiiformes 
|2={{Cladex
|1=tarsiers (superfamily Tarsioidea) (File:Säugethiere vom Celebes- und Philippinen-Archipel (Taf. III) (white background) (1).jpg|25 px)|barbegin1=purple
}}
}}
|label2= Strepsirrhini 
|2={{Cladex
|label1=Lemuriformes{{efn|name=Lemuriformes}} 
|1={{Cladex
|1=lemurs (superfamily Lemuroidea) (File:FMIB 46849 Primates Maki Moccoe Lemur catta (white background).jpeg|35 px)|bar1=purple
|2=lorises and allies (superfamily Lorisoidea) (File:Nycticebus (white background).jpg|25 px)|barend2=purple
}}
}}
}}
}}}}}}All groups with scientific names are monophyletic (that is, they are clades), and the sequence of scientific classification reflects the evolutionary history of the related lineages. Groups that are traditionally named are shown on the right; they form an "ascending series" (per Clark, see above), and several groups are paraphyletic:
  • "prosimians" contain two monophyletic groups (the suborder Strepsirrhini, or lemurs, lorises and allies, as well as the tarsiers of the suborder Haplorhini); it is a paraphyletic grouping because it excludes the Simiiformes, which also are descendants of the common ancestor Primates.
  • "monkeys" comprise two monophyletic groups, New World monkeys and Old World monkeys, but is paraphyletic because it excludes hominoids, superfamily Hominoidea, also descendants of the common ancestor Simiiformes.
  • "apes" as a whole, and the "great apes" in particular, are paraphyletic because they exclude humans.
Thus, the members of the two sets of groups, and hence names, do not match, which causes problems in relating scientific names to common (usually traditional) names. Consider the superfamily Hominoidea: In terms of the common names on the right, this group consists of apes and humans and there is no single common name for all the members of the group. One remedy is to create a new common name, in this case "hominoids". Another possibility is to expand the use of one of the traditional names. For example, in his 2005 book, the vertebrate palaeontologist Benton wrote, "The apes, Hominoidea, today include the gibbons and orang-utan ... the gorilla and chimpanzee ... and humans";{{Sfn|Benton|2005|p=371}} thereby Benton was using "apes" to mean "hominoids". In that case, the group heretofore called "apes" must now be identified as the "non-human apes".{{As of|2005}}, there is no consensus as to whether to accept traditional (that is, common), but paraphyletic, names or to use monophyletic names only; or to use 'new' common names or adaptations of old ones. Both competing approaches can be found in biological sources, often in the same work, and sometimes by the same author. Thus, Benton defines "apes" to include humans, then he repeatedly uses "ape-like" to mean "like an ape rather than a human"; and when discussing the reaction of others to a new fossil he writes of "claims that Orrorin ... was an ape rather than a human".{{Sfn|Benton|2005|pp=378–380}}

Classification of living primates

File:Primates-drawing.jpg|thumb|upright|A 1927 drawing of chimpanzees, a gibbon (top right) and two (orangutan]]s (center and bottom center): The chimpanzee in the upper left is brachiating; the orangutan at the bottom center is knuckle-walking.)File:Akha cropped.png|right|upright|thumb|Homo sapiensHomo sapiensFile:NILGIRI LANGUR (Trachypithecus johnii).jpg|thumb|upright|Nilgiri langur (Trachypithecus johnii), an Old World monkeyOld World monkeyA list of the families of the living primates is given below, together with one possible classification into ranks between order and family.BOOK, Rylands, A. B., Mittermeier, R. A., yes, South American Primates: Comparative Perspectives in the Study of Behavior, Ecology, and Conservation, The Diversity of the New World Primates (Platyrrhini), Springer, Garber, P. A., Estrada, A., Bicca-Marques, J. C., Heymann, E. W., Strier, K. B., Karen B. Strier, 978-0-387-78704-6, 2009, Other classifications are also used. For example, an alternative classification of the living Strepsirrhini divides them into two infraorders, Lemuriformes and Lorisiformes.{{Sfn|Hartwig|2011|pp=20–21}}
  • Order Primates
    • Suborder Strepsirrhini: lemurs, galagos and lorisids
      • Infraorder Lemuriformes{{efn|name=Lemuriformes|Although the monophyletic relationship between lemurs and lorisoids is widely accepted, their clade name is not. The term "lemuriform" is used here because it derives from one popular taxonomy that clumps the clade of toothcombed primates into one infraorder and the extinct, non-toothcombed adapiforms into another, both within the suborder Strepsirrhini.{{Sfn|Szalay|Delson|1980|p=149}}{{Sfn|Cartmill|2010|p=15}} However, another popular alternative taxonomy places the lorisoids in their own infraorder, Lorisiformes.{{Sfn|Hartwig|2011|pp=20–21}}}}
Superfamily Lemuroidea* Family Cheirogaleidae: dwarf lemurs and mouse-lemurs (34 species)* Family Daubentoniidae: aye-aye (one species)* Family Lemuridae: ring-tailed lemur and allies (21 species)* Family Lepilemuridae: sportive lemurs (26 species)* Family Indriidae: woolly lemurs and allies (19 species) Superfamily Lorisoidea* Family Lorisidae: lorisids (14 species)* Family Galagidae: galagos (19 species) Family Tarsiidae: tarsiers (11 species) Parvorder Platyrrhini: New World monkeys* Family Callitrichidae: marmosets and tamarins (42 species)* Family Cebidae: capuchins and squirrel monkeys (14 species)* Family Aotidae: night or owl monkeys (douroucoulis) (11 species)* Family Pitheciidae: titis, sakis and uakaris (43 species)* Family Atelidae: howler, spider, woolly spider and woolly monkeys (29 species) Parvorder Catarrhini* Superfamily Cercopithecoidea** Family Cercopithecidae: Old World monkeys (138 species)* Superfamily Hominoidea** Family Hylobatidae: gibbons or "lesser apes" (18 species)** Family Hominidae: great apes, including humans (8 species)Order Primates was established by Carl Linnaeus in 1758, in the tenth edition of his book Systema Naturae,BOOK, Linnaeus, C., 1758, Sistema naturae per regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonimis locis. Tomus I, Impensis direct. Laurentii Salvii, Holmia, 20–32, for the genera Homo (humans), Simia (other apes and monkeys), Lemur (prosimians) and Vespertilio (bats). In the first edition of the same book (1735), he had used the name Anthropomorpha for Homo, Simia and Bradypus (sloths).BOOK, Linnaeus, C., 1735, Sistema naturae sive regna tria Naturae systematice proposita per classes, ordines, genera, & species, apud Theodorum Haak, Lugduni Batavorum, s.p, In 1839, Henri Marie Ducrotay de Blainville, following Linnaeus and imitating his nomenclature, established the orders Secundates (including the suborders Chiroptera, Insectivora and Carnivora), Tertiates (or Glires) and Quaternates (including Gravigrada, Pachydermata and Ruminantia),BOOK, Blainville, H., 1839, Nouvelle classification des Mammifères, Annales Françaises et Etrangères d'Anatomie et de Physiologie Appliquées à la Médicine et à l'Histoire Naturelle, 3, 268–269, but these new taxa were not accepted.Before Anderson and Jones introduced the classification of Strepsirrhini and Haplorhini in 1984,BOOK, Thorington, R. W., Anderson, S., yes, 1984, Primates, 187–217, Anderson, S., Jones, J. K., Orders and Families of Recent Mammals of the World, John Wiley and Sons, New York, 978-0-471-08493-8,weblink (followed by McKenna and Bell's 1997 work (Mammal classification#McKenna/Bell classification|Classification of Mammals: Above the species level)),BOOK, McKenna, M. C., Bell, S. K., yes, 1997, Classification of Mammals: Above the species level, Columbia University Press, New York, 631, 0-231-11013-8, the Primates were divided into two superfamilies: Prosimii and Anthropoidea.BOOK, Primate Behavioral Ecology, Third, Strier, K., Karen B. Strier, 2007, 978-0-205-44432-8, 50–53, Prosimii included all of the prosimians: Strepsirrhini plus the tarsiers. Anthropoidea contained all of the simians.

Phylogeny and genetics

{hide}cladogram|clades={{Clade|style=font-size:75%; width:35em;|label1=Euarchontoglires  |1={{Clade
|label1=Glires 
|1={{Clade
|1=Rodentia (rodents)
|2=Lagomorpha (rabbits, hares, pikas){edih}
|label2= Euarchonta 
|2={{Clade
|1=Scandentia (treeshrews)
| label2=Primatomorpha
|2={{Clade
|1=Dermoptera (colugos)
|label2=Primates
|2={{Clade
|state1=double
|1=†Plesiadapiformes
|2=crown primates}}
}}
}}
}}
}}}}Order Primates is part of the clade Euarchontoglires, which is nested within the clade Eutheria of Class Mammalia. Recent molecular genetic research on primates, colugos, and treeshrews has shown that the two species of colugos are more closely related to primates than to treeshrews,JOURNAL, Molecular and Genomic Data Identify the Closest Living Relative of Primates, Science, 2 November 2007, Janečka, J. E., Miller, W., Pringle, T. H., Wiens, F., Zitzmann, A., Helgen, K. M., Springer, M. S., Murphy, W. J., 10.1126/science.1147555, 318, 5851, 792–794, 17975064, 2007Sci...318..792J, even though treeshrews were at one time considered primates.BOOK, A Complete Guide to Monkeys, Apes and Other Primates, Kavanagh, M., Viking Press, New York, 18, 1983, 0-670-43543-0, These three orders make up the clade Euarchonta. The combination of this clade with the clade Glires (composed of Rodentia and Lagomorpha) forms the clade Euarchontoglires. Variously, both Euarchonta and Euarchontoglires are ranked as superorders. Some scientists consider Dermoptera to be a suborder of Primates and use the suborder Euprimates for the "true" primates.BOOK, McKenna, M. C., Bell, S. K., yes, 1997, Classification of Mammals Above the Species Level, Columbia University Press, New York, 329, 0-231-11012-X,

