Array ( [0] => {{short description|Order of mammals}} [1] => {{About|the type of animal}} [2] => {{pp-move|small=yes}} [3] => {{automatic taxobox [4] => | name = Primates [5] => | fossil_range = {{Fossil range|65.9|0|earliest=85}} [6] => [[Early Paleocene]] to [[Holocene|Present]] [7] => | image = {{Multiple image [8] => | perrow = 2/2/2/2 [9] => | total_width = 280 [10] => | caption_align = center [11] => | image1 = Lemur catta 001.jpg [12] => | caption1 = [[Ring-tailed lemur]] [13] => | image3 = Loris tardigradus tardigradus 003.jpg [14] => | caption3 = [[Red slender loris]] [15] => | image5 = Garnett's Galago (Greater Bushbaby).jpg [16] => | caption5 = [[Northern greater galago]] [17] => | image6 = Bohol Tarsier.jpg [18] => | caption6 = [[Philippine tarsier]] [19] => | image7 = Ateles paniscus -Brazil-8.jpg [20] => | caption7 = [[Black spider monkey]] [21] => | image8 = Papio hamadryas (aka).jpg [22] => | caption8 = [[Hamadryas baboon]] [23] => | image9 = RickMurphy graueri72.jpg [24] => | caption9 = [[Eastern gorilla]] [25] => | image10 = Akha cropped hires.JPG [26] => | caption10 = [[Human]]s [27] => | border = infobox [28] => }} [29] => | taxon = Primates [30] => | authority = [[Carl Linnaeus|Linnaeus]], 1758{{MSW3 Groves|pages=111–184|id=12100001}} [31] => | subdivision_ranks = Suborders [32] => | subdivision = *[[Strepsirrhini]] [33] => *[[Haplorhini]] [34] => *†''[[Altiatlasius]]'' [35] => '''sister:''' [[Dermoptera]] [36] => | range_map = Primate_SpeciesDensity.png [37] => | range_map_caption = Range and density of non-human primates. [38] => | synonyms = [[Plesiadapiformes]] ([[cladistically]] including crown primates{{Cite journal|last1=Silcox|first1=Mary T.|last2=Bloch|first2=Jonathan I.|last3=Boyer|first3=Doug M.|last4=Chester|first4=Stephen G. B.|last5=López-Torres|first5=Sergi|date=2017|title=The evolutionary radiation of plesiadapiforms|journal=Evolutionary Anthropology: Issues, News, and Reviews|language=en|volume=26|issue=2|pages=74–94|doi=10.1002/evan.21526|pmid=28429568|issn=1520-6505|doi-access=free}}) [39] => }} [40] => [41] => [42] => '''Primates''' are the members of a diverse [[order (biology)|order]] of [[mammal]]s. They are divided into the [[Strepsirrhini|strepsirrhine]]s, which include the [[lemurs]], [[galagos]], and [[lorisid]]s, and the [[Haplorhini|haplorhine]]s, which include the [[Tarsiiformes|tarsiers]] and the [[Simiiformes|simian]]s ([[monkey]]s and [[ape]]s). Primates arose 85–55 million years ago first from small [[Terrestrial animal|terrestrial]] mammals, which adapted to living in the trees of [[tropical forest]]s: many primate characteristics represent adaptations to life in this challenging environment, including large brains, [[visual acuity]], [[color vision]], a [[shoulder girdle]] allowing a large degree of movement in the shoulder joint, 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 376–524 [[species]] of living primates, depending on which classification is used. New primate species continue to be discovered: over 25 species were described in the 2000s, 36 in the 2010s, and [[List of primates described in the 2020s|six in the 2020s]]. [43] => [44] => 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 [[olfaction|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. Most primates also have [[thumb|opposable thumbs]]. Some primates, including gorillas, humans, and baboons, are primarily [[terrestrial animal|terrestrial]] rather than [[arboreal]], but all species have adaptations for climbing trees. Arboreal [[Animal locomotion|locomotion]] techniques used include leaping from tree to tree and swinging between branches of trees ([[brachiation]]); terrestrial locomotion techniques include walking on two limbs ([[bipedal]]ism) and modified walking on four limbs ([[knuckle-walking]]). [45] => [46] => 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 structure|social systems]], many defined by the amount of movement by adolescent females between groups. Primates have slower rates of development than other similarly sized mammals, reach maturity later, and have longer lifespans. Primates are also the most [[primate cognition|intelligent]] animals and non-human primates are recorded to [[Tool use by animals|use tools]]. They may communicate using facial and hand gestures, smells and vocalizations. [47] => [48] => Close interactions between humans and non-human primates (NHPs) can create opportunities for the transmission of [[Zoonosis|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 drive]]s, and primate hunting for use in medicines, as pets, and for food. Large-scale tropical forest clearing for agriculture most threatens primates. [49] => [50] => ==Etymology== [51] => The English name ''[[wikt:primate#English|primates]]'' is derived from [[Old French]] or French {{lang|fr|primat}}, from a noun use of Latin {{lang|la|primat-}}, from {{lang|la|primus}} ('prime, first rank').{{cite encyclopedia | title=Primate | encyclopedia=Merriam-Webster Online Dictionary |publisher= [[Merriam-Webster]] | url=http://www.merriam-webster.com/dictionary/primate | access-date=2008-07-21}} The name was given by [[Carl Linnaeus]] because he thought this the "highest" order of animals.{{cite book|title=The Book of Popular Science|page=257|year=1963|url=https://books.google.com/books?id=qNtjfh57ipsC&q=primate+carl+linnaeus+highest}} 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.{{cite EB1911|wstitle= Ape | volume= 02 | page = 160 }}{{cite news |last=Weisberger |first=Mindy |title=Why don't humans have tails? Scientists find answers in an unlikely place |url=https://www.cnn.com/2024/03/23/world/humans-tails-genetic-mutation-junk-dna-scn/index.html |date=March 23, 2024 |work=[[CNN]] |url-status=live |archiveurl=https://archive.today/20240324031927/https://www.cnn.com/2024/03/23/world/humans-tails-genetic-mutation-junk-dna-scn/index.html |archivedate=March 24, 2024 |accessdate=March 24, 2024 }} [52] => [53] => 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 ''[[prosimian]]s'', ''[[monkey]]s'', ''[[lesser ape]]s'', and ''[[great ape]]s'' reflect this methodology. According to our current understanding of the evolutionary history of the primates, several of these groups are [[paraphyletic]], or rather they do not include all the descendants of a common ancestor.Definitions of [[paraphyly]] vary; for the one used here see e.g. {{Citation|last=Stace |first=Clive A. |author-link=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 |access-date=2010-02-07 |url-status=dead |archive-url=https://web.archive.org/web/20110726104027/https://www.watsonia.org.uk/Wats28p103.pdf |archive-date=2011-07-26}}. [54] => [55] => 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 |name-list-style=amp}}. [56] => [57] => The [[cladogram]] below shows one possible classification sequence of the living primates:{{cite book | last1 = Cartmill | first1 = M. | last2 = Smith | first2 = F. H. | title = The Human Lineage | publisher = John Wiley & Sons | year = 2011 | isbn = 978-1-118-21145-8 | url = https://books.google.com/books?id=X058kYnhxC0C&pg=PA90}} groups that use common (traditional) names are shown on the right. [58] => {{Barlabel|style=font-size:90%;|labelwidth=6 [59] => |size=9 [60] => |at1=-3.5|color1=purple|label1=prosimians [61] => |at2=1.3|color2=crimson|label2=monkeys [62] => |at3=4.5|color3=darkgreen|label3=lesser apes [63] => |at4=12|color4=green|label4=great apes [64] => |cladogram= [65] => {{Clade [66] => |label1=[[Primatomorpha]] [67] => |1={{Clade [68] => |1=[[Dermoptera]][[File:Cynocephalus volans Brehm1883 (white background).jpg|50 px]] [69] => |label2='''Primates''' [70] => |2={{Clade [71] => |label1=[[Strepsirrhini]] [72] => |1={{Clade [73] => |label1=[[Lemuriformes]]{{efn|name=Lemuriformes}} [74] => |1={{Clade [75] => |1=[[lemur]]s (superfamily Lemuroidea)[[File:FMIB 46849 Primates Maki Moccoe Lemur catta (white background).jpeg|35 px]]|barbegin1=purple [76] => |2=[[Lorisoidea|lorises and allies]] (superfamily Lorisoidea)[[File:Nycticebus (white background).jpg|25 px]]|bar2=purple [77] => }} [78] => }} [79] => |label2=[[Haplorhini]] [80] => |2={{Clade [81] => |label1=[[Tarsiiformes]] [82] => |1=[[tarsier]]s (superfamily Tarsioidea)[[File:Säugethiere vom Celebes- und Philippinen-Archipel (Taf. III) (white background) (1).jpg|25 px]]|barend1=purple [83] => |label2=[[Simiiformes]] [84] => |2={{Clade [85] => |1=[[New World monkey]]s (parvorder Platyrrhini)[[File:Die Säugthiere in Abbildungen nach der Natur, mit Beschreibungen (Plate 8) (white background).jpg|50 px]]|barbegin1=crimson [86] => |label2=[[Catarrhini]] [87] => |2={{Clade [88] => |1=[[Old World monkey]]s (superfamily Cercopithecoidea)[[File:Cynocephalus doguera - 1700-1880 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - (white background).tiff|60 px]]|barend1=crimson [89] => |label2=[[Hominoidea]] [90] => |2={{Clade [91] => |1=[[gibbon]]s (family Hylobatidae)[[File:Le gibbon (white background).jpg|50 px]]|barbegin1=green|barend1=green [92] => |label2=[[Hominidae]] [93] => |2={{Clade [94] => |1=[[orangutan]]s (subfamily Ponginae)[[File:Simia satyrus - 1837 - Print - Iconographia Zoologica - Special Collections University of Amsterdam - White Background.jpg|60 px]]|barbegin1=darkgreen [95] => |label2=[[Homininae]] [96] => |2={{Clade [97] => |1=[[gorilla]]s (tribe Gorillini)[[File:Gorila de llanura occidental. Gorilla gorilla - Blanca Martí de Ahumada (white background).jpg|60 px]]|bar1=darkgreen [98] => |label2=[[Hominini]] [99] => |2={{Clade [100] => |1=[[Homo|human]]s (g. ''Homo'')[[File:Silhouette of a woman walking.svg|60 px]]|bar1=darkgreen [101] => |2=[[Pan (genus)|chimpanzees, bonobo]]s (g. ''Pan'')[[File:PanTroglodytesSmit (white background).jpg|50 px]]|barend2=darkgreen [102] => }} [103] => }} [104] => }} [105] => }} [106] => }} [107] => }} [108] => }} [109] => }} [110] => }} [111] => }} [112] => }} [113] => [114] => All groups with scientific names are [[clade]]s, or monophyletic groups, 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: [115] => [116] => * 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. [117] => * 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. [118] => * Apes as a whole, and the [[great ape]]s, are paraphyletic if the terms are used such that they exclude humans. [119] => [120] => 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]] [[palaeontology|palaeontologist]] Benton wrote, "The apes, Hominoidea, today include the [[gibbon]]s and [[orangutan]] ... the [[gorilla]] and [[chimpanzee]] ... and [[human]]s";{{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. [121] => [122] => {{As of|2021}}, 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}} [123] => [124] => ==Classification of living primates== [125] => [[File:Primates-drawing.jpg|thumb|upright|A 1927 drawing of [[chimpanzee]]s, a [[gibbon]] (top right) and two [[orangutan]]s (center and bottom center): The chimpanzee in the upper left is [[Brachiation|brachiating]]; the orangutan at the bottom center is [[knuckle-walking]].]] [126] => [[File:AFR V4 D225 Kafir man and woman.png|upright|thumb|''[[Homo sapiens]]'' is the only living primate species that is fully bipedal]] [127] => [[File:NILGIRI LANGUR (Trachypithecus johnii).jpg|thumb|upright|[[Nilgiri langur]] (''Trachypithecus johnii''), an [[Old World monkey]]]] [128] => [129] => A list of the families of the living primates is given below, together with one possible classification into ranks between order and family.