Evolution

{{Human timeline}}{{Further|Evolution of primates}}The primate lineage is thought to go back at least near the Cretaceous–Paleogene boundary or around 63–74 (mya),JOURNAL, Williams, B.A., Kay, R.F., Kirk, E.C., New perspectives on anthropoid origins, Proceedings of the National Academy of Sciences of the United States of America, 107, 4797–4804, 10.1073/pnas.0908320107, 11, 20212104, 2010, 2841917, 2010PNAS..107.4797W, JOURNAL, Stanyon, Roscoe, Springer, Mark S., Meredith, Robert W., Gatesy, John, Emerling, Christopher A., Park, Jong, Rabosky, Daniel L., Stadler, Tanja, Steiner, Cynthia, Ryder, Oliver A., Janečka, Jan E., Fisher, Colleen A., Murphy, William J., Macroevolutionary Dynamics and Historical Biogeography of Primate Diversification Inferred from a Species Supermatrix, PLOS ONE, 7, 11, 2012, e49521, 1932-6203, 10.1371/journal.pone.0049521, 23166696, 3500307, 2012PLoSO...749521S, JOURNAL, Jameson, Natalie M., Hou, Zhuo-Cheng, Sterner, Kirstin N., Weckle, Amy, Goodman, Morris, Steiper, Michael E., Wildman, Derek E., Genomic data reject the hypothesis of a prosimian primate clade, Journal of Human Evolution, 61, 3, September 2011, 295–305, 0047-2484, 10.1016/j.jhevol.2011.04.004, 21620437, JOURNAL, Pozzi, Luca, Hodgson, Jason A., Burrell, Andrew S., Sterner, Kirstin N., Raaum, Ryan L., Disotell, Todd R., Primate phylogenetic relationships and divergence dates inferred from complete mitochondrial genomes, Molecular Phylogenetics and Evolution, 75, June 2014, 165–183, 1055-7903, 10.1016/j.ympev.2014.02.023, 24583291, 4059600, JOURNAL, Stanyon, Roscoe, Finstermeier, Knut, Zinner, Dietmar, Brameier, Markus, Meyer, Matthias, Kreuz, Eva, Hofreiter, Michael, Roos, Christian, A Mitogenomic Phylogeny of Living Primates, PLOS ONE, 8, 7, 16 July 2013, e69504, 1932-6203, 10.1371/journal.pone.0069504, 23874967, 3713065, 2013PLoSO...869504F, even though the oldest known primates from the fossil record date to the Late Paleocene of Africa (Altiatlasius)JOURNAL, Williams, B. A., Kay, R. F., Kirk, E. C., New perspectives on anthropoid origins, 10.1073/pnas.0908320107, Proceedings of the National Academy of Sciences of the United States of America, 107, 11, 4797–4804, 2010, 20212104, 2841917, harv, 2010PNAS..107.4797W, or the Paleocene-Eocene transition in the northern continents, c. 55 mya (Cantius, Donrussellia, Altanius, Plesiadapis and Teilhardina).JOURNAL, Miller, E. R., Gunnell, G. F., Martin, R. D., Deep Time and the Search for Anthropoid Origins, 10.1002/ajpa.20352, American Journal of Physical Anthropology, 128, 60–95, 2005, 16369958,weblink harv, JOURNAL, 27 October 2009, Helen J, Chatterjee, Simon Y.W., Ho, Ian, Barnes, Colin, Groves, Colin Groves, Estimating the phylogeny and divergence times of primates using a supermatrix approach, BMC Evolutionary Biology, 9, 10.1186/1471-2148-9-259, 19860891, 259, 2774700, 1, JOURNAL, O'Leary, M. A., 8 February 2013, The placental mammal ancestor and the post–K-Pg radiation of placentals, Science, 339, 6120, 662–667, 10.1126/science.1229237, 23393258, etal, 2013Sci...339..662O, 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.JOURNAL, Molecular Clock Calibrations and Metazoan Divergence Dates, Lee, M., Journal of Molecular Evolution, 49, 3, September 1999, 385–391, 10.1007/PL00006562, 10473780, 1999JMolE..49..385L, WEB, Scientists Push Back Primate Origins From 65 Million To 85 Million Years Ago,weblink Science Daily, 2008-10-24, JOURNAL, Using the fossil record to estimate the age of the last common ancestor of extant primates, Tavaré, S., Marshall, C. R., Will, O., Soligo, C., Martin R.D., Nature, April 18, 2002, 726–729, 416, 6882, 10.1038/416726a, 11961552, 2002Natur.416..726T, By modern cladistic reckoning, the order Primates is monophyletic. The suborder Strepsirrhini, the "wet-nosed" primates, is generally thought to have split off from the primitive primate line about 63 mya,JOURNAL, Molecular Remodeling of Members of the Relaxin Family During Primate Evolution, Molecular Biology and Evolution, 18, 393–403,weblink Klonisch, T., Froehlich, C., Tetens, F., Fischer, B., Hombach-Klonisch, S., 2001, 2008-08-22, 11230540, 3, 10.1093/oxfordjournals.molbev.a003815, although earlier dates are also supported.JOURNAL, Development and Application of a Phylogenomic Toolkit: Resolving the Evolutionary History of Madagascar's Lemurs, Genome Research, 18, 489–499, 2008, Horvath, J., 10.1101/gr.7265208, 18245770, 3, 2259113, 1, Weisrock, D. W., Embry, S. L., Fiorentino, I., Balhoff, J. P., Kappeler, P., Wray, G. A., Willard, H. F., Yoder, A. D., The seven strepsirrhine families are the five related lemur families and the two remaining families that include the lorisids and the galagos.JOURNAL, Lemur Diversity in Madagascar, Russell Mittermeier, Mittermeier, R., Ganzhorn, J., Konstant, W., Glander, K., Tattersall, I., Groves, C., Rylands, A., Hapke, A., Ratsimbazafy, J., Mayor, M., Louis, E., Rumpler, Y., Schwitzer, C., Rasoloarison, R., International Journal of Primatology, 10.1007/s10764-008-9317-y, 1607–1656, 29, 6, December 2008, Older classification schemes wrap Lepilemuridae into Lemuridae and Galagidae into Lorisidae, yielding a four-one family distribution instead of five-two as presented here. During the Eocene, most of the northern continents were dominated by two groups, the adapiforms and the omomyids.BOOK, Primates in Perspective, Hartwig, W., Primate Evolution, Campbell, C., Fuentes, A., MacKinnon, K., Panger, M., Bearder, S., 2007, Oxford University Press, 978-0-19-517133-4, 13–17, The former are considered members of Strepsirrhini, but did not have a toothcomb like modern lemurs; recent analysis has demonstrated that Darwinius masillae fits into this grouping.JOURNAL, Williams, B. A., Kay, R. F., Christopher Kirk, E., Ross, C. F., Darwinius masillae is a strepsirrhine—a reply to Franzen et al. (2009), 10.1016/j.jhevol.2010.01.003, Journal of Human Evolution, 59, 5, 567–573; discussion 573–9, 2010, 20188396,weblinkweblink" title="web.archive.org/web/20130517155919weblink">weblink 2013-05-17, harv, The latter was closely related to tarsiers, monkeys, and apes. How these two groups relate to extant primates is unclear. Omomyids perished about 30 mya, while adapiforms survived until about 10 mya.BOOK, Primate Evolution and Human Origins, Ciochon, R., Fleagle, J., yes, Menlo Park, California, Benjamin/Cummings, 1987, 978-0-202-01175-2, 72, According to genetic studies, the lemurs of Madagascar diverged from the lorisoids approximately 75 mya. 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.BOOK, Garbutt, N., Mammals of Madagascar, A Complete Guide, A&C Black Publishers, 2007, 978-0-300-12550-4, 85–86, However, Madagascar split from Africa 160 mya and from India 90 mya.BOOK, Mittermeier, R.A., Russell Mittermeier, Lemurs of Madagascar, 2nd, Conservation International, 2006, 1-881173-88-7, 23–26, etal, 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. Other colonization options have been suggested, such as multiple colonizations from Africa and India,WEB,weblink Primate Evolution, 2008-10-23, Sellers, Bill, 2000-10-20, University of Edinburgh, 13–17, dead,weblink" title="web.archive.org/web/20081029184647weblink">weblink 2008-10-29, but none are supported by the genetic and molecular evidence.File:Brown Lemur in Andasibe.jpg|right|thumb|Common brown lemur, a Strepsirrhine primate]]Until recently, the aye-aye has been difficult to place within Strepsirrhini. 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.Suborder Haplorhini, the simple-nosed or "dry-nosed" primates, is composed of two sister clades. Prosimian tarsiers in the family Tarsiidae (monotypic in its own infraorder Tarsiiformes), represent the most basal division, originating about 58 mya.CONFERENCE, Evolutionary Biology of Tarsiers, Shekelle, M., 2005,weblink 2008-08-22, dead,weblink" title="web.archive.org/web/20080907073204weblink">weblink 2008-09-07, JOURNAL, Rapid electrostatic evolution at the binding site for cytochrome c on cytochrome c oxidase in anthropoid primates, Schmidt, T., Proceedings of the National Academy of Sciences of the United States of America, 102, 18, 6379–6384, 10.1073/pnas.0409714102, 3 May 2005, 15851671, 1088365, 1, Wildman, DE, Uddin, M, Opazo, JC, Goodman, M, Grossman, LI, 2005PNAS..102.6379S, The earliest known haplorhine skeleton, that of 55 MA old tarsier-like Archicebus, was found in central China,WEB, Wade, Lizzie, Early Primate Weighed Less Than an Ounce,weblink ScienceNow, June 5, 2013, 2013-06-07, dead,weblink 2013-06-08, supporting an already suspected Asian origin for the group.JOURNAL, Kay, R. F., Evidence for an Asian origin of stem anthropoid s, Proceedings of the National Academy of Sciences of the United States of America, 109, 26, 2012, 10132–10133, 10.1073/pnas.1207933109, 2012PNAS..10910132K, 22699505, 3387095, The infraorder Simiiformes (simian primates, consisting of monkeys and apes) emerged about 40 mya, possibly also in Asia; if so, they dispersed across the Tethys Sea from Asia to Africa soon afterwards.JOURNAL, Chaimanee, Y., Chavasseau, O., Beard, K. C., Kyaw, A. A., Soe, A. N., Sein, C., Lazzari, V., Marivaux, L., Marandat, B., Swe, M., Rugbumrung, M., Lwin, T., Valentin, X., Zin-Maung-Maung-Thein, Jaeger, J. -J., Late Middle Eocene primate from Myanmar and the initial anthropoid colonization of Africa, 10.1073/pnas.1200644109, Proceedings of the National Academy of Sciences of the United States of America, 109, 26, 10293–10297, 2012, 22665790, 3387043, harv, 2012PNAS..10910293C, There are two simian clades, both (wikt:parvorder|parvorder)s: Catarrhini, which developed in Africa, consisting of Old World monkeys, humans and the other apes, and Platyrrhini, which developed in South America, consisting of New World monkeys. A third clade, which included the eosimiids, developed in Asia, but became extinct millions of years ago.JOURNAL, Marivaux, L., Anthropoid primates from the Oligocene of Pakistan (Bugti Hills): Data on early anthropoid evolution and biogeography, Proceedings of the National Academy of Sciences of the United States of America, 2005-06-14, 102, 24, 8436–8441, 10.1073/pnas.0503469102, 15937103, 1150860, 1, Antoine, PO, Baqri, SR, Benammi, M, Chaimanee, Y, Crochet, JY, De Franceschi, D, Iqbal, N, Jaeger, JJ, 2005PNAS..102.8436M, 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.JOURNAL, Schrago, C.G., Russo, C.A.M., yes, Timing the Origin of New World Monkeys, Molecular Biology and Evolution, 20, 10, 2003, 1620–1625,weblink PDF Reprint, 10.1093/molbev/msg172, 12832653, The anthropoid primates possibly traversed the Atlantic Ocean from Africa to South America during the Eocene by island hopping, facilitated by Atlantic Ocean ridges and a lowered sea level. Alternatively, a single 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. Research suggests that a small {{convert|1|kg|lb|abbr=on}} primate could have survived 13 days on a raft of vegetation.JOURNAL, Houle, A., The origin of platyrrhines: An evaluation of the Antarctic scenario and the floating island model, American Journal of Physical Anthropology, 109, 4, 1999, 541–559, 10.1002/(SICI)1096-8644(199908)109:43.0.CO;2-N, 10423268, 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 monkeyNew World monkeyApes and monkeys spread from Africa into Europe and Asia starting in the Miocene.JOURNAL, Andrews, P., Kelley, J., yes, Middle Miocene Dispersals of Apes, Folia Primatologica, 78, 2007, 328–343, 10.1159/000105148, 17855786, 5–6, 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. Old World monkeys disappeared from Europe about 1.8 mya.BOOK, Primate Behavioral Ecology, Strier, K., Allyn & Bacon, 3rd, 2007, 978-0-205-44432-8, 7, 64, 71, 77, 182–185, 273–280, 284, 287–298, Molecular and fossil studies generally show that modern humans originated in Africa 100,000–200,000 years ago.BOOK, Pough, F. W., Janis, C. M., Heiser, J. B., Vertebrate Life, Primate Evolution and the Emergence of Humans, 2005, 1979, 7th, Pearson, 650, 0-13-127836-3, Although primates are well studied in comparison to other animal groups, several new species have been discovered recently, and genetic tests have revealed previously unrecognised species in known populations. Primate Taxonomy listed about 350 species of primates in 2001;BOOK, Primate Taxonomy, Smithsonian Institution Press, 2001, Groves, C. P., 1-56098-872-X, the author, Colin Groves, increased that number to 376 for his contribution to the third edition of Mammal Species of the World (MSW3). However, publications since the taxonomy in MSW3 was compiled in 2003 have pushed the number to 424 species, or 658 including subspecies.

Hybrids

Primate hybrids usually arise in captivity,JOURNAL, Songs of hybrid gibbons (Hylobates lar × H. muelleri), Tenaza, R., 1984, American Journal of Primatology, 8, 3, 249–253, 10.1002/ajp.1350080307, but there have also been examples in the wild.JOURNAL, 1966, Naturally occurring primate hybrid, Bernsteil, I. S., Science, 154, 3756, 1559–1560, 10.1126/science.154.3756.1559, 4958933, 1966Sci...154.1559B, JOURNAL, Sugawara, K., Sociological study of a wild group of hybrid baboons between Papio anubis and P. hamadryas in the Awash Valley, Ethiopia, January 1979, Primates, 20, 1, 10.1007/BF02373827, 21–56, Hybridization occurs where two species' range overlap to form hybrid zones; hybrids may be created by humans when animals are placed in zoos or due to environmental pressures such as predation. Intergeneric hybridizations, hybrids of different genera, have also been found in the wild. Although they belong to genera that have been distinct for several million years, interbreeding still occurs between the gelada and the hamadryas baboon.JOURNAL, Jolly, Intergeneric Hybrid Baboons, International Journal of Primatology, 1997, 18, 4, 10.1023/A:1026367307470, 597–627, C. J., 2, Woolley-Barker, Tamsin, Beyene, Shimelis, Disotell, Todd R., Phillips-Conroy, Jane E.,

Clones

On 24 January 2018, scientists in China reported in the journal Cell the creation of two crab-eating macaque clones, named Zhong Zhong and Hua Hua, using the complex DNA transfer method that produced Dolly the sheep, for the first time.JOURNAL, Liu, Zhen, etal, Cloning of Macaque Monkeys by Somatic Cell Nuclear Transfer,weblink 24 January 2018, Cell (journal), Cell, 10.1016/j.cell.2018.01.020, 29395327, 24 January 2018, 172, 4, 881–887.e7, JOURNAL, Normile, Dennis, These monkey twins are the first primate clones made by the method that developed Dolly,weblink 24 January 2018, Science (journal), Science, 10.1126/science.aat1066, 24 January 2018, JOURNAL, Cyranoski, David, First monkeys cloned with technique that made Dolly the sheep - Chinese scientists create cloned primates that could revolutionize studies of human disease., 24 January 2018, Nature (journal), Nature, 553, 7689, 387–388, 10.1038/d41586-018-01027-z, 29368720, 2018Natur.553..387C, NEWS, Briggs, Helen, First monkey clones created in Chinese laboratory,weblink 24 January 2018, BBC News, 24 January 2018, NEWS, Associated Press, Scientists Successfully Clone Monkeys; Are Humans Up Next?,weblink 24 January 2018, The New York Times, 24 January 2018,

Anatomy and physiology

Head

File:Primate skull series with legend cropped.png|thumb|Primate skulls showing postorbital barpostorbital barThe primate skull has a large, domed cranium, which is particularly prominent in anthropoids. The cranium protects the large brain, a distinguishing characteristic of this group. The endocranial volume (the volume within the skull) is three times greater in humans than in the greatest nonhuman primate, reflecting a larger brain size. The mean endocranial volume is 1,201 cubic centimeters in humans, 469 cm3 in gorillas, 400 cm3 in chimpanzees and 397 cm3 in orangutans.BOOK, Aiello, L., Dean, C., yes, 1990, An Introduction to Human Evolutionary Anatomy, Academic Press, 193, 0-12-045590-0, The primary evolutionary trend of primates has been the elaboration of the brain, in particular the neocortex (a part of the cerebral cortex), which is involved with sensory perception, generation of motor commands, spatial reasoning, conscious thought and, in humans, language. While other mammals rely heavily on their sense of smell, the arboreal life of primates has led to a tactile, visually dominant sensory system, a reduction in the olfactory region of the brain and increasingly complex social behavior.WEB,weblink "Primates" (On-line), Animal Diversity Web, Myers, P., 1999, 2008-06-03, Primates have forward-facing eyes on the front of the skull; binocular vision allows accurate distance perception, useful for the brachiating ancestors of all great apes. A bony ridge above the eye sockets reinforces weaker bones in the face, which are put under strain during chewing. Strepsirrhines have a postorbital bar, a bone around the eye socket, to protect their eyes; in contrast, the higher primates, haplorhines, have evolved fully enclosed sockets.BOOK, Campbell, B. G., Loy, J. D., yes, 2000, Humankind Emerging (8th edition), Allyn & Bacon, 85, 0-673-52364-0, (File:PrimateFeet.jpg|thumb|right|upright|An 1893 drawing of the hands and feet of various primates)Primates show an evolutionary trend towards a reduced snout.WEB, White, T., Kazlev, A., yes,weblink Archonta: Primates, Palaeos, 2008-06-03, 2006-01-08,weblink" title="web.archive.org/web/20080512022901weblink">weblink 2008-05-12, Technically, Old World monkeys are distinguished from New World monkeys by the structure of the nose, and from apes by the arrangement of their teeth. In New World monkeys, the nostrils face sideways; in Old World monkeys, they face downwards. Dental pattern in primates vary considerably; although some have lost most of their incisors, all retain at least one lower incisor. In most strepsirrhines, the lower incisors form a toothcomb, which is used in grooming and sometimes foraging. Old World monkeys have eight premolars, compared with 12 in New World monkeys. The Old World species are divided into apes and monkeys depending on the number of cusps on their molars: monkeys have four, apes have five - although humans may have four or five.BOOK, Wheeler's Dental Anatomy, Physiology, and Occlusion, Ash, M. M., Nelson, S. J., Wheeler, R. C., W.B. Saunders, 2003, 978-0-7216-9382-8, 12,weblink The main hominid molar cusp (hypocone) evolved in early primate history, while the cusp of the corresponding primitive lower molar (paraconid) was lost. Prosimians are distinguished by their immobilized upper lips, the moist tip of their noses and forward-facing lower front teeth.

Body

(File:Chlorocebus pygerythrus01.jpg|thumb|Vervet hindfoot showing fingerprint ridges on the sole)Primates generally have five digits on each limb (pentadactyly), with keratin nails on the end of each finger and toe. The bottom sides of the hands and feet have sensitive pads on the fingertips. Most have opposable thumbs, a characteristic primate feature most developed in humans, though not limited to this order, (opossums and koalas, for example, also have them). Thumbs allow some species to use tools. In primates, the combination of opposing thumbs, short fingernails (rather than claws) and long, inward-closing fingers is a relict of the ancestral practice of gripping branches, and has, in part, allowed some species to develop brachiation (swinging by the arms from tree limb to tree limb) as a significant means of locomotion. Prosimians have clawlike nails on the second toe of each foot, called toilet-claws, which they use for grooming.BOOK, Pough, F. W., Janis, C. M., Heiser, J. B., Vertebrate Life, Characteristics of Primates, 2005, 1979, 7th, Pearson, 630, 0-13-127836-3, The primate collar bone is a prominent element of the pectoral girdle; this allows the shoulder joint broad mobility. Compared to Old World monkeys, apes have more mobile shoulder joints and arms due to the dorsal position of the scapula, broad ribcages that are flatter front-to-back, a shorter, less mobile spine, and with lower vertebrae greatly reduced - resulting in tail loss in some species. Prehensile tails are found in atelids, including the howler, spider, woolly spider, woolly monkeys; and in capuchins.JOURNAL, Garber PA, Rehg JA, The ecological role of the prehensile tail in white-faced capuchins (Cebus capucinus), American Journal of Physical Anthropology, 110, 3, 325–39, November 1999, 10516564, 10.1002/(SICI)1096-8644(199911)110:33.0.CO;2-D, JOURNAL, Russo GA, Young JW, Tail growth tracks the ontogeny of prehensile tail use in capuchin monkeys (Cebus albifrons and C. apella), American Journal of Physical Anthropology, 146, 3, 465–73, November 2011, 21953012, 10.1002/ajpa.21617, Male primates have a pendulous penis and scrotal testes.BOOK, Friderun Ankel-Simons, Primate Anatomy: An Introduction,weblink 27 July 2010, Academic Press, 978-0-08-046911-9, ENCYCLOPEDIA, Macdonald, David, David W. Macdonald, Primates, The Encyclopedia of Mammals, 282–307, The Brown Reference Group plc, 2006, 0-681-45659-0,

Sexual dimorphism

File:Hamadryas Baboon.jpg|thumb|right|Distinct sexual size dimorphism can be seen between the female and two male hamadryas baboonhamadryas baboonSexual dimorphism is often exhibited in simians, though to a greater degree in Old World species (apes and some monkeys) than New World species. Recent studies involve comparing DNA to examine both the variation in the expression of the dimorphism among primates and the fundamental causes of sexual dimorphism. Primates usually have dimorphism in body massJOURNAL, Lindenfors, Patrik, Tullberg, Birgitta S., 1998, Phylogenetic analyses of primate size evolution: the consequences of sexual selection, Biological Journal of the Linnean Society, en, 64, 4, 413–447, 10.1111/j.1095-8312.1998.tb00342.x, 0024-4066, JOURNAL, Ralls, K., 1976, Mammals in Which Females are Larger Than Males, The Quarterly Review of Biology, 51, 2, 10.1086/409310, 245–76, 785524, JOURNAL, Lindstedtand & Boyce, July 1985, The American Naturalist, 125, 873, 10.1086/284385, Seasonality, Fasting Endurance, and Body Size in Mammals, Boyce, Mark S., 6, and canine tooth sizeJOURNAL, Thorén, Sandra, Lindenfors, Patrik, Kappeler, Peter M., 2006, Phylogenetic analyses of dimorphism in primates: Evidence for stronger selection on canine size than on body size, American Journal of Physical Anthropology, en, 130, 1, 50–59, 10.1002/ajpa.20321, 16345072, 0002-9483, JOURNAL, Frisch, J. E., 1963, Sex-differences in the canines of the gibbon (Hylobates lar), Primates, 4, 2, 10.1007/BF01659148, 1–10, JOURNAL, Kay, R. F., 1975, The functional adaptations of primate molar teeth, American Journal of Physical Anthropology, 43, 2, 195–215, 10.1002/ajpa.1330430207, 810034, along with pelage and skin color.BOOK, Crook, J. H., Campbell, B. G., 1972, Sexual selection, dimorphism, and social organization in the primates, Sexual selection and the descent of man, Aldine Transaction, 246, 978-0-202-02005-1,weblink The dimorphism can be attributed to and affected by different factors, including mating system,JOURNAL, Cheverud, J. M., Dow, M. M., Leutenegger, W., November 1985, The quantitative assessment of phylogenetic constraints in comparative analyses: Sexual dimorphism in body weight among primates, Evolution, 39, 6, 1335–1351, 10.2307/2408790, 28564267, 2408790, size, habitat and diet.JOURNAL, Leutenegger, W., Cheverud, J. M., 1982, Correlates of sexual dimorphism in primates: Ecological and size variables, International Journal of Primatology, 3, 4, 10.1007/BF02693740, 387–402, Comparative analyses have generated a more complete understanding of the relationship between sexual selection, natural selection, and mating systems in primates. Studies have shown that dimorphism is the product of changes in both male and female traits.JOURNAL, Plavcan, J. M., 2001, Sexual dimorphism in primate evolution, American Journal of Physical Anthropology, 33, 25–53, 10.1002/ajpa.10011, 11786990, Ontogenetic scaling, where relative extension of a common growth trajectory occurs, may give some insight into the relationship between sexual dimorphism and growth patterns.JOURNAL, O'Higgins, P., Collard, M., 2002, Sexual dimorphism and facial growth in papionine monkeys, Journal of Zoology, 257, 2, 255–72, 10.1017/S0952836902000857, Some evidence from the fossil record suggests that there was convergent evolution of dimorphism, and some extinct hominids probably had greater dimorphism than any living primate.