{{cite book |author1=Rylands, A. B. |author2=Mittermeier, R. A. |name-list-style=amp | title = South American Primates: Comparative Perspectives in the Study of Behavior, Ecology, and Conservation | chapter = The Diversity of the New World Primates (Platyrrhini) | publisher = Springer | editor = Garber, P. A. |editor2=Estrada, A. |editor3=Bicca-Marques, J. C. |editor4=Heymann, E. W. |editor5=Strier, K. B.|editor5-link=Karen B. Strier | isbn = 978-0-387-78704-6 | year = 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}} [130] => * '''Order Primates''' [131] => ** '''Suborder [[Strepsirrhini]]''': lemurs, galagos and lorisids [132] => *** Infraorder [[Lemuriformes]]{{efn|name=Lemuriformes|Although the [[Monophyly|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 [[Taxonomic rank|infraorder]] and the extinct, non-toothcombed [[Adapiformes|adapiforms]] into another, both within the [[Taxonomic rank|suborder]] Strepsirrhini.{{Sfn|Szalay|Delson|1980|p=149}}{{Sfn|Cartmill|2010|p=15}} However, another popular alternative taxonomy places the [[Lorisoidea|lorisoids]] in their own infraorder, Lorisiformes.{{Sfn|Hartwig|2011|pp=20–21}}}} [133] => **** Superfamily [[Lemuroidea]] [134] => ***** Family [[Cheirogaleidae]]: dwarf lemurs and mouse-lemurs (41 species) [135] => ***** Family [[Aye-aye|Daubentoniidae]]: aye-aye (1 species) [136] => ***** Family [[Lemuridae]]: ring-tailed lemur and allies (21 species) [137] => ***** Family [[Lepilemuridae]]: sportive lemurs (26 species) [138] => ***** Family [[Indriidae]]: woolly lemurs and allies (19 species) [139] => **** Superfamily [[Lorisoidea]] [140] => ***** Family [[Lorisidae]]: lorisids (16 species) [141] => ***** Family [[Galagidae]]: galagos (23 species) [142] => ** '''Suborder [[Haplorhini]]''': tarsiers, monkeys and apes [143] => *** Infraorder [[Tarsiiformes]] [144] => **** Family [[tarsier|Tarsiidae]]: tarsiers (14 species) [145] => *** Infraorder [[Simiiformes]] (or Anthropoidea) [146] => **** Parvorder [[Platyrrhini]]: New World monkeys [147] => ***** Family [[Callitrichidae]]: marmosets and tamarins (49 species) [148] => ***** Family [[Cebidae]]: capuchins and squirrel monkeys (29 species) [149] => ***** Family [[Aotidae]]: night or owl monkeys (douroucoulis) (11 species) [150] => ***** Family [[Pitheciidae]]: titis, sakis and uakaris (56 species) [151] => ***** Family [[Atelidae]]: howler, spider, woolly spider and woolly monkeys (26 species) [152] => **** Parvorder [[Catarrhini]] [153] => ***** Superfamily [[Old World monkey|Cercopithecoidea]] [154] => ****** Family [[Old World monkey|Cercopithecidae]]: Old World monkeys (165 species) [155] => ***** Superfamily [[Hominoidea]] [156] => ****** Family [[Hylobatidae]]: gibbons or "lesser apes" (20 species) [157] => ****** Family [[Hominidae]]: great apes, including humans (8 species) [158] => [159] => [160] => Order Primates was established by [[Carl Linnaeus]] in 1758, in the [[10th edition of Systema Naturae|tenth edition]] of his book ''[[Systema Naturae]]'',{{cite book | author = Linnaeus, C. | year = 1758 | title = Sistema naturae per regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonimis locis. Tomus I | publisher = Impensis direct. Laurentii Salvii, Holmia | pages = 20–32}} for the genera ''[[Homo]]'' (humans), ''[[Simia]]'' (other apes and monkeys), ''[[Lemur (genus)|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).{{cite book | author = Linnaeus, C. | year = 1735 | title = Sistema naturae sive regna tria Naturae systematice proposita per classes, ordines, genera, & species| publisher = apud Theodorum Haak, Lugduni Batavorum | pages = s.p}} In 1839, [[Henri Marie Ducrotay de Blainville]], following Linnaeus and aping his nomenclature, established the orders [[Secundates]] (including the suborders [[Bat|Chiroptera]], [[Insectivora]] and [[Carnivora]]), [[Tertiates]] (or [[Glires]]) and [[Quaternates]] (including [[Gravigrada]], [[Pachydermata]] and [[Ruminantia]]),{{cite book | author = Blainville, H. | year = 1839 | chapter = Nouvelle classification des Mammifères | title = Annales Françaises et Etrangères d'Anatomie et de Physiologie Appliquées à la Médicine et à l'Histoire Naturelle, 3| pages = 268–269}} but these new taxa were not accepted. [161] => [162] => Before Anderson and Jones introduced the classification of Strepsirrhini and Haplorhini in 1984,{{cite book |author1=Thorington, R. W. |author2=Anderson, S. |name-list-style=amp |year=1984 |chapter=Primates |pages=[https://archive.org/details/ordersfamiliesof0000unse/page/187 187–217] |editor=Anderson, S. |editor2=Jones, J. K. |title=Orders and Families of Recent Mammals of the World |publisher=John Wiley and Sons |location=New York |isbn=978-0-471-08493-8 |chapter-url=https://archive.org/details/ordersfamiliesof0000unse/page/187 }} (followed by McKenna and Bell's 1997 work [[Mammal classification#McKenna/Bell classification|''Classification of Mammals: Above the species level'']]),{{cite book|author1=McKenna, M. C. |author2=Bell, S. K. |name-list-style=amp |year=1997 |title=Classification of Mammals: Above the species level |publisher=Columbia University Press |location=New York |pages=631 |isbn=0-231-11013-8}} Primates was divided into two superfamilies: [[Prosimii]] and [[Anthropoidea]].{{cite book|title=Primate Behavioral Ecology|edition=Third|author=Strier, K.|author-link=Karen B. Strier|year=2007|isbn=978-0-205-44432-8 |pages=50–53|publisher=Pearson Allyn and Bacon }} Prosimii included all of the [[prosimian]]s: Strepsirrhini plus the [[tarsier]]s. Anthropoidea contained all of the [[simian]]s. [163] => [164] => ==Phylogeny and genetics== [165] => {{cladogram|clades={{Clade|style=font-size:85%; width:35em; [166] => |label1=[[Euarchontoglires]]   [167] => |1={{Clade [168] => |label1=[[Glires]]  [169] => |1={{Clade [170] => |1=[[Rodent]]ia (rodents) [171] => |2=[[Lagomorpha]] (rabbits, hares, pikas)}} [172] => |label2= [[Euarchonta]]  [173] => |2={{Clade [174] => |1=[[Treeshrew|Scandentia]] (treeshrews) [175] => | label2=[[Primatomorpha]] [176] => |2={{Clade [177] => |1=[[Colugo|Dermoptera]] (colugos) [178] => |label2='''Primates''' [179] => |2={{Clade [180] => |state1=double [181] => |1=†[[Plesiadapiformes]] [182] => |2=crown primates}} [183] => }} [184] => }} [185] => }} [186] => }}}} [187] => [[Order (biology)|Order]] Primates is part of the clade [[Euarchontoglires]], which is nested within the clade [[Eutheria]] of Class [[Mammal]]ia. Recent molecular genetic research on primates, [[colugo]]s, and [[treeshrew]]s has shown that the two species of colugos are more closely related to primates than to treeshrews,{{cite journal |title=Molecular and Genomic Data Identify the Closest Living Relative of Primates |journal=Science |date=2 November 2007 |last1=Janečka |first1=J. E. |last2=Miller |first2=W. |last3=Pringle |first3=T. H. |last4=Wiens |first4=F. |last5=Zitzmann |first5=A. |last6=Helgen |first6=K. M. |last7=Springer |first7=M. S. |last8=Murphy |first8=W. J. |doi=10.1126/science.1147555 |volume=318 |issue=5851 |pages=792–794 |pmid=17975064 |bibcode=2007Sci...318..792J|s2cid=12251814 }} even though treeshrews were at one time considered primates.{{cite book|title=A Complete Guide to Monkeys, Apes and Other Primates|url=https://archive.org/details/completeguidetom0000kava_z0w7|url-access=registration|author=Kavanagh, M. |publisher=Viking Press |location=New York|pages=[https://archive.org/details/completeguidetom0000kava_z0w7/page/18 18]|year=1983|isbn=0-670-43543-0}} These three orders make up the [[clade]] [[Euarchonta]]. The combination of this clade with the clade [[Glires]] (composed of [[Rodent]]ia 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.{{cite book |author1=McKenna, M. C. |author2=Bell, S. K. |name-list-style=amp | year = 1997 | title = Classification of Mammals Above the Species Level| publisher = Columbia University Press |location=New York| pages = 329 |isbn=0-231-11012-X}} [188] => [189] => ===Evolutionary history=== [190] => {{Further|Evolution of primates}} [191] => The primate lineage is thought to go back at least near the [[Cretaceous–Paleogene boundary]] or around 63–74 ([[mya (unit)|mya]]).{{cite journal | last1 = Williams | first1 = B.A. | last2 = Kay | first2 = R.F. | last3 = Kirk | first3 = E.C. | title = New perspectives on anthropoid origins | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 107 | pages = 4797–4804 | doi = 10.1073/pnas.0908320107 | issue = 11 | pmid = 20212104 | year = 2010 | pmc = 2841917 | bibcode=2010PNAS..107.4797W| doi-access = free }}{{cite journal |last1=Stanyon |first1=Roscoe |last2=Springer |first2=Mark S.|last3=Meredith|first3=Robert W. |last4=Gatesy |first4=John|last5=Emerling|first5=Christopher A. |last6=Park |first6=Jong|last7=Rabosky|first7=Daniel L. |last8=Stadler |first8=Tanja |last9=Steiner |first9=Cynthia|last10=Ryder|first10=Oliver A.|last11=Janečka|first11=Jan E. |last12=Fisher |first12=Colleen A. |last13=Murphy |first13=William J. |title=Macroevolutionary Dynamics and Historical Biogeography of Primate Diversification Inferred from a Species Supermatrix |journal=PLOS ONE |volume=7 |issue=11|year=2012 |pages=e49521 |issn=1932-6203 |doi=10.1371/journal.pone.0049521 |pmid=23166696 |pmc=3500307|bibcode=2012PLoSO...749521S|doi-access=free }}{{cite journal|last1=Jameson|first1=Natalie M. |last2=Hou |first2=Zhuo-Cheng |last3=Sterner |first3=Kirstin N. |last4=Weckle |first4=Amy |last5=Goodman |first5=Morris |last6=Steiper |first6=Michael E. |last7=Wildman |first7=Derek E. |title=Genomic data reject the hypothesis of a prosimian primate clade |journal=Journal of Human Evolution |volume=61 |issue=3|date=September 2011|pages=295–305 |issn=0047-2484 |doi=10.1016/j.jhevol.2011.04.004 |pmid=21620437}}{{cite journal |last1=Pozzi |first1=Luca |last2=Hodgson|first2=Jason A. |last3=Burrell |first3=Andrew S. |last4=Sterner |first4=Kirstin N.|last5=Raaum |first5=Ryan L.|last6=Disotell|first6=Todd R. |title=Primate phylogenetic relationships and divergence dates inferred from complete mitochondrial genomes|journal=Molecular Phylogenetics and Evolution |volume=75 |date=June 2014|pages=165–183|issn=1055-7903|doi=10.1016/j.ympev.2014.02.023 |pmid=24583291 |pmc=4059600}}{{cite journal |last1=Stanyon |first1=Roscoe |last2=Finstermeier |first2=Knut|last3=Zinner |first3=Dietmar|last4=Brameier|first4=Markus |last5=Meyer |first5=Matthias |last6=Kreuz |first6=Eva |last7=Hofreiter|first7=Michael|last8=Roos|first8=Christian|title=A Mitogenomic Phylogeny of Living Primates |journal=PLOS ONE |volume=8|issue=7|date=16 July 2013 |pages=e69504 |issn=1932-6203 |doi=10.1371/journal.pone.0069504 |pmid=23874967 |pmc=3713065 |bibcode=2013PLoSO...869504F|doi-access=free }} The earliest possible primate/proto-primate may be ''[[Purgatorius]]'', which dates back to [[Danian|Early Paleocene]] of North America ~66mya.