Locomotion

File:Diademed ready to push off.jpg|thumb|Diademed sifakaDiademed sifakaPrimate species move by brachiation, bipedalism, leaping, arboreal and terrestrial quadrupedalism, climbing, knuckle-walking or by a combination of these methods. Several prosimians are primarily vertical clingers and leapers. These include many bushbabies, all indriids (i.e., sifakas, avahis and indris), sportive lemurs, and all tarsiers.BOOK, Primate Ecology and Social Structure Volume 1: Lorises, Lemurs and Tarsiers, Sussman, R. W., 1999, 0-536-02256-9, Needham Heights, MA, Pearson Custom Publishing & Prentice Hall, 78, 89–90, 108, 121–123, 233, Other prosimians are arboreal quadrupeds and climbers. Some are also terrestrial quadrupeds, while some are leapers. Most monkeys are both arboreal and terrestrial quadrupeds and climbers. Gibbons, muriquis and spider monkeys all brachiate extensively, with gibbons sometimes doing so in remarkably acrobatic fashion. Woolly monkeys also brachiate at times. Orangutans use a similar form of locomotion called quadramanous climbing, in which they use their arms and legs to carry their heavy bodies through the trees. Chimpanzees and gorillas knuckle walk, and can move bipedally for short distances. Although numerous species, such as australopithecines and early hominids, have exhibited fully bipedal locomotion, humans are the only extant species with this trait.BOOK, S. D., Glazier, C. A., Flowerday, Selected Readings in the Anthropology of Religion: Theoretical and Methodological Essays, Greenwood Publishing Group, 2003, 9780313300905, 53,

Vision

missing image!
- GarnettsGalago CincinnatiZoo.jpg -
The tapetum lucidum of a northern greater galago, typical of prosimians, reflects the light of the photographers flash
The evolution of color vision in primates is unique among most eutherian mammals. While the remote vertebrate ancestors of the primates possessed three color vision (trichromaticism), the nocturnal, warm-blooded, mammalian ancestors lost one of three cones in the retina during the Mesozoic era. Fish, reptiles and birds are therefore trichromatic or tetrachromatic, while all mammals, with the exception of some primates and marsupials,JOURNAL, Arrese, 2005, Cone topography and spectral sensitivity in two potentially trichromatic marsupials, the quokka (Setonix brachyurus) and quenda (Isoodon obesulus), Proceedings of the Royal Society B, 272, 1565, 10.1098/rspb.2004.3009, 791–6, 15888411, 1599861, C. A., Oddy, Alison Y., 2, Runham, P. B, Hart, N. S, Shand, J., Hunt, D. M, Beazley, L. D, are dichromats or monochromats (totally color blind). Nocturnal primates, such as the night monkeys and bush babies, are often monochromatic. Catarrhines are routinely trichromatic due to a gene duplication of the red-green opsin gene at the base of their lineage, 30 to 40 million years ago.JOURNAL, Bowmaker, J. K., Astell, S., Hunt, D. M., Mollon, J. D., 1991, Photosensitive and photostable pigments in the retinae of Old World monkeys, The Journal of Experimental Biology, 156, 1–19, 0022-0949,weblink 2008-06-16, 2051127, 1, Platyrrhines, on the other hand, are trichromatic in a few cases only.JOURNAL, Surridge, A. K., D. Osorio, yes, 2003, Evolution and selection of trichromatic vision in primates, Trends in Ecology and Evolution, 18, 198–205, 10.1016/S0169-5347(03)00012-0, 4, Specifically, individual females must be heterozygous for two alleles of the opsin gene (red and green) located on the same locus of the X chromosome. Males, therefore, can only be dichromatic, while females can be either dichromatic or trichromatic. Color vision in strepsirrhines is not as well understood; however, research indicates a range of color vision similar to that found in platyrrhines.Like catarrhines, howler monkeys (a family of platyrrhines) show routine trichromatism that has been traced to an evolutionarily recent gene duplication.JOURNAL, Lucas, P. W., Dominy, N. J., Riba-Hernandez, P., Stoner, K. E., Yamashita, N., Loría-Calderón, E., Petersen-Pereira, W., Rojas-Durán, Y., Salas-Pena, R., Solis-Madrigal, S., Osorio, D., Darvell, B. W., 2003, Evolution and function of routine trichromatic vision in primates, Evolution, 57, 2636–43, 10.1554/03-168, 14686538, 11, Howler monkeys are one of the most specialized leaf-eaters of the New World monkeys; fruits are not a major part of their diets,BOOK, Primate Ecology and Social Structure, Volume 2: New World Monkeys, Revised First, Sussman, R. W., 2003, 0-536-74364-9, Needham Heights, MA, Pearson Custom Publishing & Prentice Hall, 77–80, 132–133, 141–143, and the type of leaves they prefer to consume (young, nutritive, and digestible) are detectable only by a red-green signal. Field work exploring the dietary preferences of howler monkeys suggests that routine trichromaticism was selected by environment.

Behavior

Social systems

Richard Wrangham stated that social systems of non-human primates are best classified by the amount of movement by females occurring between groups.BOOK, Wrangham, R. W., Richard Wrangham, Mutualism, kinship and social evolution, 1982, Current Problems in Sociobiology, Cambridge University Press, 269–89, 0-521-24203-7, He proposed four categories:
  • Female transfer systems â€“ 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. This organization can be seen in chimpanzees, where the males, who are typically related, will cooperate in defense of the group's territory. Among New World Monkeys, spider monkeys and muriquis use this system.BOOK, Primates in Perspective, Fiore, A. D., Campbell, C. J., yes, The Atelines, 2007, 175, Oxford University Press, Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M., Bearder, S. K., 978-0-19-517133-4,
File:Ringstaartmakis - Ring-tailed Lemur.jpg|thumb|right|A social huddle of ring-tailed lemurring-tailed lemur
  • Male transfer systems â€“ while the females remain in their natal groups, the males will emigrate as adolescents. Polygynous and multi-male societies are classed in this category. Group sizes are usually larger. This system is common among the ring-tailed lemur, capuchin monkeys and cercopithecine monkeys.
  • Monogamous species â€“ 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. Gibbons essentially use this system, although "monogamy" in this context does not necessarily mean absolute sexual fidelity.BOOK, Primates in Perspective, Bartlett, T. Q., The Hylobatidae, 2007, Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M., Bearder, S. K., Oxford University Press, 978-0-19-517133-4, 283, These species do not live in larger groups.
  • Solitary species â€“ often males who defend territories that include the home ranges of several females. This type of organization is found in the prosimians such as the slow loris. Orangutans do not defend their territory but effectively have this organization.BOOK, Primates in Perspective, Knott, C. D., Kahlenberg, S. M., yes, Orangutans in Perspective, 2007, Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M., Bearder, S. K., Oxford University Press, 978-0-19-517133-4, 294,
Other systems are known to occur as well. For example, with howler monkeys 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. Some prosimians, colobine monkeys and callitrichid monkeys use this system.The transfer of females or males from their native group is likely an adaptation for avoiding inbreeding.JOURNAL, Charpentier MJ, Widdig A, Alberts SC, Inbreeding depression in non-human primates: a historical review of methods used and empirical data, American Journal of Primatology, 69, 12, 1370–86, December 2007, 17486606, 10.1002/ajp.20445, 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.JOURNAL, Ralls K, Ballou J, 1982, Effect of inbreeding on infant mortality in captive primates, 10.1007/BF02693747, International Journal of Primatology, 3, 4, 491–505,weblink 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|The chimpanzees are social great apes.]]Primatologist Jane Goodall, who studied in the Gombe Stream National Park, noted fission-fusion societies in chimpanzees.JOURNAL, Constable, J. L., Ashley, M. V., Goodall, J., Pusey, A. E., Noninvasive paternity assignment in Gombe chimpanzees, Molecular Ecology, 10, 5, 1279–300, May 2001, 11380884, 10.1046/j.1365-294X.2001.01262.x, 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,BOOK, The Pictorial Guide to the Living Primates, Rowe, N., 1996, Pogonias Press, 0-9648825-0-7, 4, 139, 143, 154, 185, 223, spider monkeys and the bonobo. The gelada has a similar social structure in which many smaller groups come together to form temporary herds of up to 600 monkeys.These social systems are affected by three main ecological factors: distribution of resources, group size, and predation.BOOK, Pough, F. W., Janis, C. M., Heiser, J. B., Vertebrate Life, Primate Societies, 2005, 1979, 7th, Pearson, 621–623, 0-13-127836-3, Within a social group there is a balance between cooperation and competition. Cooperative behaviors include social grooming (removing 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 hierarchies.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.

Interspecific associations

Several species of primates are known to associate in the wild. Some of these associations have been extensively studied. In the 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 and sooty mangabey, which coordinate anti-predator alarm calls.BOOK, Monkeys of Tai Forest, An African Primate Community, Interactions between African Crowned Eagles and Their Prey Community, Shultz, S., Thomsett, S., yes, McGraw, W., Zuberbuhler, K., Noe, R., 2007, Cambridge University Press, 978-0-521-81633-5, 181, Among the predators of these monkeys is the common chimpanzee.BOOK, Monkeys of Tai Forest, An African Primate Community, Interactions between Red Colobus Monkeys and Chimpanzees, Bshary, R., McGraw, W., Zuberbuhler, K., Noe, R., 2007, Cambridge University Press, 978-0-521-81633-5, 155–170, 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. Several of these species are preyed upon by the common chimpanzee.BOOK, Chimpanzee and Red Colobus : the ecology of predator and prey, Stanford, C., 1998, Harvard University Press, 0-674-00722-0, 130–138, 233, In South America, squirrel monkeys associate with capuchin monkeys.BOOK, On the Move : how and why animals travel in groups, Social Manipulation Within and Between Troops Mediates Primate Group Movement, Boinski, S., Boinski, S., Garber, P., 2000, University of Chicago Press, 0-226-06340-2, 447–448, This may have more to do with foraging benefits to the squirrel monkeys than anti-predation benefits.

Cognition and communication

{{Further|Great ape language}}File:howler monkey.jpg|thumb|Pair of black howler monkeys vocalizing]]Primates have advanced cognitive abilities: some make tools and use them to acquire food and for social displays;JOURNAL, Boesch, C., Boesch, H., Tool Use and Tool Making in Wild Chimpanzees, Folia Primatologica, 1990, 54, 86–99, 10.1159/000156428, 2157651, 1–2, JOURNAL, Westergaard, 1998, Why some capuchin monkeys (Cebus apella) use probing tools (and others do not), Journal of Comparative Psychology, 112, 2, 207–211, 10.1037/0735-7036.112.2.207, 9642788, G. C., Lundquist, A. L., 2, Haynie, Michael Katherine, Kuhn, Heather E., Suomi, Stephen J., some can perform tasks requiring cooperation, influence and rank;JOURNAL, de Waal, F. B. M., Davis, J. M., 2003, Capuchin cognitive ecology: cooperation based on projected returns, Neuropsychologia, 41, 221–228, 10.1016/S0028-3932(02)00152-5, 12459220, 2, they are status conscious, manipulative and capable of deception;JOURNAL, Paar, L. A., Winslow, J. T., Hopkins, W. D., de Waal, F. B. M., 2000, Recognizing facial cues: Individual discrimination by chimpanzees (Pan troglodytes) and rhesus monkeys (Macaca mulatta), Journal of Comparative Psychology, 114, 1, 47–60, 10.1037/0735-7036.114.1.47, 10739311, 2018744, JOURNAL, Byrne, Richard, Corp, Nadia, Neocortex size predicts deception rate in primates, Proceedings of the Royal Society of London. Series B: Biological Sciences, 2004, 271, 1549, 1693–1699, 10.1098/rspb.2004.2780, 15306289, 1691785, they can recognise kin and conspecifics;JOURNAL, Paar, L. A., de Waal, F. B. M., 1999, Visual kin recognition in chimpanzees, Nature, 399, 647–648, 10.1038/21345, 10385114, 6737, 1999Natur.399..647P, JOURNAL, 1997, Discrimination of macaques by macaques: The case of sulawesi species, Primates, 38, 3, 233–245, Fujita, K., Watanabe, K., Widarto, T. H., Suryobroto, B., 10.1007/BF02381612, and they can learn to use symbols and understand aspects of human language including some relational syntax and concepts of number and numerical sequence.JOURNAL, Call, J., 2001, Object permanence in orangutans (Pongo pygmaeus), chimpanzees (Pan troglodytes), and children (Homo sapiens), Journal of Comparative Psychology, 115, 2, 159–171, 10.1037/0735-7036.115.2.159, 11459163, JOURNAL, Itakura, S., Tanaka, M., June 1998, Use of experimenter-given cues during object-choice tasks by chimpanzees (Pan troglodytes), an orangutan (Pongo pygmaeus), and human infants (Homo sapiens), Journal of Comparative Psychology, 112, 2, 119–126, 10.1037/0735-7036.112.2.119, 9642782, JOURNAL, Gouteux, S., Thinus-Blanc, C., Vauclair, J., 2001, Rhesus monkeys use geometric and nongeometric information during a reorientation task, Journal of Experimental Psychology: General, 130, 3, 505–519, 10.1037/0096-3445.130.3.505,weblink Research in primate cognition explores problem solving, memory, social interaction, a theory of mind, and numerical, spatial, and abstract concepts.BOOK, Tomasello, M., Call, J., yes, Primate Cognition, Oxford University Press US, 1997, 978-0-19-510624-4, 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.JOURNAL, Deaner, R. O., van Schaik, C. P., Johnson, V. E., 2006, Do some taxa have better domain-general cognition than others? A metaanalysis of nonhuman primate studies, Evolutionary Psychology (journal), Evolutionary Psychology, 4,weblink 149–196, 2011-07-04, dead,weblink" title="web.archive.org/web/20110728153456weblink">weblink 2011-07-28, 10.1177/147470490600400114, JOURNAL, Reader, S. M., Hager, Y., Laland, K. N., The evolution of primate general and cultural intelligence, Philosophical Transactions of the Royal Society B, 366, 1567, 1017–1027, 2011,weblink 10.1098/rstb.2010.0342, 21357224, 2011-07-04, dead,weblink" title="web.archive.org/web/20111003152030weblink">weblink 2011-10-03, 3049098, However, there is a great deal of variation in each group (e.g., among New World monkeys, both spider and capuchin monkeys have scored highly by some measures), as well as in the results of different studies.Lemurs, lorises, tarsiers, and New World monkeys rely on olfactory signals for many aspects of social and reproductive behavior. Specialized glands are used to mark territories with pheromones, which are detected by the vomeronasal organ; this process forms a large part of the communication behavior of these primates. In Old World monkeys and apes this ability is mostly vestigial, having regressed as trichromatic eyes evolved to become the main sensory organ.JOURNAL, Relaxed selective pressure on an essential component of pheromone transduction in primate evolution, Liman, E. R., Innan, H., Proceedings of the National Academy of Sciences of the United States of America, 2003, 100, 6, 3328–3332,weblink 2008-07-23, 10.1073/pnas.0636123100, 12631698, 152292, 2003PNAS..100.3328L, Primates also use vocalizations, gestures, and facial expressions to convey psychological state.BOOK, Egnor, R., Miller, C., Hauser, M.D., 2004, Nonhuman Primate Communication, Encyclopedia of Language and Linguistics, 2nd, Elsevier, 0-08-044299-4,weblink dead,weblink" title="web.archive.org/web/20080910043811weblink">weblink 2008-09-10, Facial musculature is complex in primates, particularly in monkeys and apes. Like humans, chimpanzees can distinguish the faces of familiar and unfamiliar individuals.JOURNAL, Burrows, A. M., 2008, The facial expression musculature in primates and its evolutionary significance, BioEssays, 30, 3, 212–225, 10.1002/bies.20719, 18293360,weblink Hand and arm gestures are also important forms of communication for great apes and a single gesture can have multiple functions.JOURNAL, Pollick, A. S., de Waal, F. B. M., 2007, Ape gestures and language evolution, Proceedings of the National Academy of Sciences, 104, 19, 8184–8189, 10.1073/pnas.0702624104, 17470779, 1876592, 2007PNAS..104.8184P, {{sound|filename=Howler monkey.ogg|right|title=Howler monkey roaring|description=Howler monkey roaring}}The Philippine tarsier, has a high-frequency limit of auditory sensitivity of approximately 91 kHz with a dominant frequency of 70 kHz. Such values are among the highest recorded for any terrestrial mammal, and a relatively extreme example of ultrasonic communication. For Philippine tarsiers, 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.JOURNAL, Ramsier, M.A., Cunningham, A.J., Moritz, G.L., Finneran, J.J., Williams, C.V., Ong, P.S., Gursky-Doyen, S.L., Dominy, N.J., 2012, Primate communication in the pure ultrasound, Biology Letters, 8, 4, 508–511, 10.1098/rsbl.2011.1149, 22319094, 3391437, Male howler monkeys are among the loudest land mammals and their roars can be heard up to {{Convert|4.8|km|mi|abbr=on}}.WEB, Black howler monkey, Smithsonian’s National Zoo & Conservation Biology Institute,weblink 2016-07-10, Roars are produced by modified larynx and enlarged hyoid bone which contains an air sac.JOURNAL, Kelemen, G., Sade, J., 1960, The vocal organ of the Howling monkey (Alouatta palliata), Journal of Morphology, 107, 2, 123–140, 10.1002/jmor.1051070202, 13752246, These calls are thought to relate to intergroup spacing and territorial protection as well as possibly mate-guarding.JOURNAL, da Cunha, R. G. T., Byrne, R., 2006, Roars of Black Howler Monkeys (Alouatta caraya): Evidence for a Function in Inter-Group Spacing, Behaviour, 143, 10, 1169–1199, 4536401, 10.1163/156853906778691568, 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. For example, if an alarm call signals a python, the monkeys climb into the trees, whereas the eagle alarm causes monkeys to seek a hiding place on the ground.JOURNAL, Seyfarth, R. M., D. L., Cheney, Peter, Marler, Vervet Monkey Alarm Calls: Semantic communication in a Free-Ranging Primate, Animal Behaviour, 28, 4, 1070–1094, 1980, 10.1016/S0003-3472(80)80097-2,weblink Many non-human primates have the vocal anatomy to produce human speech but lack the proper brain wiring.JOURNAL, Fitch, W. T., de Boer, B., Mathur, N., Ghazanfar, A. A., 2016, Monkey vocal tracts are speech-ready, Science Advances, 2, 12, e1600723, 10.1126/sciadv.1600723, 27957536, 2016SciA....2E0723F, 5148209, Vowel-like vocal patterns have been recorded in baboons which has implications for the origin of speech in humans.JOURNAL, Boë L.-J., Berthommier, F., Legou, T., Captier, G., Kemp, C., Sawallis, T. R., Becker Y., Rey, A., Fagot, J., 6, 2017, Evidence of a Vocalic Proto-System in the Baboon (Papio papio) Suggests Pre-Hominin Speech Precursors, PLOS ONE, 12, 1, e0169321, 10.1371/journal.pone.0169321, 28076426, 2017PLoSO..1269321B, 5226677,

Aggression

Non-human primates and humans have been observed to be very similar in terms of personality, such as chimpanzees having “’Big Five’ personality factors found in humans, i.e. neuroticism, extraversion, openness, agreeableness, and conscientiousness”. Primates seem to possess a sixth personality trait, dominance.JOURNAL, 2006-01-01, Psychopathology in great apes: Concepts, treatment options and possible homologies to human psychiatric disorders, Neuroscience & Biobehavioral Reviews, en, 30, 8, 1246–1259, 10.1016/j.neubiorev.2006.09.002, 17141312, 0149-7634, Brune, M., Brunecohrs, U., McGrew, W., Preuschoft, S., Chimpanzees and bonobos tend to be more social than male orangutans, who cannot be in the same living space. Male gorillas display aggression during a majority of their interactions while chimpanzees reconcile after their aggressive encounters. Killing males in differing groups in order to expand territory is an act of violence only seen in chimps and humans. Bonobos tend to live in cohesive communities with lower aggressive incidents.JOURNAL, 2015-05-01, Behavioural, hormonal and neurobiological mechanisms of aggressive behaviour in human and nonhuman primates, Physiology & Behavior, en, 143, 121–135, 10.1016/j.physbeh.2015.02.053, 25749197, 0031-9384, De Almeida, Rosa Maria Martins, Cabral, João Carlos Centurion, Narvaes, Rodrigo, This reduction in reactive aggression committed by male bonobos may be due to the evolution of prominent female groups.Both humans and nonhuman primates, such as chimpanzees, exhibit proactive aggression, a type of preplanned aggression with a reward. This aggression is expressed between neighboring groups. Proactive aggression ultimately increases the fitness of the community as a whole as the size of the community increases. Unlike proactive aggression, reactive aggression is low in humans but high in chimpanzees. Reactive aggression is a result of anger in order to cease a stressful stimulus. Low reactive aggression in humans can be attributed to tolerance and cooperation. Wrangham found “evolution of within-group tolerance, such as individual selection for cooperative breeding, group selection for parochial altruism, and cultural group selection for prosocial norms”. Ranking in nonhuman primates stems from the most aggressive male, while nomadic hunter-gatherers are respected for their prestige and ability to form alliances and negotiations. Boehm proposed the “execution hypothesis”, or the approval by the community of killing group members who violate the norm of the group or are deemed a bully. Hunter-gatherers who attempted to utilize aggression were viewed as a threat and removed from the group. The removal of community threats can be likened to modern capital punishment, more specifically the death penalty.JOURNAL, Wrangham, Richard W., 2018-01-09, Two types of aggression in human evolution, Proceedings of the National Academy of Sciences, en, 115, 2, 245–253, 10.1073/pnas.1713611115, 1091-6490, 29279379, 5777045,