{{Cite journal |last1=Wilson Mantilla |first1=Gregory P. |last2=Chester |first2=Stephen G. B. |last3=Clemens |first3=William A. |last4=Moore |first4=Jason R. |last5=Sprain |first5=Courtney J. |last6=Hovatter |first6=Brody T. |last7=Mitchell |first7=William S. |last8=Mans |first8=Wade W. |last9=Mundil |first9=Roland |last10=Renne |first10=Paul R. |title=Earliest Palaeocene purgatoriids and the initial radiation of stem primates |journal=Royal Society Open Science |year=2021 |volume=8 |issue=2 |pages=210050 |doi=10.1098/rsos.210050 |pmc=8074693 |pmid=33972886|bibcode=2021RSOS....810050W }} The oldest known primates from the fossil record date to the Late Paleocene of Africa, c.57 mya (''[[Altiatlasius]]''){{Cite journal | last1 = Williams | first1 = B. A. | last2 = Kay | first2 = R. F. | last3 = Kirk | first3 = E. C. | title = New perspectives on anthropoid origins | doi = 10.1073/pnas.0908320107 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 107 | issue = 11| pages = 4797–4804 | year = 2010 | pmid = 20212104 | pmc = 2841917|bibcode = 2010PNAS..107.4797W| doi-access = free }} or the Paleocene-Eocene transition in the northern continents, c. 55 mya (''[[Cantius]]'', ''[[Donrussellia]]'', ''[[Altanius]]'', ''[[Plesiadapis]]'' and ''[[Teilhardina]]'').{{Cite journal | last1 = Miller | first1 = E. R. | last2 = Gunnell | first2 = G. F. | last3 = Martin | first3 = R. D. | title = Deep Time and the Search for Anthropoid Origins | doi = 10.1002/ajpa.20352 | journal = American Journal of Physical Anthropology | volume = 128 | pages = 60–95 | year = 2005 | pmid = 16369958| url = http://www.paleontology.lsa.umich.edu/Accomplishments/deeptime.ajpa2005.pdf }}{{Cite journal|date=27 October 2009 |first1=Helen J |last1=Chatterjee |first2=Simon Y.W. |last2=Ho |first3=Ian |last3=Barnes |first4=Colin |last4=Groves |author-link4=Colin Groves |title=Estimating the phylogeny and divergence times of primates using a supermatrix approach |journal=BMC Evolutionary Biology |volume= 9 |doi=10.1186/1471-2148-9-259|pmid=19860891|pages=259|pmc=2774700 |issue=1 |doi-access=free }}{{cite journal|author=O'Leary, M. A.|date=8 February 2013|title= The placental mammal ancestor and the post–K-Pg radiation of placentals|journal=Science|volume=339|issue=6120|pages=662–667|doi=10.1126/science.1229237|pmid=23393258|display-authors=etal |bibcode=2013Sci...339..662O|hdl=11336/7302 |s2cid=206544776|hdl-access=free}} 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.{{cite journal|title=Molecular Clock Calibrations and Metazoan Divergence Dates|url=https://archive.org/details/sim_journal-of-molecular-evolution_1999-09_49_3/page/385|author=Lee, M.|journal=Journal of Molecular Evolution|volume=49|issue=3|date=September 1999 |pages=385–391|doi=10.1007/PL00006562|pmid=10473780|bibcode=1999JMolE..49..385L|s2cid=1629316}}{{cite web|title=Scientists Push Back Primate Origins From 65 Million To 85 Million Years Ago|url=https://www.sciencedaily.com/releases/2002/04/020418073440.htm|website=Science Daily|access-date=2008-10-24}}{{cite journal|title=Using the fossil record to estimate the age of the last common ancestor of extant primates|url=https://archive.org/details/sim_nature-uk_2002-04-18_416_6882/page/726|author1=Tavaré, S. |author2=Marshall, C. R. |author3=Will, O. |author4=Soligo, C. |author5=Martin R.D.|journal=Nature|date=April 18, 2002|pages=726–729 |volume=416|issue=6882 |doi=10.1038/416726a |pmid=11961552|bibcode = 2002Natur.416..726T|s2cid=4368374 }} [192] => [193] => By modern [[Cladistics|cladistic]] reckoning, the order Primates is [[monophyletic]]. The suborder [[Strepsirrhini]], the "[[rhinarium|wet-nosed]]" primates, is generally thought to have split off from the primitive primate line about 63 mya,{{cite journal |title=Molecular Remodeling of Members of the Relaxin Family During Primate Evolution |journal=Molecular Biology and Evolution|volume=18 |pages=393–403 |author1=Klonisch, T. |author2=Froehlich, C. |author3=Tetens, F. |author4=Fischer, B. |author5=Hombach-Klonisch, S. |year=2001 |pmid=11230540|issue=3 |doi=10.1093/oxfordjournals.molbev.a003815|doi-access=free }} although earlier dates are also supported.{{cite journal |title=Development and Application of a Phylogenomic Toolkit: Resolving the Evolutionary History of Madagascar's Lemurs |journal=Genome Research|volume=18 |pages=489–499 |year=2008 |author=Horvath, J.|doi=10.1101/gr.7265208 |pmid=18245770 |issue=3 |pmc=2259113 |display-authors=1 |last2=Weisrock|first2=D. W.|last3=Embry|first3=S. L.|last4=Fiorentino|first4=I. |last5=Balhoff |first5=J. P.|last6=Kappeler|first6=P.|last7=Wray|first7=G. A.|last8=Willard|first8=H. F.|last9=Yoder |first9=A. D.}} The seven strepsirrhine families are the five related [[lemur]] families and the two remaining families that include the [[Lorisidae|lorisids]] and the [[galago]]s.{{cite journal |title=Lemur Diversity in Madagascar |author-link1=Russell Mittermeier |author1=Mittermeier, R. |author2=Ganzhorn, J. |author3=Konstant, W. |author4=Glander, K. |author5=Tattersall, I. |author6=Groves, C. |author7=Rylands, A. |author8=Hapke, A. |author9=Ratsimbazafy, J. |author10=Mayor, M. |author11=Louis, E. |author12=Rumpler, Y. |author13=Schwitzer, C. |author14=Rasoloarison, R. |journal=International Journal of Primatology |doi=10.1007/s10764-008-9317-y |pages=1607–1656 |volume=29 |issue=6 |date=December 2008 |s2cid=17614597 |url=https://dukespace.lib.duke.edu/dspace/bitstream/10161/6237/1/08%20lemur%20diversity.pdf |access-date=2019-09-24 |archive-date=2021-02-15 |archive-url=https://web.archive.org/web/20210215163911/https://dukespace.lib.duke.edu/dspace/bitstream/handle/10161/6237/08 |url-status=dead }} Older classification schemes wrap [[Sportive lemur|Lepilemuridae]] into [[Lemuridae]] and [[Galago|Galagidae]] into [[Lorisidae]], yielding a four-one family distribution instead of five-two as presented here. During the [[Eocene]], most of the northern continents were dominated by two groups, the [[adapiform]]s and the [[omomyid]]s.{{cite book|title=Primates in Perspective|author=Hartwig, W.|chapter=Primate Evolution|editor=Campbell, C. |editor2=Fuentes, A. |editor3=MacKinnon, K. |editor4=Panger, M. |editor5=Bearder, S.|year=2007|publisher=Oxford University Press|isbn=978-0-19-517133-4|pages=13–17}} The former are considered members of Strepsirrhini, but did not have a [[toothcomb]] like modern lemurs; recent analysis has demonstrated that ''[[Darwinius|Darwinius masillae]]'' fits into this grouping.{{Cite journal | last1 = Williams | first1 = B. A. | last2 = Kay | first2 = R. F. | last3 = Christopher Kirk | first3 = E. | last4 = Ross | first4 = C. F. | title = ''Darwinius masillae'' is a strepsirrhine—a reply to Franzen ''et al.'' (2009) | doi = 10.1016/j.jhevol.2010.01.003 | journal = Journal of Human Evolution | volume = 59 | issue = 5 | pages = 567–573; discussion 573–9 | year = 2010 | pmid = 20188396 | url = http://strainlab.uchicago.edu/publications/Williams%20et%20al%202010.pdf | archive-url = https://web.archive.org/web/20130517155919/http://strainlab.uchicago.edu/publications/Williams%20et%20al%202010.pdf | archive-date = 2013-05-17 }} 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.{{cite book|title=Primate Evolution and Human Origins|author1=Ciochon, R. |author2=Fleagle, J. |name-list-style=amp |location=Menlo Park, California|publisher=Benjamin/Cummings|year=1987|isbn= 978-0-202-01175-2|pages=72}} [194] => [195] => 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.{{cite book | last = Garbutt | first = N. | title = Mammals of Madagascar, A Complete Guide | publisher = A&C Black Publishers | year = 2007 | isbn = 978-0-300-12550-4 | pages = 85–86}} However, Madagascar split from Africa 160 mya and from India 90 mya.{{cite book | last1 = Mittermeier | first1 = R.A. | author-link = Russell Mittermeier | title = Lemurs of Madagascar | edition = 2nd | publisher = Conservation International | year = 2006 | isbn=1-881173-88-7 | pages = 23–26|display-authors=etal}} 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,{{cite web|url=http://homepage.mac.com/wis/Personal/lectures/human-origins/PrimateEvolution.pdf |title=Primate Evolution |access-date=2008-10-23 |last=Sellers |first=Bill |date=2000-10-20 |publisher=University of Edinburgh |pages=13–17 |url-status=dead |archive-url=https://web.archive.org/web/20081029184647/http://homepage.mac.com/wis/Personal/lectures/human-origins/PrimateEvolution.pdf |archive-date=2008-10-29}} but none are supported by the genetic and molecular evidence. [196] => [[File:Brown Lemur in Andasibe.jpg|right|thumb|[[Common brown lemur]], a [[Strepsirrhini|Strepsirrhine]] primate]] [197] => 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. [198] => [199] => 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 (phylogenetics)|basal]] division, originating about 58 mya.{{cite conference|title=Evolutionary Biology of Tarsiers |author=Shekelle, M. |year=2005 |url=http://rmbr.nus.edu.sg/bejc/ |access-date=2008-08-22 |url-status=dead |archive-url=https://web.archive.org/web/20080907073204/http://rmbr.nus.edu.sg/bejc/ |archive-date=2008-09-07}}{{cite journal |title=Rapid electrostatic evolution at the binding site for cytochrome c on cytochrome c oxidase in anthropoid primates |author=Schmidt, T. |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=102 |issue=18|pages=6379–6384 |doi=10.1073/pnas.0409714102 |date=3 May 2005 |pmid=15851671 |pmc=1088365 |display-authors=1 |last2=Wildman |first2=DE |last3=Uddin |first3=M |last4=Opazo |first4=JC |last5=Goodman |first5=M |last6=Grossman |first6=LI |bibcode = 2005PNAS..102.6379S|doi-access=free }} The earliest known haplorhine skeleton, that of 55 MA old tarsier-like ''[[Archicebus]]'', was found in central China,{{cite web|last=Wade |first=Lizzie |title=Early Primate Weighed Less Than an Ounce |url=http://news.sciencemag.org/sciencenow/2013/06/crucial-link-in-primate-evolutio.html |publisher=[[ScienceNow]] |date=June 5, 2013 |access-date=2013-06-07 |url-status=dead |archive-url=https://web.archive.org/web/20130608230639/https://news.sciencemag.org/sciencenow/2013/06/crucial-link-in-primate-evolutio.html |archive-date=2013-06-08}} supporting an already suspected Asian origin for the group.{{cite journal|last1=Kay |first1=R. F. |title=Evidence for an Asian origin of stem anthropoid s|journal=Proceedings of the National Academy of Sciences of the United States of America |volume= 109|issue= 26 |date= 2012 |pages= 10132–10133 |doi=10.1073/pnas.1207933109 |bibcode = 2012PNAS..10910132K |pmid=22699505 |pmc=3387095|doi-access=free }} The infraorder [[Simian|Simiiformes]] (simian primates, consisting of monkeys and apes) emerged about 40 mya, possibly also in Asia; if so, they [[Oceanic dispersal|dispersed]] across the [[Tethys Ocean|Tethys Sea]] from Asia to Africa soon afterwards.