Life history

File:Crab-eating Macaque nursing.jpg|A thumbPrimates have slower rates of development than other mammals. All primate infants are 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 weaned the mother can reproduce again. This often leads to weaning conflict with infants who attempt to continue breastfeeding.Infanticide is common in polygynous species such as gray langurs 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.JOURNAL, Opie, Christopher, Atkinson, Quentin D., Dunbarc, Robin I. M., Shultz, Susanne, 2013, Male infanticide leads to social monogamy in primates, Proceedings of the National Academy of Sciences of the United States of America, 110, 33, 13328–13332, 10.1073/pnas.1307903110, 23898180, 3746880, 2013PNAS..11013328O, Promiscuity may also lessen the risk of infanticide since paternity becomes uncertain.JOURNAL, De Ruiter, Jan R., Van Hooff, Jan A. R. A. M., Jan van Hooff, Scheffrahn, Wolfgang, yes, 1994, Social and genetic aspects of paternity in wild long-tailed macaques (Macaca fascicularis), Behaviour, 129, 3–4, 203–24, 4535195, 10.1163/156853994x00613, Primates have a longer juvenile period between weaning and sexual maturity than other mammals of similar size. Some primates such as galagos and new world monkeys use tree-holes for 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.JOURNAL, Kappeler, Peter M., 1998, Nests, Tree Holes, and the Evolution of Primate Life Histories, American Journal of Primatology, 1, 7–33, 9730211, 10.1002/(SICI)1098-2345(1998)46:13.0.CO;2-#, 46, JOURNAL, Ross, Caroline, 1991, Park or ride? Evolution of infant carrying in primates., International Journal of Primatology, Kluwer Academic Publishing, 22, 5, 749–771, 10.1023/A:1012065332758, 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. They learn social and fighting skills, often through playing. Primates, especially females, have longer lifespans than other similarly sized mammals, this may be partially due to their slower metabolisms.WEB,weblink Humans And Primates Burn 50 Percent Fewer Calories Each Day Than Other Mammals, Mintz, Zoe, 14 January 2014, www.ibtimes.com, IBT Media Inc., 2014-01-14, Late in life, female catarrhine primates appear to undergo a cessation of reproductive function known as menopause; other groups are less studied.JOURNAL, Walker ML, Herndon JG, Menopause in nonhuman primates?, Biology of Reproduction, 79, 398–406, 2008, 18495681, 10.1095/biolreprod.108.068536, 3, 2553520, Herndon,

Diet, feeding and hunting

File:Colubusmonkey.JPG|thumb|Leaf eating mantled guereza, a species of black-and-white colobusblack-and-white colobusFile:Microcebus murinus -Artis Zoo, Amsterdam, Netherlands-8c.jpg|thumb|right|A mouse lemurmouse lemurPrimates 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.MAGAZINE, Milton, K., 1993, Diet and Primate Evolution,weblink Scientific American, 269, 2, 86–93, 8351513, 10.1038/scientificamerican0893-86, 1993SciAm.269b..86M, Most primates include fruit in their diets to obtain easily digested nutrients including carbohydrates and lipids for energy. Primates in the suborder Strepsirrhini (non-tarsier prosimians) are able to synthesize vitamin C, like most other mammals, while primates of the suborder Haplorrhini (tarsiers, monkeys and apes) have lost this ability, and require the vitamin in their diet.JOURNAL, Vitamin C biosynthesis in prosimians: Evidence for the anthropoid affinity of Tarsius, Pollock, J. I., Mullin, R. J., American Journal of Physical Anthropology, 1986, 73, 1, 65–70, 10.1002/ajpa.1330730106,weblinkweblink" title="archive.today/20120628232930weblink">weblink dead, 2012-06-28, 3113259, Many primates have anatomical specializations that enable them to exploit particular foods, such as fruit, leaves, gum or insects. For example, leaf eaters such as howler monkeys, black-and-white colobuses and sportive lemurs have extended digestive tracts which enable them to absorb nutrients from leaves that can be difficult to digest. Marmosets, 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. 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.JOURNAL, Milliken, G. W., Ward, J. P., Erickson, C. J., 1991, Independent digit control in foraging by the aye-aye (Daubentonia madagascariensis), Folia Primatologica, 56, 4, 219–224, 10.1159/000156551, 1937286, Some species have additional specializations. For example, the grey-cheeked mangabey has thick enamel on its teeth, enabling it to open hard fruits and seeds that other monkeys cannot. The gelada is the only primate species that feeds primarily on grass.WEB, Hiller, C., 2000,weblink Theropithecus gelada, 2008-08-08, Animal Diversity Web,

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.)Tarsiers are the only extant obligate carnivorous primates, exclusively eating insects, crustaceans, small vertebrates and snakes (including venomous species).BOOK, Tarsiers Past, Present and Future, Introduction, Wright, P., Simmons, E., Gursky, S., Wright, P., Simmons, E., Gursky, S., 2003, Rutgers University Press, 0-8135-3236-1, 1, Capuchin monkeys can exploit many different types of plant matter, including fruit, leaves, flowers, buds, nectar and seeds, but also eat insects and other invertebrates, bird eggs, and small vertebrates such as birds, lizards, squirrels and bats.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%.BOOK, Goodall, Jane, Jane Goodall, 1986, The Chimpanzees of Gombe: Patterns of Behavior, 0-674-11649-6,weblink WEB, Guernsey, Paul, WHAT DO CHIMPS EAT?,weblink All About Wildlife, 2013-04-22, The meat consumption includes predation on other primate species, such as the western red colobus monkey. This sometimes involves tool use. Common chimpanzees sharpen sticks to use as weapons when hunting mammals. This is considered the first evidence of systematic use of weapons in a species other than humans. Researchers documented 22 occasions where wild chimpanzees fashioned sticks into "spears" to hunt lesser bush babies (Galago senegalensis). In each case, a chimpanzee modified a branch by breaking off one or both ends and, frequently using its teeth, sharpened the stick. The tools, on average, were about 60 cm (24 in) long and 1.1 cm (0.4 in) in circumference. The chimpanzees then jabbed their spears into hollows in tree trunks where bush babies slept. There was a single case in which a chimpanzee successfully extracted a bush baby with the tool. The bonobo is an omnivorous frugivore – the majority of its diet is fruit, but it supplements this with leaves, meat from small vertebrates, such as anomalures, flying squirrels and duikers,JOURNAL, Ihobe H, Observations on the meat-eating behavior of wild bonobos (Pan paniscus) at Wamba, Republic of Zaire, Primates, 33, 2, 247–250, 1992, 10.1007/BF02382754, and invertebrates.BOOK, Rafert, J., E.O., Vineberg, 1997, Bonobo Nutrition – relation of captive diet to wild diet,weblink dead,weblink" title="web.archive.org/web/20120425232556weblink">weblink 2012-04-25, Bonobo Husbandry Manual, American Association of Zoos and Aquariums, In some instances, bonobos have been shown to consume lower-order primates.JOURNAL, Surbeck, M, Fowler, A, Deimel, C, Hohmann, G, Evidence for the consumption of arboreal, diurnal primates by bonobos (Pan paniscus), American Journal of Primatology, 71, 2, 171–4, 2008, 10.1002/ajp.20634, 19058132, JOURNAL, Surbeck M, Hohmann G, Primate hunting by bonobos at LuiKotale, Salonga National Park, Current Biology, 18, 19, R906–7, 14 October 2008, 10.1016/j.cub.2008.08.040, 18957233, Hohmann,

As prey

Predators of primates include various species of carnivorans, birds of prey, reptiles, 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 adaptations including crypsis, alarm calls and 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.BOOK, Fichtel, Claudia, Predation, 2012, The Evolution of Primate Societies, Mitani, John C., Call, Josep, Kappeler, Peter M., Palombit, Ryne A., Silk, Joan B, University of Chicago Press, 169–84, 978-0-226-53172-4,

Tool use and manufacture

Tool use

File:Gorilla tool use.png|thumb|right|A western lowland gorillawestern lowland gorillaFile:Macaca fascicularis aurea stone tools - journal.pone.0072872.g002.png|thumb|Crab-eating macaques with stone toolstone toolThere are many reports of non-human primates using tools, both in the wild or when captive. The use of tools by primates is varied and includes hunting (mammals, invertebrates, fish), collecting honey, processing food (nuts, fruits, vegetables and seeds), collecting water, weapons and shelter.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.WEB,weblink Tool use, hunting & other discoveries, The Jane Goodall Institute, 2013-08-01, 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.WEB,weblink Bonobos, ApeTag, 2010, 2013-08-03, dead,weblink 2013-11-02, It has been reported that females, both chimpanzee and bonobo, use tools more avidly than males.JOURNAL, Gruber, T., Clay, Z., Zuberbühler, K., 2010, A comparison of bonobo and chimpanzee tool use: evidence for a female bias in the Pan lineage, Animal Behaviour, 80, 6, 1023–1033,weblink 10.1016/j.anbehav.2010.09.005, Orangutans in Borneo scoop catfish out of small ponds. Anthropologist Anne Russon saw several animals on these forested islands learn on their own to jab at catfish with sticks, so that the panicked prey would flop out of ponds and into the orangutan's waiting handsMAGAZINE,weblink Orangutans use simple tools to catch fish, Bower, B., Wired (website), Wired, 18 April 2011, 2013-08-05, There are few reports of gorillas 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.JOURNAL, Breuer, T., Ndoundou-Hockemba, M., Fishlock, V., 2005, First observation of tool use in wild gorillas, PLOS Biology, 3, 11, e380, 10.1371/journal.pbio.0030380, 16187795, 1236726, The black-striped capuchin was the first non-ape primate for which tool use was documented in the wild; individuals were observed cracking nuts by placing them on a stone anvil and hitting them with another large stone.JOURNAL, Fragaszy, D., Izar, P., Visalberghi, E., Ottoni, E.B., de Oliveira, M.G., 2004, Wild capuchin monkeys (Cebus libidinosus) use anvils and stone pounding tools, American Journal of Primatology, 64, 4, 359–366, 10.1002/ajp.20085, 15580579, In Thailand and Myanmar, crab-eating macaques use stone tools to open nuts, oysters and other bivalves, and various types of sea snails.JOURNAL, Gumert, M.D., Kluck, M., Malaivijitnond, S., The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the Andaman Sea region of Thailand, American Journal of Primatology, 2009, 71, 7, 594–608, 10.1002/ajp.20694, 19405083, 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.JOURNAL, Hamilton, W.J., Buskirk, R.E., Buskirk, W.H., 1975, Defensive stoning by baboons, Nature, 256, 488–489, 10.1038/256488a0, 5517, 1975Natur.256..488H, 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.BOOK, Fichtel, C., Kappeler, P. M., Chapter 19: Human universals and primate symplesiomorphies: Establishing the lemur baseline, Kappeler, P. M., Silk, J. B., Mind the Gap: Tracing the Origins of Human Universals, Springer, 2010, 978-3-642-02724-6,weblink

Tool manufacture

Tool manufacture is much rarer than simple tool use and probably represents higher cognitive functioning. 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. Both bonobos and chimpanzees have also been observed making "sponges" out of leaves and moss that suck up water and are used as grooming tools. Sumatran orangutans have been observed making and using tools. They will break off a tree branch that is about 30 cm long, snap off the twigs, fray one end and then use the stick to dig in tree holes for termites.WEB,weblink Sumatran orangutans, OrangutanIslands.com, 2013-08-02, dead,weblink 2013-11-26, JOURNAL, van Schaik, C., Fox, E., Sitompul, A., 1996, Manufacture and use of tools in wild Sumatran orangutans, Naturwissenschaften, 83, 4, 186–188, 10.1007/BF01143062, 8643126, 1996NW.....83..186V, 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.WEB,weblink Mandrill monkey makes 'pedicuring' tool, Victoria, Gill, BBC, 22 July 2011, 2013-08-11, Captive gorillas have made a variety of tools.WEB,weblink Vancatova, M., Gorillas and Tools – Part I, 2008, 2013-08-04,