{{Cite journal | last1 = Chaimanee | first1 = Y. | last2 = Chavasseau | first2 = O. | last3 = Beard | first3 = K. C. | last4 = Kyaw | first4 = A. A. | last5 = Soe | first5 = A. N. | last6 = Sein | first6 = C. | last7 = Lazzari | first7 = V. | last8 = Marivaux | first8 = L. | last9 = Marandat | first9 = B. | last10 = Swe | first10 = M. | last11 = Rugbumrung | first11 = M. | last12 = Lwin | first12 = T. | last13 = Valentin | first13 = X. | last14 = Zin-Maung-Maung-Thein | last15 = Jaeger | first15 = J. -J. | title = Late Middle Eocene primate from Myanmar and the initial anthropoid colonization of Africa | doi = 10.1073/pnas.1200644109 | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 109 | issue = 26 | pages = 10293–10297 | year = 2012 | pmid = 22665790| pmc = 3387043| bibcode = 2012PNAS..10910293C| doi-access = free }} There are two simian clades, both [[wikt:parvorder|parvorder]]s: [[Catarrhini]], which developed in Africa, consisting of [[Old World monkey]]s, humans and the other apes, and Platyrrhini, which developed in South America, consisting of [[New World monkey]]s. A third clade, which included the [[eosimiids]], developed in Asia, but became extinct millions of years ago.{{cite journal |author=Marivaux, L. |title=Anthropoid primates from the Oligocene of Pakistan (Bugti Hills): Data on early anthropoid evolution and biogeography|journal=Proceedings of the National Academy of Sciences of the United States of America |date=2005-06-14 |volume=102 |issue=24 |pages=8436–8441 |doi=10.1073/pnas.0503469102 |pmid= 15937103|pmc=1150860|display-authors=1|last2= Antoine |first2=PO |last3= Baqri |first3=SR|last4= Benammi |first4=M|last5= Chaimanee |first5=Y |last6=Crochet |first6=JY |last7=De Franceschi|first7= D|last8=Iqbal |first8= N|last9=Jaeger|first9= JJ|bibcode = 2005PNAS..102.8436M|doi-access=free}} [200] => [201] => 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.{{cite journal |author1=Schrago, C.G. |author2=Russo, C.A.M. |name-list-style=amp | title = Timing the Origin of New World Monkeys | journal = Molecular Biology and Evolution | volume = 20 | issue = 10 | year = 2003 | pages = 1620–1625 | format = PDF Reprint | doi = 10.1093/molbev/msg172 | pmid = 12832653| doi-access = free }} The anthropoid primates possibly traversed the Atlantic Ocean from Africa to South America during the [[Eocene]] by [[island hopping]], facilitated by [[Mid-Atlantic Ridge|Atlantic Ocean ridges]] and a lowered sea level. Alternatively, a single [[Oceanic dispersal|rafting event]] may explain this transoceanic colonization. Due to [[continental drift]], the Atlantic Ocean was not nearly as wide at the time as it is today. Research suggests that a small {{convert|1|kg|lb|abbr=on}} primate could have survived 13 days on a raft of vegetation.{{cite journal | author = Houle, A. | title = The origin of platyrrhines: An evaluation of the Antarctic scenario and the floating island model | url = https://archive.org/details/sim_american-journal-of-physical-anthropology_1999-08_109_4/page/541 | journal = [[American Journal of Physical Anthropology]] | volume = 109 | issue = 4 | year = 1999 | pages = 541–559 | doi = 10.1002/(SICI)1096-8644(199908)109:4<541::AID-AJPA9>3.0.CO;2-N | pmid = 10423268}} Given estimated current and wind speeds, this would have provided enough time to make the voyage between the continents. [202] => [[File:Tamarin portrait 2 edit3.jpg|thumb|[[Emperor tamarin]], a [[New World monkey]]]] [203] => Apes and monkeys spread from Africa into Europe and Asia starting in the [[Miocene]].{{cite journal |author1=Andrews, P. |author2=Kelley, J. |name-list-style=amp | title = Middle Miocene Dispersals of Apes |journal=Folia Primatologica | volume = 78 | year = 2007 | pages = 328–343 | doi = 10.1159/000105148 | pmid = 17855786 | issue = 5–6|s2cid=19293586 }} 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.{{cite book|title=Primate Behavioral Ecology |author=Strier, K. |publisher = [[Allyn & Bacon]] | edition = 3rd|year=2007|isbn=978-0-205-44432-8|pages=7, 64, 71, 77, 182–185, 273–280, 284, 287–298}} Molecular and fossil studies generally show that modern humans originated in Africa 100,000–200,000 years ago.{{cite book |author1=Pough, F. W. |author2=Janis, C. M. |author3=Heiser, J. B. |title=Vertebrate Life |url=https://archive.org/details/vertebratelife0000poug |chapter=Primate Evolution and the Emergence of Humans |year=2005 |orig-year=1979 |edition=7th |publisher= Pearson |pages=[https://archive.org/details/vertebratelife0000poug/page/650 650] |isbn=0-13-127836-3}} [204] => [205] => Although primates are well studied in comparison to other animal groups, several new species have [[Primates discovered in the 2000s|been discovered recently]], and genetic tests have revealed previously unrecognised species in known populations. ''Primate Taxonomy'' listed about 350 species of primates in 2001;{{cite book |title=Primate Taxonomy |publisher=Smithsonian Institution Press |year=2001 |author=Groves, C. P. |isbn=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 522 species, or 708 including subspecies.{{cite web |title=Primate diversity by region |author=IUCN/SSC Primate Specialist Group |date=1 March 2021 |publisher=International Union for the Conservation of Nature |url=http://www.primate-sg.org/primate_diversity_by_region/}} [206] => [207] => ===Hybrids=== [208] => Primate [[Hybrid (biology)|hybrids]] usually arise in captivity,{{cite journal |title=Songs of hybrid gibbons (''Hylobates lar'' × ''H. muelleri'') |author=Tenaza, R. |year=1984 |journal=American Journal of Primatology |volume=8 |issue=3 |pages=249–253 |doi=10.1002/ajp.1350080307|pmid=31986810 |s2cid=84957700 }} but there have also been examples in the wild.{{cite journal |year=1966 |title=Naturally occurring primate hybrid |author=Bernsteil, I. S. |journal=Science |volume=154 |issue=3756 |pages=1559–1560 |doi=10.1126/science.154.3756.1559 |pmid=4958933|bibcode = 1966Sci...154.1559B|s2cid=85898043 }}{{cite journal |author=Sugawara, K. |title=Sociological study of a wild group of hybrid baboons between ''Papio anubis'' and ''P. hamadryas'' in the Awash Valley, Ethiopia |date=January 1979 |journal=Primates |volume=20 |issue=1 |doi=10.1007/BF02373827 |pages=21–56|s2cid=23061688 }} Hybridization occurs where two species' range overlap to form [[hybrid zone]]s; 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]].{{cite journal |last1=Jolly |title=Intergeneric Hybrid Baboons |journal=International Journal of Primatology |year=1997 |volume=18 |issue=4 |doi=10.1023/A:1026367307470 |pages=597–627 |first1=C. J. |display-authors=2|last2=Woolley-Barker|first2=Tamsin |last3=Beyene |first3=Shimelis |last4=Disotell |first4=Todd R. |last5=Phillips-Conroy |first5=Jane E.|s2cid=27900830 }} [209] => [210] => ===Clones=== [211] => On 24 January 2018, scientists in China reported in the journal ''[[Cell (journal)|Cell]]'' the creation of two [[crab-eating macaque]] [[Clone (cell biology)|clone]]s, named ''[[Zhong Zhong and Hua Hua|Zhong Zhong]]'' and ''[[Zhong Zhong and Hua Hua|Hua Hua]]'', using the [[Nuclear transfer|complex DNA transfer method]] that produced [[Dolly (sheep)|''Dolly'' the sheep]], for the first time.{{cite journal |last=Liu |first=Zhen |display-authors=etal |title=Cloning of Macaque Monkeys by Somatic Cell Nuclear Transfer |date=24 January 2018 |journal=[[Cell (journal)|Cell]] |doi=10.1016/j.cell.2018.01.020 |pmid=29395327 |volume=172 |issue=4 |pages=881–887.e7|doi-access=free }}{{cite journal |last=Normile |first=Dennis |title=These monkey twins are the first primate clones made by the method that developed Dolly |url=https://www.science.org/content/article/these-monkey-twins-are-first-primate-clones-made-method-developed-dolly |date=24 January 2018 |journal=[[Science (journal)|Science]] |doi=10.1126/science.aat1066 |access-date=24 January 2018}}{{cite journal |last=Cyranoski |first=David |title=First monkeys cloned with technique that made Dolly the sheep - Chinese scientists create cloned primates that could revolutionize studies of human disease. |date=24 January 2018 |journal=[[Nature (journal)|Nature]] |volume=553 |issue=7689 |pages=387–388 |doi=10.1038/d41586-018-01027-z|pmid=29368720 |bibcode=2018Natur.553..387C|doi-access=free }}{{cite news |last=Briggs |first=Helen |title=First monkey clones created in Chinese laboratory |url=https://www.bbc.com/news/health-42809445 |date=24 January 2018 |work=[[BBC News]] |access-date=24 January 2018}}{{cite news |agency=Associated Press |title=Scientists Successfully Clone Monkeys; Are Humans Up Next? |url=https://www.nytimes.com/aponline/2018/01/24/science/ap-us-sci-cloned-monkeys.html |date=24 January 2018 |work=[[The New York Times]] |access-date=24 January 2018}} [212] => [213] => ==Anatomy and physiology== [214] => [215] => ===Head=== [216] => [[File:Primate skull series with legend cropped.png|thumb|Primate skulls showing [[postorbital bar]], and increasing brain sizes]] [217] => The primate skull has a large, domed [[Skull|cranium]], which is particularly prominent in [[Simian|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 [[human]]s 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 [[gorilla]]s, 400 cm3 in [[chimpanzee]]s and 397 cm3 in [[orangutan]]s.{{cite book |author1=Aiello, L. |author2=Dean, C. |name-list-style=amp | year=1990 | title=An Introduction to Human Evolutionary Anatomy |url=https://archive.org/details/introductiontohu00aiel | publisher=Academic Press | pages=[https://archive.org/details/introductiontohu00aiel/page/193 193] | isbn=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 [[sense|sensory perception]], generation of [[motor cortex|motor commands]], spatial reasoning, [[consciousness|conscious thought]] and, in humans, [[language]]. While other mammals rely heavily on their [[Olfaction|sense of smell]], the arboreal life of primates has led to a [[Somatosensory system|tactile]], [[Visual perception|visually]] dominant sensory system, a reduction in the olfactory region of the brain and increasingly complex social behavior.{{cite web | url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Primates.html | title="Primates" (On-line) |publisher=Animal Diversity Web | author=Myers, P. | year=1999 | access-date=2008-06-03}} The visual acuity of [[human]]s and other [[hominid]]s is exceptional; they have the most [[Spatial resolution|acute vision]] known among all vertebrates, with the exception of certain species of [[Bird of prey|predatory birds]].{{Cite journal |last=Caves |first=Eleanor M. |date=May 2018 |title=Visual Acuity and the Evolution of Signals |journal=Trends in Ecology & Evolution |volume=33 |issue=5 |pages=358–372 |doi=10.1016/j.tree.2018.03.001 |pmid=29609907 |url=https://www.cell.com/trends/ecology-evolution/fulltext/S0169-5347(18)30052-1 |access-date=29 July 2018}}{{Citation |last1=Kirk |first1=E. Christopher |title=The Evolution of High Visual Acuity in the Anthropoidea |date=2004 |url=https://doi.org/10.1007/978-1-4419-8873-7_20 |work=Anthropoid Origins: New Visions |pages=539–602 |editor-last=Ross |editor-first=Callum F. |access-date=2023-07-30 |series=Developments in Primatology: Progress and Prospects |place=Boston, MA |publisher=Springer US |language=en |doi=10.1007/978-1-4419-8873-7_20 |isbn=978-1-4419-8873-7 |last2=Kay |first2=Richard F. |editor2-last=Kay |editor2-first=Richard F.