Ecology

{{See also|List of primates by population}}File:Macaque India 3.jpg|thumb|upright|right|Rhesus macaque at Agra Fort, IndiaIndiaNon-human primates primarily live in the tropical latitudes of Africa, Asia, and the Americas. Species that live outside of the tropics; include the Japanese macaque which lives in the Japanese islands of HonshÅ« and Hokkaido; the Barbary macaque which lives in North Africa and several species of langur which live in China. Primates tend to live in tropical rainforests but are also found in temperate forests, savannas, deserts, mountains and coastal areas.BOOK, Cowlishaw, G., Clutton-Brock, T., Primates, 2009, The Princeton Encyclopedia of Mammals, MacDonald, D., Princeton and Oxford University Press, 270–280, 978-0-691-14069-8, The number of primate species within tropical areas has been shown to be positively correlated to the amount of rainfall and the amount of rain forest area.JOURNAL, Geographic and climatic control of primate diversity, Reed, K., Fleagle, J., Proceedings of the National Academy of Sciences of the United States of America, 92, 17, 7874–7876, August 15, 1995, 10.1073/pnas.92.17.7874, 7644506, 41248, 1995PNAS...92.7874R, Accounting for 25% to 40% of the fruit-eating animals (by weight) within tropical rainforests, primates play an important ecological role by dispersing seeds of many tree species.BOOK, Primates in Perspective, Chapman, C., Russo, S., yes, Primate Seed Dispersal, 2007, Oxford University Press, Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M., Bearder, S. K., 978-0-19-517133-4, 510, Primate habitats span a range of altitudes: the black snub-nosed monkey has been found living in the Hengduan Mountains at altitudes of 4,700 meters (15,400 ft),JOURNAL, Long, Y. C., Kirkpatrick, R. C., Zhong, T., Xiao, L., April 1994, Report on the distribution, population, and ecology of the Yunnan snub-nosed monkey (Rhinopithecus bieti), Primates, 35, 2, 241–250, 10.1007/BF02382060, the mountain gorilla can be found at 4,200 meters (13,200 ft) crossing the Virunga Mountains,BOOK, Schaller, G. B., 1963, The Mountain Gorilla: Ecology and Behavior, Chicago, University Chicago Press, 978-0-226-73635-8,weblink and the gelada has been found at elevations of up to {{Convert|5,000|m|ft|abbr=on}} in the Ethiopian Highlands.BOOK, Stammbach, E., 1987, Desert, Forest, and Montane Baboons: Multilevel-Societies, 112–120, Smuts, B, Cheney, D, Seyfarth, R, Wrangham, R, Struhsaker, T., Primate Societies, The University of Chicago Press, 978-0226767161, Some species interact with aquatic environments and may swim or even dive, including the proboscis monkey, De Brazza's monkey and Allen's swamp monkey.JOURNAL, Kemp, E., 2009, Patterns of Water Use in Primates, Folia Primatologica, 80, 4, 275–294, 10.1159/000252586, 19864919, Some primates, such as the rhesus macaque and gray langurs, can exploit human-modified environments and even live in cities.BOOK, Primates in Perspective, Wolfe, L. D., Fuentes, A., yes, Ethnoprimatology, 2007, Oxford University Press, Campbell, C. J., Fuentes, A., MacKinnon, K. C., Panger, M., Bearder, S. K., 978-0-19-517133-4, 692,

Interactions between humans and other primates

Disease transmission

Close interactions between humans and non-human primates (NHPs) can create pathways for the transmission of zoonotic diseases. Viruses such as Herpesviridae (most notably Herpes B Virus), Poxviridae, measles, ebola, rabies, the Marburg virus and viral hepatitis can be transmitted to humans; in some cases the viruses produce potentially fatal diseases in both humans and non-human primates.JOURNAL, Renquist, D. M., Whitney, R. A., yes, 1987, Zoonoses Acquired from Pet Primates, Veterinary Clinics of North America: Small Animal Practice, 17, 1, 219–240,weblink 2008-08-11, 10.1016/s0195-5616(87)50614-3,

Legal and social status

{{further|Great ape personhood}}File:Slow Loris Female.jpg|thumb|left|Slow lorises are popular in the exotic petexotic petOnly humans are recognized as persons and protected in law by the United Nations Universal Declaration of Human Rights.{{refn|group=lower-alpha|Article 6: Everyone has the right to recognition everywhere as a person before the law.WEB,weblink 1948, 2008-12-02, The Universal Declaration of Human Rights, United Nations, dead,weblink" title="web.archive.org/web/20141208080853weblink">weblink 2014-12-08, }} The legal status of NHPs, on the other hand, is the subject of much debate, with organizations such as the Great Ape Project (GAP) campaigning to award at least some of them legal rights.WEB, Cavalieri, P., Singer, P., yes,weblink Declaration on Great Apes, Great Ape Project, 2008-06-16, dead,weblink" title="web.archive.org/web/20080820011354weblink">weblink 2008-08-20, In June 2008, Spain became the first country in the world to recognize the rights of some NHPs, when its parliament's cross-party environmental committee urged the country to comply with GAP's recommendations, which are that chimpanzees, bonobos, orangutans, and gorillas are not to be used for animal experiments.NEWS, Glendinning, L.,weblink Spanish parliament approves 'human rights' for apes, The Guardian, 26 June 2008, 2008-11-10, NEWS, Singer, P.,weblink Of great apes and men, The Guardian, 18 July 2008, 2008-11-10, Many species of NHP are kept as pets by humans, the Allied Effort to Save Other Primates (AESOP) estimates that around 15,000 NHPs live as exotic pets in the United States.MAGAZINE, The Perils of Keeping Monkeys as Pets,weblink M., Mott, National Geographic, 16 September 2003, 2013-02-06, dead,weblink" title="web.archive.org/web/20130608041322weblink">weblink 2013-06-08, The expanding Chinese middle class has increased demand for NHPs as exotic pets in recent years.JOURNAL, Workman, C., June 2004, Primate conservation in Vietnam: toward a holistic environmental narrative, American Anthropologist, 106, 2, 346–352, 10.1525/aa.2004.106.2.346, Although NHP import for the pet trade was banned in the U.S. in 1975, smuggling still occurs along the United States – Mexico border, with prices ranging from US$3000 for monkeys to $30,000 for apes.WEB, International Primate Protection League, 2003, IPPL News: The US Pet Monkey Trade,weblink 2008-08-04, dead,weblink" title="web.archive.org/web/20080725190533weblink">weblink 2008-07-25, Primates are used as model organisms in laboratories and have been used in space missions.WEB,weblink The Beginnings of Research in Space Biology at the Air Force Missile Development Center, 1946–1952, 2008-08-18, Bushnell, D., 1958, History of Research in Space Biology and Biodynamics, NASA, They serve as service animals for disabled humans. Capuchin monkeys can be trained to assist quadriplegic humans; their intelligence, memory, and manual dexterity make them ideal helpers.NEWS, Monkeys as Helpers To Quadriplegics At Home, Blumenthal, D., 1987-06-17, The New York Times,weblink 2008-10-08, NHPs are kept in zoos around the globe. Historically, zoos were primarily a form of entertainment, but more recently have shifted their focus towards conservation, education and research. GAP does not insist that all NHPs should be released from zoos, primarily because captive-born primates lack the knowledge and experience to survive in the wild if released.BOOK, Newman, James L., 2013, Encountering Gorillas: A Chronicle of Discovery, Exploitation, Understanding, and Survival, Rowman and Littlefield, Plymouth, United Kingdom, 978-1-4422-1957-1, 173,

Role in scientific research

{{Further|Animal testing on non-human primates|International trade in primates}}File:Monkey Sam Before The Flight On Little Joe 2.jpg|thumb|right|upright|Sam, a rhesus macaque, was flown into space by NASANASAThousands of non-human primates are used around the world in research because of their psychological and physiological similarity to humans.JOURNAL, Chen, F. C., Li, W. H., February 2001, Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees, American Journal of Human Genetics, 68, 2, 444–456, 10.1086/318206, 11170892, 1235277, In particular, the brains and eyes of NHPs more closely parallel human anatomy than those of any other animals. NHPs are commonly used in preclinical trials, neuroscience, ophthalmology studies, and toxicity studies. Rhesus macaques are often used, as are other macaques, African green monkeys, chimpanzees, baboons, squirrel monkeys, and marmosets, both wild-caught and purpose-bred.WEB,weblink The supply and use of primates in the EU, European Biomedical Research Association, 1996, 2008-08-18, dead,weblink" title="web.archive.org/web/20120117061036weblink">weblink 2012-01-17, JOURNAL, Conlee, K. M., Hoffeld, E. H., Stephens, M. L., 2004, A Demographic Analysis of Primate Research in the United States, Alternatives to Laboratory Animals, 32, Sup 1, 315–322, 10.1177/026119290403201s52, 23577480, In 2005, GAP reported that 1,280 of the 3,100 NHPs living in captivity in the United States were used for experiments. In 2004, the European Union used around 10,000 NHPs in such experiments; in 2005 in Great Britain, 4,652 experiments were conducted on 3,115 NHPs.BOOK, Presented to Parliament by the Secretary of State for the Home Department by Command of Her Majesty, Statistics of scientific procedures on living animals: Great Britain 2005, July 2006, The Stationery Office, 0-10-168772-9,weblink 2008-06-16, Governments of many nations have strict care requirements of NHPs kept in captivity. In the US, federal guidelines extensively regulate aspects of NHP housing, feeding, enrichment, and breeding.WEB, Nonhuman Primates: Research Animals, United States Department of Agriculture, Animal Welfare Information Center,weblink 2008-07-14, dead,weblink" title="web.archive.org/web/20081003004620weblink">weblink 2008-10-03, European groups such as the European Coalition to End Animal Experiments are seeking a ban on all NHP use in experiments as part of the European Union's review of animal testing legislation.WEB,weblink Directive 86/609, European Coalition to End Animal Experiments, 2008-10-08, dead,weblink" title="web.archive.org/web/20080929214536weblink">weblink 2008-09-29,