}} [218] => [219] => Primates have forward-facing eyes on the front of the skull; [[binocular vision]] allows accurate distance perception, useful for the [[Brachiation|brachiating]] ancestors of all great apes. A [[supraorbital ridge|bony ridge]] above the eye sockets reinforces weaker bones in the face, which are put under strain during chewing. [[Strepsirrhini|Strepsirrhines]] have a [[postorbital bar]], a bone around the eye socket, to protect their eyes; in contrast, the higher primates, [[Haplorhini|haplorhines]], have evolved fully enclosed sockets.{{cite book |author1=Campbell, B. G. |author2=Loy, J. D. |name-list-style=amp | year = 2000 | title = Humankind Emerging | publisher = Allyn & Bacon | pages = 85 |isbn=0-673-52364-0|edition=8th }} [220] => [[File:PrimateFeet.jpg|thumb|right|upright|An 1893 drawing of the hands and feet of various primates]] [221] => Primates show an evolutionary trend towards a reduced [[snout]].{{cite web |author1=White, T. |author2=Kazlev, A. |name-list-style=amp |url=http://www.palaeos.com/Vertebrates/Units/480Archonta/480.400.html#Primates | title=Archonta: Primates |publisher=[[Palaeos]]| access-date=2008-06-03 | date=2006-01-08 |archive-url=https://web.archive.org/web/20080512022901/http://www.palaeos.com/Vertebrates/Units/480Archonta/480.400.html |archive-date=2008-05-12}} Technically, Old World monkeys are distinguished from New World monkeys by the structure of the nose, and from apes by the [[dentition|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 [[incisor]]s, 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 [[premolar]]s, compared with 12 in New World monkeys. The Old World species are divided into apes and monkeys depending on the number of [[Cusp (dentistry)|cusps]] on their [[Molar (tooth)|molars]]: monkeys have four, apes have five - although humans may have four or five.{{cite book |title=Wheeler's Dental Anatomy, Physiology, and Occlusion |last1=Ash |first1=M. M. |last2=Nelson |first2=S. J. |last3=Wheeler |first3=R. C. |publisher=W.B. Saunders |year=2003 |isbn=978-0-7216-9382-8 |page=[https://archive.org/details/dentalanatomyphy00majo/page/12 12] |url=https://archive.org/details/dentalanatomyphy00majo/page/12}} 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. [222] => [223] => ===Body=== [224] => [[File:Chlorocebus pygerythrus01.jpg|thumb|Vervet hindfoot showing fingerprint ridges on the sole]] [225] => Primates generally have five digits on each limb ([[Dactyly#Pentadactyly|pentadactyly]]), with a characteristic type of keratin [[Nail (anatomy)|fingernail]] on the end of each finger and toe. The bottom sides of the hands and feet have [[tactile pad|sensitive pads]] on the [[distal phalanges|fingertips]]. Most have [[thumb|opposable thumbs]], a characteristic primate feature most developed in [[human]]s, though not limited to this order ([[opossum]]s and [[koala]]s, for example, also have them). Thumbs allow some species to use [[Tool use by animals|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-claw]]s, which they use for grooming.{{cite book |author1=Pough, F. W. |author2=Janis, C. M. |author3=Heiser, J. B. |title=Vertebrate Life |url=https://archive.org/details/vertebratelife0000poug |chapter=Characteristics of Primates |year=2005 |orig-year=1979 |edition=7th |publisher= Pearson |pages=[https://archive.org/details/vertebratelife0000poug/page/630 630] |isbn=0-13-127836-3}} [226] => [227] => The primate [[clavicle|collar bone]] is a prominent element of the [[pectoral girdle]]; this allows the [[Glenohumeral joint|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 [[vertebra]]e greatly reduced - resulting in tail loss in some species. [[Prehensile tail]]s are found in the New World [[Atelidae|atelids]], including the [[howler monkey|howler]], [[spider monkey|spider]], [[woolly spider monkey|woolly spider]], [[woolly monkey]]s; and in [[Capuchin monkey|capuchin]]s.{{cite journal |vauthors=Garber PA, Rehg JA |title=The ecological role of the prehensile tail in white-faced capuchins (Cebus capucinus) |url=https://archive.org/details/sim_american-journal-of-physical-anthropology_1999-11_110_3/page/325 |journal=American Journal of Physical Anthropology |volume=110 |issue=3 |pages=325–39 | date=November 1999 |pmid=10516564 |doi=10.1002/(SICI)1096-8644(199911)110:3<325::AID-AJPA5>3.0.CO;2-D}}{{cite journal |vauthors=Russo GA, Young JW |title=Tail growth tracks the ontogeny of prehensile tail use in capuchin monkeys (Cebus albifrons and C. apella) |url=https://archive.org/details/sim_american-journal-of-physical-anthropology_2011-11_146_3/page/465 |journal=American Journal of Physical Anthropology |volume=146 |issue=3 |pages=465–73 | date=November 2011 |pmid=21953012 |doi=10.1002/ajpa.21617}} Male primates have a [[Penis#Primates|low-hanging penis]] and testes descended into a scrotum.{{cite book|author=Friderun Ankel-Simons|title=Primate Anatomy: An Introduction|url=https://books.google.com/books?id=Mwl3M6c5KzoC|date=27 July 2010|publisher=Academic Press|isbn=978-0-08-046911-9|pages=442, 521 }}{{cite encyclopedia | last = Macdonald | first = David | author-link = David W. Macdonald | title = Primates | encyclopedia = The Encyclopedia of Mammals | pages = 282–307 | publisher = The Brown Reference Group plc | year = 2006 | isbn = 0-681-45659-0}} [228] => [229] => ===Sexual dimorphism=== [230] => {{Main|Sexual dimorphism in non-human primates}} [231] => [[File:Silverback.JPG|thumb|right|Distinct sexual size dimorphism can be seen between the male and female mountain gorilla.]] [232] => [[Sexual dimorphism]] is often exhibited in [[simian]]s, 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 weight|body mass]]{{cite journal |author=Ralls, K. |year=1976 |title=Mammals in Which Females are Larger Than Males |journal=The Quarterly Review of Biology |volume=51 |issue=2 |doi=10.1086/409310 | pages=245–76 |pmid=785524|s2cid=25927323 }}{{cite journal | author=Lindstedtand & Boyce | date=July 1985 | journal=The American Naturalist | volume=125 |pages=873 |doi = 10.1086/284385 |title = Seasonality, Fasting Endurance, and Body Size in Mammals | last2=Boyce | first2=Mark S. | issue=6| s2cid=84308684 }} and canine tooth size{{cite journal |author=Frisch, J. E. |year=1963 |title=Sex-differences in the canines of the gibbon (''Hylobates lar'') |journal = Primates |volume = 4 |issue = 2 |doi=10.1007/BF01659148 |pages=1–10|s2cid=189798134 }}{{cite journal |author=Kay, R. F. |year=1975 |title=The functional adaptations of primate molar teeth |journal=American Journal of Physical Anthropology |volume=43 |issue=2 |pages=195–215 |doi=10.1002/ajpa.1330430207 |pmid=810034}} along with [[pelage]] and skin color.{{cite book |author=Crook, J. H. |editor=Campbell, B. G. |year=1972 |chapter=Sexual selection, dimorphism, and social organization in the primates |title=Sexual selection and the descent of man |publisher=Aldine Transaction |pages=[https://archive.org/details/sexualselection00camp/page/246 246] |isbn=978-0-202-02005-1 |chapter-url=https://archive.org/details/sexualselection00camp/page/246}} The dimorphism can be attributed to and affected by different factors, including [[mating system]],{{cite journal |last1=Cheverud |first1=J. M. |last2=Dow |first2=M. M. |last3=Leutenegger |first3=W. |date=November 1985 |title=The quantitative assessment of phylogenetic constraints in comparative analyses: Sexual dimorphism in body weight among primates |url=https://archive.org/details/sim_evolution_1985-11_39_6/page/1335 |journal=Evolution |volume=39 |issue=6 |pages=1335–1351 |doi=10.2307/2408790 |pmid=28564267 |jstor=2408790}} size, habitat and diet.{{cite journal |last1=Leutenegger |first1=W. |last2=Cheverud |first2=J. M. | year=1982 | title=Correlates of sexual dimorphism in primates: Ecological and size variables | journal = International Journal of Primatology | volume = 3 | issue = 4 | doi=10.1007/BF02693740 | pages=387–402|s2cid=38220186 | doi-access=free }} [233] => [234] => 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.{{cite journal |author=Plavcan, J. M. |year=2001 |title=Sexual dimorphism in primate evolution |journal=American Journal of Physical Anthropology |volume=33 |pages=25–53 |doi=10.1002/ajpa.10011 |pmid=11786990|s2cid=31722173 |doi-access=free }} Ontogenetic scaling, where relative extension of a common growth trajectory occurs, may give some insight into the relationship between sexual dimorphism and growth patterns.{{cite journal |last1=O'Higgins |first1=P. |last2=Collard |first2=M. |year=2002 |title=Sexual dimorphism and facial growth in papionine monkeys |journal=Journal of Zoology |volume=257 |issue=2 |pages=255–72 |doi=10.1017/S0952836902000857}} Some evidence from the fossil record suggests that there was [[convergent evolution]] of dimorphism, and some extinct [[hominid]]s probably had greater dimorphism than any living primate. [235] => [236] => ===Locomotion=== [237] => [[File:Diademed ready to push off.jpg|thumb|[[Diademed sifaka]], a lemur that is a vertical clinger and leaper]] [238] => Primate species move by [[brachiation]], [[bipedalism]], [[Jumping|leaping]], arboreal and terrestrial [[quadruped]]alism, [[arboreal locomotion|climbing]], [[knuckle-walking]] or by a combination of these methods. Several prosimians are primarily vertical clingers and leapers. These include many [[Galago|bushbabies]], all [[Indriidae|indriids]] (i.e., [[sifaka]]s, [[avahi (genus)|avahis]] and [[indri]]s), [[sportive lemur]]s, and all [[tarsier]]s.{{cite book|title=Primate Ecology and Social Structure Volume 1: Lorises, Lemurs and Tarsiers|author=Sussman, R. W.|year=1999|isbn=0-536-02256-9|location=Needham Heights, MA|publisher=Pearson Custom Publishing & Prentice Hall|pages=[https://archive.org/details/primateecologyso0001suss/page/78 78, 89–90, 108, 121–123, 233]|url=https://archive.org/details/primateecologyso0001suss/page/78}} 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. [[Gibbon]]s, [[muriqui]]s and [[spider monkey]]s all brachiate extensively, with gibbons sometimes doing so in remarkably acrobatic fashion. [[Woolly monkey]]s also brachiate at times. [[Orangutan]]s 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. [[Chimpanzee]]s and [[gorilla]]s knuckle walk, and can move bipedally for short distances. Although numerous species, such as [[australopithecine]]s and [[Homo|early hominids]], have exhibited fully bipedal locomotion, humans are the only extant species with this trait.{{Cite book | first1 = S. D. | last1 = Glazier | first2 = C. A. | last2 = Flowerday | title = Selected Readings in the Anthropology of Religion: Theoretical and Methodological Essays | url = https://archive.org/details/selectedreadings00glaz | publisher = Greenwood Publishing Group | year = 2003 | isbn = 9780313300905 | pages = [https://archive.org/details/selectedreadings00glaz/page/53 53]}} [239] => [240] => ===Vision=== [241] => [[File:GarnettsGalago CincinnatiZoo.jpg|thumb|The ''[[tapetum lucidum]]'' of a [[northern greater galago]], typical of prosimians, reflects the light of the photographer's flash.]] [242] => The [[evolution of color vision in primates]] is unique among most [[eutheria]]n [[mammal]]s. While the remote [[vertebrate]] ancestors of the primates possessed [[trichromacy|three color vision]] (trichromaticism), the [[Nocturnality|nocturnal]], [[warm-blooded]], mammalian ancestors lost one of three cones in the [[retina]] during the [[Mesozoic]] era. Fish, [[reptile]]s and birds are therefore trichromatic or [[Tetrachromacy|tetrachromatic]], while all mammals, with the exception of some primates and [[marsupial]]s,{{cite journal | last1=Arrese | year=2005 | title=Cone topography and spectral sensitivity in two potentially trichromatic marsupials, the quokka (''Setonix brachyurus'') and quenda (''Isoodon obesulus'') | journal=Proceedings of the Royal Society B | volume=272 | issue=1565 | doi=10.1098/rspb.2004.3009 | pages=791–6 | pmid=15888411 | pmc=1599861 | first1=C. A. |last2=Oddy|first2=Alison Y. | display-authors=2 | last3=Runham | first3=P. B | last4=Hart | first4=N. S | last5=Shand | first5=J. | last6=Hunt | first6=D. M | last7=Beazley | first7=L. D}} are dichromats or [[Monochromacy|monochromats]] (totally color blind). Nocturnal primates, such as the [[night monkey]]s and [[Galago|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.{{cite journal |last1 = Bowmaker |first1=J. K. |last2=Astell |first2=S. |last3=Hunt |first3=D. M. |last4 = Mollon |first4=J. D. | year = 1991 | title = Photosensitive and photostable pigments in the retinae of Old World monkeys | journal = The Journal of Experimental Biology | volume = 156 | pages = 1–19 |issn=0022-0949 |url=http://jeb.biologists.org/cgi/reprint/156/1/1.pdf |access-date=2008-06-16 | pmid = 2051127 | issue=1|doi=10.1242/jeb.156.1.1 }} Platyrrhines, on the other hand, are trichromatic in a few cases only.{{cite journal |author1=Surridge, A. K. |author2=D. Osorio |name-list-style=amp | year = 2003 | title = Evolution and selection of trichromatic vision in primates | journal = Trends in Ecology and Evolution | volume = 18 | pages = 198–205 | doi = 10.1016/S0169-5347(03)00012-0 | issue = 4}} Specifically, individual females must be [[Zygosity#Heterozygous|heterozygous]] for two [[allele]]s of the opsin gene (red and green) located on the same [[Locus (genetics)|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. [243] => [244] => Like catarrhines, howler monkeys (a family of platyrrhines) show routine trichromatism that has been traced to an evolutionarily recent [[gene duplication]].{{cite journal |last1=Lucas |first1=P. W. |last2=Dominy |first2=N. J. |last3=Riba-Hernandez |first3=P. |last4=Stoner |first4=K. E. |last5=Yamashita |first5=N. |last6=Loría-Calderón |first6=E. |last7=Petersen-Pereira |first7=W. |last8=Rojas-Durán |first8=Y. |last9=Salas-Pena |first9=R. |last10=Solis-Madrigal |first10=S. |last11=Osorio |first11=D. |last12=Darvell |first12=B. W. | year = 2003 | title = Evolution and function of routine trichromatic vision in primates |url=https://archive.org/details/sim_evolution_2003-11_57_11/page/2636 | journal = Evolution | volume = 57 |pages = 2636–43 | doi = 10.1554/03-168 | pmid = 14686538 | issue = 11|s2cid=739130 }} Howler monkeys are one of the most specialized leaf-eaters of the New World monkeys; fruits are not a major part of their diets,{{cite book|title=Primate Ecology and Social Structure, Volume 2: New World Monkeys|edition=Revised First|author=Sussman, R. W.|year=2003|isbn=0-536-74364-9|location = Needham Heights, MA | publisher = Pearson Custom Publishing & Prentice Hall|pages=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. [245] => [246] => ==Behavior== [247] => [248] => ===Social systems=== [249] => [[Richard Wrangham]] stated that [[Social structure|social systems]] of primates are best classified by the amount of movement by females occurring between groups.{{cite book |author=Wrangham, R. W. |author-link=Richard Wrangham |chapter=Mutualism, kinship and social evolution |year=1982 |title=Current Problems in Sociobiology |publisher=Cambridge University Press |pages=269–89 |isbn=0-521-24203-7}} He proposed four categories: [250] => [251] => * Female transfer systems – 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 [[chimpanzee]]s, where the males, who are typically related, will cooperate in defense of the group's territory.{{cite journal |author=Goldberg, T. L. |author2=Wrangham, R. W. | title=Genetic correlates of social behavior in wild chimpanzees: evidence from mitochondrial DNA | journal=Animal Behaviour | volume=54 | issue=3 | pages=559–70 | date=September 1997 | pmid=9299041 | doi=10.1006/anbe.1996.0450|s2cid=18223362 }} Evidence of this social system has also been found among [[Neanderthal]] remains in [[Spain]]{{cite news | last = Bowdler | first = Neil | title = Neanderthal family found cannibalised in cave in Spain | url = https://www.bbc.co.uk/news/science-environment-12049854 | work = [[BBC News]] | date = 21 December 2010}} and in remains of ''[[Australopithecus]]'' and ''[[Paranthropus robustus]]'' groups in southern Africa.{{cite news | first=Neil | last=Bowdler | url=https://www.bbc.co.uk/news/science-environment-13609260 | title=Ancient cave women 'left childhood homes' | date=2 June 2011 | work=[[BBC News]] | access-date=2 June 2011}}{{cite journal | last1 = Copeland | first1 = Sandi R. |display-authors=etal|title=Strontium isotope evidence for landscape use by early hominins |volume=474 | issue = 7349 | date=1 June 2011 |pages=76–78 |journal=Nature |doi=10.1038/nature10149 |pmid=21637256| s2cid = 205225222 }} Among New World Monkeys, [[spider monkey]]s and [[muriqui]]s use this system.{{cite book|title=Primates in Perspective|author1=Fiore, A. D. |author2=Campbell, C. J. |name-list-style=amp |chapter=The Atelines|year=2007|page=175|publisher=Oxford University Press|editor=Campbell, C. J. |editor2=Fuentes, A. |editor3=MacKinnon, K. C. |editor4=Panger, M. |editor5=Bearder, S. K.|isbn=978-0-19-517133-4}} [252] => [[File:Ringstaartmakis - Ring-tailed Lemur.jpg|thumb|right|A social huddle of [[ring-tailed lemur]]s. The two individuals on the right exposing their white ventral surface are sunning themselves.]] [253] => * Male transfer systems – while the females remain in their natal groups, the males will emigrate as adolescents. [[Polygyny in nature|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 monkey]]s and [[Cercopithecinae|cercopithecine monkeys]]. [254] => * Monogamous species – 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. [[Gibbon]]s essentially use this system, although "monogamy" in this context does not necessarily mean absolute sexual fidelity.{{cite book|title=Primates in Perspective|author=Bartlett, T. Q.|chapter=The Hylobatidae|year=2007|editor=Campbell, C. J. |editor2=Fuentes, A. |editor3=MacKinnon, K. C. |editor4=Panger, M. |editor5=Bearder, S. K.|publisher=Oxford University Press|isbn=978-0-19-517133-4|page=283}} These species do not live in larger groups. [255] => * Solitary species – 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]].{{Cite thesis | degree = Ph.D. | title = Behavior and ecology of wild slow lorises (''Nycticebus coucang''): social organization, infant care system, and diet | url = http://deposit.ddb.de/cgi-bin/dokserv?idn=966021282&dok_var=d1&dok_ext=pdf&filename=966021282.pdf | last = Wiens | first = Frank | year = 2002 | publisher = Bayreuth University | url-status = dead | archive-url = https://web.archive.org/web/20120309212455/http://deposit.ddb.de/cgi-bin/dokserv?idn=966021282&dok_var=d1&dok_ext=pdf&filename=966021282.pdf |pages=31–32 |archive-date = 9 March 2012 | df = dmy-all }} [[Orangutan]]s do not defend their territory but effectively have this organization.{{cite book|title=Primates in Perspective|author1=Knott, C. D. |author2=Kahlenberg, S. M. |name-list-style=amp |chapter=Orangutans in Perspective|year=2007|editor=Campbell, C. J. |editor2=Fuentes, A. |editor3=MacKinnon, K. C. |editor4=Panger, M. |editor5=Bearder, S. K.|publisher=Oxford University Press|isbn=978-0-19-517133-4|page=294}} [256] => [257] => Other systems are known to occur as well. For example, with [[howler monkey]]s and [[gorilla]]s both the males and females typically transfer from their natal group on reaching sexual maturity, resulting in groups in which neither the males nor females are typically related.{{Cite book|author=Watts D. P.|year=1996|contribution=Comparative socio-ecology of gorillas|editor1=McGrew W. C. |editor2=Marchant L. F. |editor3=Nishida, T.|title=Great Ape Societies|url=https://archive.org/details/greatapesocietie00mcgr|location=Cambridge (England|publisher=Cambridge Univ Press|pages=[https://archive.org/details/greatapesocietie00mcgr/page/16 16]–28|isbn=978-0521555364}} Some prosimians, [[Colobinae|colobine]] monkeys and [[Callitrichinae|callitrichid]] monkeys also use this system. [258] => [259] => The transfer of females or males from their native group is likely an adaptation for avoiding inbreeding.{{cite journal |vauthors=Charpentier MJ, Widdig A, Alberts SC |title=Inbreeding depression in non-human primates: a historical review of methods used and empirical data |journal=American Journal of Primatology |volume=69 |issue=12 |pages=1370–86 |date=December 2007 |pmid=17486606 |doi=10.1002/ajp.20445 |s2cid=46626761 }} 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.{{cite journal | vauthors = Ralls K, Ballou J | year = 1982 | title = Effect of inbreeding on infant mortality in captive primates | doi = 10.1007/BF02693747 | journal = International Journal of Primatology | volume = 3 | issue = 4 | pages = 491–505 | s2cid = 10954608 | url = http://si-pddr.si.edu/dspace/bitstream/10088/6162/1/8133A167-A994-450A-ADF4-DAD3F4EAA6F6.pdf }}{{Dead link|date=October 2023 |bot=InternetArchiveBot |fix-attempted=yes }} This effect of inbreeding on infant mortality is probably largely a result of increased expression of deleterious recessive alleles (see [[Inbreeding depression]]). [260] => [[File:Three chimpanzees with apple.jpg|thumb|right|[[Chimpanzee]]s are social great apes.]] [261] => Primatologist [[Jane Goodall]], who studied in the [[Gombe Stream National Park]], noted [[fission-fusion society|fission-fusion societies]] in chimpanzees.{{cite journal |last1=Constable |first1=J. L. |last2=Ashley |first2=M. V. |last3=Goodall |first3=J. |last4=Pusey |first4=A. E. |title=Noninvasive paternity assignment in Gombe chimpanzees |url=https://archive.org/details/sim_molecular-ecology_2001-05_10_5/page/1279 |journal=Molecular Ecology |volume=10 |issue=5 |pages=1279–300 | date=May 2001 |pmid=11380884 |doi=10.1046/j.1365-294X.2001.01262.x|s2cid=46604532 }} 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]],{{cite book|title=The Pictorial Guide to the Living Primates|url=https://archive.org/details/pictorialguideto0000rowe|url-access=registration|author=Rowe, N.|year=1996|publisher=Pogonias Press|isbn=0-9648825-0-7|pages=[https://archive.org/details/pictorialguideto0000rowe/page/n15 4], 139, 143, 15 185, 223}} [[spider monkey]]s 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. [[Human]]s also form fission-fusion societies. In hunter-gatherer societies, humans form groups which are made up of several individuals that may split up to obtain different resources.{{Cite journal|last1=Couzin|first1=Iain D.|last2=Laidre|first2=Mark E.|date=August 2009|title=Fission–fusion populations|journal=Current Biology|volume=19|issue=15|pages=R633–R635|doi=10.1016/j.cub.2009.05.034|pmid=19674541|s2cid=13549970|issn=0960-9822|doi-access=free}} [262] => [263] => These social systems are affected by three main ecological factors: distribution of resources, [[Group size measures|group size]], and [[predation]].