Extinction threats

File:Hunted Silky Sifakas.jpg|thumb|left|Humans are known to hunt other primates for food, so-called bushmeat. Pictured are two men who have killed a number of silky sifaka and white-headed brown lemurs.]]The International Union for Conservation of Nature (IUCN) lists more than a third of primates as critically endangered or vulnerable. About 60% of primate species are threatened with extinction, including: 87% of species in Madagascar, 73% in Asia, 37% in Africa, and 36% in South and Central America.JOURNAL, Estrada, Alejandro, Garber, Paul A., Rylands, Anthony B., Roos, Christian, Fernandez-Duque, Eduardo, Fiore, Anthony Di, Nekaris, K. Anne-Isola, Nijman, Vincent, Heymann, Eckhard W., 2017-01-01, Impending extinction crisis of the world's primates: Why primates matter, Science Advances, en, 3, 1, e1600946, 10.1126/sciadv.1600946, 2375-2548, 5242557, 28116351, 2017SciA....3E0946E, Additionally, 75% of primate species have decreasing populations. Trade is regulated, as all species are listed by CITES in Appendix II, except 50 species and subspecies listed in Appendix I, which gain full protection from trade.BOOK, IFAW, Born to be wild: Primates are not pets, International Fund for Animal Welfare, 2005,weblink 2011-02-26, International Fund for Animal Welfare, dead,weblink" title="web.archive.org/web/20110726171340weblink">weblink 2011-07-26, WEB, CITES, Appendices I, II and III, Convention on International Trade in Endangered Species of Wild Fauna and Flora, 2010-10-14,weblink 2012-04-02, CITES, Common threats to primate species include deforestation, forest fragmentation, monkey drives (resulting from primate crop raiding),BOOK, Grubb, P., 1998, The Sierra Leone monkey drives, Mammals of Ghana, Sierra Leone, and the Gambia, St. Ives, Trendrine, 214–219, 0-9512562-4-6, and primate hunting for use in medicines, as pets, and for food. Large-scale tropical forest clearing is widely regarded as the process that most threatens primates.JOURNAL, Chapman, C. A., Peres, C. A., 2001, Primate conservation in the new millennium: the role of scientists, Evolutionary Anthropology, 10, 1, 16–33, 10.1002/1520-6505(2001)10:13.0.CO;2-O, BOOK, Mittermeier, R. A., Cheney, D. L., yes, 1987, Conservation of primates and their habitats, Primate Societies, Smuts, B. B., Cheney, D. L., Seyfarth, R. M., Wrangham, R. W., Struhsaker, T. T., University of Chicago Press, Chicago, 477–490, JOURNAL, Southwick, C. H., Siddiqi, M. F., yes, 2001, Status, conservation and management of primates in India, Envis Bulletin: Wildlife and Protected Areas, 1, 1, 81–91,weblinkweblink" title="wayback.archive-it.org/all/20081001191112weblink">weblink dead, 2008-10-01, 2008-08-04, More than 90% of primate species occur in tropical forests.BOOK, Cowlishaw, G., Dunbar, R., yes, 2000, Primate Conservation Biology, University of Chicago Press, Chicago, 978-0-226-11637-2, The main cause of forest loss is clearing for agriculture, although commercial logging, subsistence harvesting of timber, mining, and dam construction also contribute to tropical forest destruction. In Indonesia large areas of lowland forest have been cleared to increase palm oil production, and one analysis of satellite imagery concluded that during 1998 and 1999 there was a loss of 1,000 Sumatran orangutans per year in the Leuser Ecosystem alone.JOURNAL, Van Schaik, C. P., Monk, K. A., Robertson, J. M. Y., 2001, Dramatic decline in orangutan numbers in the Leuser Ecosystem, northern Sumatra, Oryx, 35, 1, 14–25, 10.1046/j.1365-3008.2001.00150.x, File:Silky Sifaka Pink Face Closeup.JPG|thumb|right|The critically endangered silky sifakasilky sifakaPrimates with a large body size (over 5 kg) are at increased extinction risk due to their greater profitability to poachers compared to smaller primates. They reach sexual maturity later and have a longer period between births. Populations therefore recover more slowly after being depleted by poaching or the pet trade.JOURNAL, Purvis, A., Gittleman, J. L., Cowlishaw, G., Mace, G. M., 2000, Predicting extinction risk in declining species, Proceedings of the Royal Society B, 267, 1947–1952, 10.1098/rspb.2000.1234, 11075706, 1456, 1690772, Data for some African cities show that half of all protein consumed in urban areas comes from the bushmeat trade.JOURNAL, Fa, J. E., Juste, J., Perez de Val, J., Castroviejo, J., 1995, Impact of market hunting on mammal species in Equatorial Guinea, Conservation Biology, 9, 5, 1107–1115, 10.1046/j.1523-1739.1995.9051107.x, Endangered primates such as guenons and the drill are hunted at levels that far exceed sustainable levels. This is due to their large body size, ease of transport and profitability per animal. As farming encroaches on forest habitats, primates feed on the crops, causing the farmers large economic losses.JOURNAL, Hill, C. M., 1997, Crop-raiding by wild vertebrates: The farmer's perspective in an agricultural community in western Uganda, International Journal of Pest Management, 43, 1, 77–84, 10.1080/096708797229022, Primate crop raiding gives locals a negative impression of primates, hindering conservation efforts.JOURNAL, Hill, C. M., 2002, Primate conservation and local communities: Ethical issues and debates, American Anthropologist, 104, 4, 1184–1194, 10.1525/aa.2002.104.4.1184, Madagascar, home to five endemic primate families, has experienced the greatest extinction of the recent past; since human settlement 1,500 years ago, at least eight classes and fifteen of the larger species have become extinct due to hunting and habitat destruction.ENCYCLOPEDIA, Primate, Encyclopædia Britannica Online, Encyclopædia Britannica, Inc., 2008,weblink 2008-07-21, Among the primates wiped out were Archaeoindris (a lemur larger than a silverback gorilla) and the families Palaeopropithecidae and Archaeolemuridae.File:Man of the woods.JPG|thumb|left|The critically endangered Sumatran orangutanSumatran orangutanIn Asia, Hinduism, Buddhism, and Islam prohibit eating primate meat; however, primates are still hunted for food. Some smaller traditional religions allow the consumption of primate meat.JOURNAL, Choudhury, A., 2001, Primates in Northeast India: an overview of their distribution and conservation status, Envis Bulletin: Wildlife and Protected Areas, 1, 1, 92–101,weblinkweblink" title="wayback.archive-it.org/all/20081001191106weblink">weblink dead, 2008-10-01, 2008-08-04, JOURNAL, Kumara, H. N., Singh, M., October 2004, Distribution and abundance of primates in rainforests of the Western Ghats, Karnataka, India and the conservation of Macaca silenus, International Journal of Primatology, 25, 5, 1001–1018, 10.1023/B:IJOP.0000043348.06255.7f, The pet trade and traditional medicine also increase demand for illegal hunting.JOURNAL, Nijman, V., 2004, Conservation of the Javan gibbon Hylobates moloch: population estimates, local extinction, and conservation priorities, The Raffles Bulletin of Zoology, 52, 1, 271–280,weblink 2008-08-04, dead,weblink" title="web.archive.org/web/20080910043814weblink">weblink 2008-09-10, JOURNAL, O'Brien, T. G., Kinnaird, M. F., Nurcahyo, A., Iqbal, M., Rusmanto, M., April 2004, Abundance and distribution of sympatric gibbons in a threatened Sumatran rain forest, International Journal of Primatology, 25, 2, 267–284, 10.1023/B:IJOP.0000019152.83883.1c, The rhesus macaque, a model organism, was protected after excessive trapping threatened its numbers in the 1960s; the program was so effective that they are now viewed as a pest throughout their range.In Central and South America forest fragmentation and hunting are the two main problems for primates. Large tracts of forest are now rare in Central America.JOURNAL, Estrada, A., Coates-Estrada, R., Meritt, D., September 1994, Non-flying mammals and landscape changes in the tropical forest region of Los Tuxtlas, Mexico, Ecography, 17, 229–241, 10.1111/j.1600-0587.1994.tb00098.x, 3, This increases the amount of forest vulnerable to edge effects such as farmland encroachment, lower levels of humidity and a change in plant life.BOOK, Marsh, L. K., 2003, The nature of fragmentation., Primates in Fragments: Ecology and Conservation, Marsh, L. K., New York, Kluwer Academic/Plenum Publishers, 1–10, 0-306-47696-7, JOURNAL, Turner, I. M., 1996, Species loss in fragments of tropical rain forest: a review of the evidence, Journal of Applied Ecology, 33, 200–209, 10.2307/2404743, 2, 2404743, Movement restriction results in a greater amount of inbreeding, which can cause deleterious effects leading to a population bottleneck, whereby a significant percentage of the population is lost.BOOK, Chiarello, A.G., 2003, Primates of the Brazilian Atlantic forest: the influence of forest fragmentation on survival, Primates in Fragments: Ecology and Conservation, Marsh, L. K., New York, Kluwer Academic/Plenum Publishers, 99–121, 978-0-306-47696-9, BOOK, Pope, T.R., 1996, Socioecology, population fragmentation, and patterns of genetic loss in endangered primates, Conservation Genetics: Case Histories from Nature, Avise, J., Hamrick, J., Norwell, Kluwer Academic Publishers, 119–159, 978-0-412-05581-2, There are 21 critically endangered primates, 7 of which have remained on the IUCN's "The World's 25 Most Endangered Primates" list since the year 2000: the silky sifaka, Delacour's langur, the white-headed langur, the gray-shanked douc, the Tonkin snub-nosed monkey, the Cross River gorilla and the Sumatran orangutan.BOOK, Mittermeier, R.A., Wallis, J., Rylands, A.B., Ganzhorn, J.U., Oates, J.F., Williamson, E.A., Palacios, E., Heymann, E.W., Kierulff, M.C.M., Yongcheng, L., Supriatna, J., Roos, C., Walker, S., Cortés-Ortiz, L., Schwitzer, C., Illustrated by S.D. Nash, Russell Mittermeier, 2009, Primates in Peril: The World's 25 Most Endangered Primates 2008–2010, IUCN/SSC Primate Specialist Group (PSG), International Primatological Society (IPS), and Conservation International (CI), Arlington, VA., 23–26, 978-1-934151-34-1,weblink Miss Waldron's red colobus was recently declared extinct when no trace of the subspecies could be found from 1993 to 1999.JOURNAL, Oates, J. F., Abedi-Lartey, M., McGraw, W. S., Struhsaker, T. T., Whitesides, G. H., Extinction of a West African Red Colobus Monkey, Conservation Biology, October 2000, 14, 5, 10.1046/j.1523-1739.2000.99230.x, 1526–1532, A few hunters have found and killed individuals since then, but the subspecies' prospects remain bleak.JOURNAL, Update on the Search for Miss Waldron's Red Colobus Monkey, McGraw, W. S., International Journal of Primatology, 26, 3, June 2005, 605–619, 10.1007/s10764-005-4368-9,

See also

{{div col|colwidth=18em}} {{div col end}}

Footnotes

{{Reflist|group=lower-alpha}}

References

{{Reflist|30em}}

Literature cited

  • BOOK, Benton, Michael J., Chapter 3: Primate evolution, Vertebrate palaeontology, Wiley-Blackwell, 2005, 978-0-632-05637-8,weblink 2011-07-10, harv,
  • BOOK, Cartmill, M., 2010, Primate Neuroethology, Platt, M., Ghazanfar, A, Primate Classification and Diversity, 10–30, Oxford University Press, 978-0-19-532659-8,weblink harv,
  • BOOK, Hartwig, W., Campbell, Chapter 3: Primate evolution, 19–31, C. J., Fuentes, A., MacKinnon, K. C., Bearder, S. K., Stumpf, R. M, Primates in Perspective, 2nd, Oxford University Press, 2011, 978-0-19-539043-8, harv,
  • BOOK, Szalay, F.S., Delson, E., Evolutionary History of the Primates, 1980, Academic Press, 978-0126801507, 893740473,weblink harv,

Further reading

{{Library resources box|onlinebooks=yes|by=no|lcheading= Primates|label=Primates}}
  • BOOK, 1984, David J. Chivers, Bernard A. Wood, Alan Bilsborough, Food Acquisition and Processing in Primates, New York & London, Plenum Press, 0-306-41701-4,

External links

{{Mammals}}{{Primates}}{{Living things in culture}}{{Featured article}}{{Taxonbar|from=Q7380}}{{Authority control}}


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