{{cite book |last1=Pough |first1=F. W. |last2=Janis |first2=C. M. |last3=Heiser |first3=J. B. |title=Vertebrate Life |url=https://archive.org/details/vertebratelife0000poug |chapter=Primate Societies |year=2005 |orig-year=1979 |edition=7th |publisher= Pearson |pages=[https://archive.org/details/vertebratelife0000poug/page/621 621]–623 |isbn=0-13-127836-3}} Within a social group there is a balance between cooperation and competition. Cooperative behaviors in many primates species include [[social grooming]] (removing [[Parasitism|skin parasites]] and cleaning wounds), food sharing, and collective defense against predators or of a territory. Aggressive behaviors often signal competition for food, sleeping sites or mates. Aggression is also used in establishing [[dominance hierarchy|dominance hierarchies]].[[Barbara Smuts|Smuts, B.B.]], Cheney, D.L. Seyfarth, R.M., Wrangham, R.W., & Struhsaker, T.T. (Eds.) (1987). ''Primate Societies''. Chicago: University of Chicago Press for articles on the structure and function of various primate societies. [264] => [265] => In November 2023, scientists reported, for the first time, evidence that groups of primates, particularly [[bonobo]]s, are capable of cooperating with each other.{{cite news |last=Zimmer |first=Carl |authorlink=Carl Zimmer |title=Scientists Find First Evidence That Groups of Apes Cooperate - Some bonobos are challenging the notion that humans are the only primates capable of group-to-group alliances. |url=https://www.nytimes.com/2023/11/16/science/bonobos-cooperation-study.html |date=16 November 2023 |work=[[The New York Times]] |url-status=live |archiveurl=https://archive.today/20231116194259/https://www.nytimes.com/2023/11/16/science/bonobos-cooperation-study.html |archivedate=16 November 2023 |accessdate=17 November 2023 }}{{cite journal |author=Samuni, Liran |display-authors=et al. |title=Cooperation across social borders in bonobos [266] => |url=https://www.science.org/doi/10.1126/science.adg0844 |date=16 November 2023 |journal=[[Science (journal)|Science]] |volume=382 |issue=6672 |pages=805–809 |doi=10.1126/science.adg0844 |url-status=live |archiveurl=https://archive.today/20231117125744/https://www.science.org/doi/10.1126/science.adg0844 |archivedate=17 November 2023 |accessdate=17 November 2023 }} [267] => [268] => ===Interspecific associations=== [269] => Several species of primates are known to associate in the wild. Some of these associations have been extensively studied. In the [[Taï National Park|Tai Forest]] of Africa several species coordinate anti-predator behavior. These include the [[Diana monkey]], [[Campbell's mona monkey]], [[lesser spot-nosed monkey]], [[western red colobus]], [[king colobus]] (western [[Black-and-white colobus|black and white colobus]]), and [[sooty mangabey]], which coordinate anti-predator alarm calls.{{cite book|title=Monkeys of Tai Forest, An African Primate Community|chapter=Interactions between African Crowned Eagles and Their Prey Community|author1=Shultz, S. |author2=Thomsett, S. |name-list-style=amp |editor=McGraw, W. |editor2=Zuberbuhler, K. |editor3=Noe, R.|year=2007|publisher=Cambridge University Press|isbn=978-0-521-81633-5|pages=181}} Among the predators of these monkeys is the [[common chimpanzee]].{{cite book|title=Monkeys of Tai Forest, An African Primate Community|chapter=Interactions between Red Colobus Monkeys and Chimpanzees|author=Bshary, R.|editor=McGraw, W. |editor2=Zuberbuhler, K. |editor3=Noe, R.|year=2007|publisher=Cambridge University Press|isbn=978-0-521-81633-5|pages=155–170}} [270] => [271] => 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.{{cite book|title=Chimpanzee and Red Colobus : the ecology of predator and prey|url=https://archive.org/details/chimpanzeeredcol0000stan|author=Stanford, C.|year=1998|publisher=Harvard University Press|isbn=0-674-00722-0|pages=[https://archive.org/details/chimpanzeeredcol0000stan/page/130 130]–138, 233}} [272] => [273] => In South America, [[squirrel monkey]]s associate with [[capuchin monkey]]s.{{cite book|title=On the Move : how and why animals travel in groups|chapter=Social Manipulation Within and Between Troops Mediates Primate Group Movement|author=Boinski, S.|editor=Boinski, S. |editor2=Garber, P.|year=2000|publisher=University of Chicago Press|isbn=0-226-06340-2|pages=447–448}} This may have more to do with foraging benefits to the squirrel monkeys than anti-predation benefits. [274] => [275] => ===Communication=== [276] => {{Further|Great ape language}} [277] => {{listen [278] => | filename = Roep Indri Indri.ogg [279] => | title = Indri lemur wailing [280] => | description = Indri lemur wailing, [[ogg]]/[[Vorbis]] format. [281] => | format = [[Vorbis]] [282] => | filename2 = Howler monkey.ogg [283] => | title2 = Howler monkey roaring [284] => | description2 = Howler monkey roaring, [[ogg]]/[[Vorbis]] format. [285] => | format2 = [[Vorbis]] [286] => | filename3 = Vervet_Monkey_(Chlorocebus_pygerythrus)_(W_CERCOPITHECUS_AETHIOPS_R2_C2).ogg [287] => | title3 = Vervet monkey alarm call [288] => | description3 = Vervet monkey alarm call, [[ogg]]/[[Vorbis]] format. [289] => | format3 = [[Vorbis]] [290] => | filename4 = Symphalangus syndactylus 20170623 Doué-la-Fontaine.ogg [291] => | title4 = Siamang singing [292] => | description4 = Siamang singing, [[ogg]]/[[Vorbis]] format. [293] => | format4 = [[Vorbis]] [294] => }} [295] => [296] => [[Lemur]]s, [[loris]]es, [[tarsier]]s, and New World monkeys rely on [[Olfaction|olfactory signals]] for many aspects of social and reproductive behavior. Specialized glands are used to [[mark territories]] with [[pheromone]]s, which are detected by the [[vomeronasal organ]]; this process forms a large part of the communication behavior of these primates. In Old World monkeys and apes this ability is mostly [[Vestigiality|vestigial]], having regressed as [[Trichromacy|trichromatic]] eyes evolved to become the main sensory organ.{{cite journal |title=Relaxed selective pressure on an essential component of pheromone transduction in primate evolution |last1=Liman |first1=E. R. |last2=Innan |first2=H. |journal=Proceedings of the National Academy of Sciences of the United States of America |year=2003 |volume=100 |issue=6 |pages=3328–3332 |url=http://www.pnas.org/cgi/reprint/100/6/3328.pdf |access-date=2008-07-23 |doi=10.1073/pnas.0636123100 |pmid=12631698 |pmc=152292|bibcode = 2003PNAS..100.3328L|doi-access=free }} Primates also use vocalizations, gestures, and facial expressions to convey psychological state.{{cite book |last1=Egnor |first1=R. |last2=Miller |first2=C. |last3=Hauser |first3=M.D. |year=2004 |chapter=Nonhuman Primate Communication |title=Encyclopedia of Language and Linguistics |edition=2nd |publisher=Elsevier |isbn=0-08-044299-4 |chapter-url=http://www.wjh.harvard.edu/~mnkylab/publications/animalcommunication/PrimateComm_ElsevierEncy.pdf |url-status=dead |archive-url=https://web.archive.org/web/20080910043811/http://www.wjh.harvard.edu/~mnkylab/publications/animalcommunication/PrimateComm_ElsevierEncy.pdf |archive-date=2008-09-10}} Facial musculature is very developed in primates, particularly in monkeys and apes, allowing for complex facial communication. Like humans, chimpanzees can distinguish the faces of familiar and unfamiliar individuals.{{cite journal|author=Burrows, A. M.|year=2008|title=The facial expression musculature in primates and its evolutionary significance|journal=BioEssays|volume=30|issue=3|pages=212–225|doi=10.1002/bies.20719|pmid=18293360|s2cid=205478149|url=https://www.researchgate.net/publication/5559596}} Hand and arm gestures are also important forms of communication for great apes and a single gesture can have multiple functions.{{cite journal|last1=Pollick |first1=A. S. |last2=de Waal |first2=F. B. M.|year=2007|title=Ape gestures and language evolution|journal=Proceedings of the National Academy of Sciences|volume=104|issue=19|pages=8184–8189|doi=10.1073/pnas.0702624104|pmid=17470779 |pmc=1876592|bibcode=2007PNAS..104.8184P|doi-access=free }} [297] => [298] => Primates are a particularly vocal group of mammals. [[Indri]]s and [[black-and-white ruffed lemur]]s make distinctive, loud songs and choruses which maintain territories and act as [[alarm call]]s.{{cite journal|url=http://gibbons.de/main/non-gibbon/pdf_files/2006indri_varecia.pdf |author1=Geissmann, Thomas |author2=Mutschler, Thomas |name-list-style=amp |title=Diurnal Distribution of Loud Calls in Sympatric Wild Indris (''Indri indri'') and Ruffed Lemurs (''Varecia variegata''): Implications for Call Functions|pmid=16736264|year=2006|volume=47|issue=4|pages=393–6|doi=10.1007/s10329-006-0189-5|journal=Primates; Journal of Primatology|s2cid=1586657 }} The [[Philippine tarsier]], has a high-frequency limit of auditory sensitivity of approximately 91 kHz with a dominant frequency of 70 kHz, among the highest recorded for any terrestrial mammal. For Philippine tarsiers, these ultrasonic vocalizations might represent a private channel of communication that subverts detection by predators, prey and competitors, enhances energetic efficiency, or improves detection against low-frequency background noise.{{cite journal |last1=Ramsier |first1=M.A. |last2=Cunningham |first2=A.J. |last3=Moritz |first3=G.L. |last4=Finneran |first4=J.J. |last5=Williams |first5=C.V. |last6=Ong |first6=P.S. |last7=Gursky-Doyen |first7=S.L. |last8=Dominy |first8=N.J. | year = 2012 | title = Primate communication in the pure ultrasound| journal = Biology Letters | volume = 8 | issue = 4| pages = 508–511 | doi = 10.1098/rsbl.2011.1149 | pmid = 22319094 | pmc=3391437}} Male [[howler monkey]]s are among the loudest land mammals as their roars can be heard up to {{Convert|4.8|km|mi|abbr=on}}, and relate to intergroup spacing, territorial protection and possibly mate-guarding.{{cite journal |last1=da Cunha |first1=R. G. T. |last2=Byrne |first2=R.|year=2006|title=Roars of Black Howler Monkeys (Alouatta caraya): Evidence for a Function in Inter-Group Spacing|journal=Behaviour|volume=143|issue=10|pages=1169–1199|jstor=4536401|doi=10.1163/156853906778691568}}{{cite web|title=Black howler monkey|date=25 April 2016|publisher=Smithsonian's National Zoo & Conservation Biology Institute|url=https://nationalzoo.si.edu/animals/black-howler-monkey|access-date=2016-07-10}} Roars are produced by a modified [[larynx]] and enlarged [[hyoid bone]] which contains an air sac.{{cite journal|author=Kelemen, G.|author2= Sade, J.|year=1960|title=The vocal organ of the Howling monkey (''Alouatta palliata'')|journal=Journal of Morphology|volume=107|issue=2|pages=123–140|doi=10.1002/jmor.1051070202|pmid= 13752246|s2cid= 30539292}} 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.{{cite journal| last1=Seyfarth| first1=R. M. |first2=D. L. |last2=Cheney |first3=Peter |last3=Marler |title=Vervet Monkey Alarm Calls: Semantic communication in a Free-Ranging Primate |journal=Animal Behaviour |volume=28 |issue=4 |pages=1070–1094 |year=1980 |doi=10.1016/S0003-3472(80)80097-2 | s2cid=53165940 |url=https://www.researchgate.net/publication/223576319}} Male and female [[siamang]]s both possess inflatable pouches in the throat with which pair -bonds use to sing "duets" to each other.{{cite journal|last1=Haimoff|first1=E. H.|year=1983|title=Brief report: Occurrence of anti-resonance in the song of the siamang (''Hylobates syndactylus'')|journal=American Journal of Primatology|volume=5|issue=3|pages=249–256|doi=10.1002/ajp.1350050309|pmid=31986856 |s2cid=85262432 }} [299] => [300] => Many non-human primates have the vocal anatomy to produce human speech but lack the proper brain wiring.{{cite journal|last1=Fitch |first1=W. T. |last2=de Boer |first2=B. |last3=Mathur |first3=N. |last4=Ghazanfar |first4=A. A.|year=2016|title=Monkey vocal tracts are speech-ready|journal=Science Advances|volume=2|issue=12|page=e1600723|doi=10.1126/sciadv.1600723|pmid=27957536 |bibcode=2016SciA....2E0723F|pmc=5148209}} Vowel-like vocal patterns have been recorded in baboons which has implications for the origin of speech in humans.{{cite journal|author1=Boë L.-J. |author2=Berthommier, F. |author3=Legou, T. |author4=Captier, G. |author5=Kemp, C. |author6=Sawallis, T. R. |author7=Becker Y. |author8=Rey, A. |author9=Fagot, J.|display-authors=6|year=2017|title=Evidence of a Vocalic Proto-System in the Baboon (''Papio papio'') Suggests Pre-Hominin Speech Precursors|journal=PLOS ONE|volume=12|issue=1|page=e0169321|doi=10.1371/journal.pone.0169321|pmid=28076426 |bibcode=2017PLoSO..1269321B|pmc=5226677|doi-access=free }} Consonant- and vowel-like sounds exist in some orangutan calls and they maintain their meaning over great distances.{{cite journal|last1=Lameira|first1=A. R.|display-authors=etal|year=2021|title=Orangutan information broadcast via consonant-like and vowel-like calls breaches mathematical models of linguistic evolution|journal=Biology Letters|volume=17|issue=9|doi=10.1098/rsbl.2021.0302|pmid=34582737 |pmc=8478518 }} The time range for the evolution of human language and/or its anatomical prerequisites extends, at least in principle, from the phylogenetic divergence of ''[[Homo]]'' (2.3 to 2.4 million years ago) from ''[[Pan (genus)|Pan]]'' (5 to 6 million years ago) to the emergence of full [[behavioral modernity]] some 50,000–150,000 years ago. Few dispute that ''[[Australopithecus]]'' probably lacked vocal communication significantly more sophisticated than that of [[great ape]]s in general.{{Cite journal | last1 = Arcadi | first1 = AC. | title = Vocal responsiveness in male wild chimpanzees: implications for the evolution of language | journal = J Hum Evol | volume = 39 | issue = 2 | pages = 205–23 |date=Aug 2000 | doi = 10.1006/jhev.2000.0415 | pmid = 10968929 | s2cid = 7403772 | doi-access = free }} [301] => [302] => ===Life history=== [303] => [[File:Bonnet macaque nursing, Bangalore.jpg|A [[crab-eating macaque]] breastfeeding her baby|thumb]] [304] => Primates have slower rates of development than other mammals. All primate infants are [[breastfeeding|breastfed]] by their mothers (with the exception of some human cultures and various zoo raised primates which are fed formula) and rely on them for grooming and transportation. In some species, infants are protected and transported by males in the group, particularly males who may be their fathers. Other relatives of the infant, such as siblings and aunts, may participate in its care as well. Most primate mothers cease [[ovulation]] while breastfeeding an infant; once the infant is [[Weaning|weaned]] the mother can reproduce again. This often leads to weaning conflict with infants who attempt to continue breastfeeding. [305] => [306] => [[Infanticide (zoology)|Infanticide]] is common in polygynous species such as [[gray langur]]s and gorillas. Adult males may kill dependent offspring that are not theirs so the female will return to estrus and thus they can sire offspring of their own. Social monogamy in some species may have evolved to combat this behavior.{{cite journal|author1=Opie, Christopher |author2=Atkinson, Quentin D. |author3=Dunbarc, Robin I. M. |author4=Shultz, Susanne |year=2013|title=Male infanticide leads to social monogamy in primates|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=110|issue=33|pages=13328–13332|doi=10.1073/pnas.1307903110|pmid=23898180 |pmc=3746880|bibcode=2013PNAS..11013328O|doi-access=free }} [[Promiscuity]] may also lessen the risk of infanticide since paternity becomes uncertain.{{cite journal|author1=De Ruiter, Jan R.|author2= Van Hooff, Jan A. R. A. M.|author-link2=Jan van Hooff|author3=Scheffrahn, Wolfgang |name-list-style=amp |year=1994|title=Social and genetic aspects of paternity in wild long-tailed macaques (''Macaca fascicularis'')|journal=Behaviour|volume=129|issue=3–4|pages=203–24|jstor=4535195|doi=10.1163/156853994x00613}} [307] => [308] => Primates have a longer [[Juvenile (organism)|juvenile]] period between weaning and sexual maturity than other mammals of similar size. Some primates such as [[galago]]s and [[new world monkey]]s use tree-holes for [[Nest-building in primates|nesting]], and park juveniles in leafy patches while foraging. Other primates follow a strategy of "riding", i.e. carrying individuals on the body while feeding. Adults may construct or use nesting sites, sometimes accompanied by juveniles, for the purpose of resting, a behavior which has developed secondarily in the great apes.{{cite journal |last1=Kappeler |first1=Peter M. |year=1998 |title=Nests, Tree Holes, and the Evolution of Primate Life Histories |journal=American Journal of Primatology |issue=1 |pages=7–33 |pmid=9730211 |doi=10.1002/(SICI)1098-2345(1998)46:1<7::AID-AJP3>3.0.CO;2-# |volume=46|s2cid=196589387 }}{{cite journal |last1=Ross |first1= Caroline|year=1991 |title=Park or ride? Evolution of infant carrying in primates. |journal=International Journal of Primatology |publisher=Kluwer Academic Publishing |volume=22 |issue=5 |pages=749–771 |doi=10.1023/A:1012065332758|s2cid= 25301078}} 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.{{cite web |url=http://www.ibtimes.com/humans-primates-burn-50-percent-fewer-calories-each-day-other-mammals-1539866 |title=Humans And Primates Burn 50 Percent Fewer Calories Each Day Than Other Mammals |last1=Mintz |first1=Zoe |date=14 January 2014 |website=www.ibtimes.com |publisher=IBT Media Inc. |access-date=2014-01-14}} Late in life, female catarrhine primates appear to undergo a cessation of reproductive function known as [[menopause]]; other groups are less studied.{{cite journal | author = Walker ML, Herndon JG | title = Menopause in nonhuman primates? | journal = Biology of Reproduction | volume = 79 | pages = 398–406 | year = 2008 | pmid = 18495681 | doi = 10.1095/biolreprod.108.068536 | issue = 3 | pmc=2553520 | last2 = Herndon}} [309] => [310] => ===Diet and feeding=== [311] => [[File:Colubusmonkey.JPG|thumb|Leaf eating [[mantled guereza]], a species of [[black-and-white colobus]]]] [312] => [[File:Microcebus murinus -Artis Zoo, Amsterdam, Netherlands-8c.jpg|thumb|right|A [[mouse lemur]] holds a cut piece of fruit in its hands and eats.]] [313] => Primates exploit a variety of food sources. It has been said that many characteristics of modern primates, including humans, derive from an early ancestor's practice of taking most of its food from the tropical canopy.{{cite magazine |last1 = Milton |first1 = K. |year=1993 |title = Diet and Primate Evolution |url=http://nature.berkeley.edu/miltonlab/pdfs/diet_primate_evolution.pdf |magazine=Scientific American |volume=269 |issue=2 |pages=86–93 |pmid=8351513 |doi=10.1038/scientificamerican0893-86|bibcode=1993SciAm.269b..86M}} Most primates include fruit in their diets to obtain easily digested nutrients including [[carbohydrate]]s and [[lipid]]s 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 [[Haplorhini]] (tarsiers, monkeys and apes) have lost this ability, and require the vitamin in their diet.{{cite journal | title = Vitamin C biosynthesis in prosimians: Evidence for the anthropoid affinity of Tarsius |last1=Pollock |first1=J. I. |last2=Mullin |first2=R. J. | journal = American Journal of Physical Anthropology | year = 1986 | volume = 73 | issue = 1 | pages = 65–70 | doi = 10.1002/ajpa.1330730106 | url = http://www3.interscience.wiley.com/journal/110488482/abstract?CRETRY=1&SRETRY=0 | archive-url = https://archive.today/20120628232930/http://www3.interscience.wiley.com/journal/110488482/abstract?CRETRY=1&SRETRY=0 |url-status=dead | archive-date = 2012-06-28 | pmid = 3113259}} [314] => [315] => Many primates have anatomical specializations that enable them to exploit particular foods, such as fruit, leaves, gum or [[insect]]s. For example, leaf eaters such as howler monkeys, [[black-and-white colobus]]es and [[sportive lemur]]s have extended digestive tracts which enable them to absorb nutrients from leaves that can be difficult to digest. [[Marmoset]]s, which are gum eaters, have strong [[incisor]] teeth, enabling them to open tree bark to get to the gum, and claws rather than nails, enabling them to cling to trees while feeding. 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.{{cite journal |last1=Milliken |first1=G. W. |last2=Ward |first2=J. P. |last3=Erickson |first3=C. J. |year=1991 |title=Independent digit control in foraging by the aye-aye (''Daubentonia madagascariensis'') |journal=Folia Primatologica |volume=56 |issue=4 |pages=219–224 |doi=10.1159/000156551 |pmid=1937286}} Some species have additional specializations. For example, the [[grey-cheeked mangabey]] has thick [[tooth enamel|enamel]] on its teeth, enabling it to open hard fruits and seeds that other monkeys cannot. The [[gelada]] is the only primate species that feeds primarily on grass.{{cite web |last=Hiller|first=C.|year=2000|url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Theropithecus_gelada.html|title = ''Theropithecus gelada''|access-date=2008-08-08|work=Animal Diversity Web}} [316] => [317] => ====Hunting==== [318] => [[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.]] [319] => [[Tarsier]]s are the only [[Extant taxon|extant]] [[obligate carnivore|obligate carnivorous]] primates, exclusively eating insects, crustaceans, small vertebrates and snakes (including [[Venomous snake|venomous]] species).{{cite book|title=Tarsiers Past, Present and Future|chapter=Introduction|last1=Wright |first1=P. |last2=Simmons |first2=E. |last3=Gursky |first3=S.|editor=Wright, P. |editor2=Simmons, E. |editor3=Gursky, S.|year=2003|publisher=Rutgers University Press|isbn=0-8135-3236-1|pages=1}} [[Capuchin monkey]]s can exploit many different types of plant matter, including fruit, leaves, flowers, buds, nectar and seeds, but also eat insects and other [[invertebrate]]s, bird eggs, and small vertebrates such as birds, [[lizard]]s, [[squirrel]]s and [[bat]]s. [320] => [321] => 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%.{{cite book | last = Goodall | first = Jane | author-link = Jane Goodall | year = 1986 | title = The Chimpanzees of Gombe: Patterns of Behavior | publisher = Belknap Press of Harvard University Press | isbn = 0-674-11649-6 | url = https://archive.org/details/chimpanzeesofgom00good}}{{cite web|last=Guernsey|first=Paul|title=WHAT DO CHIMPS EAT?|url=http://www.allaboutwildlife.com/what-do-chimps-eat|work=All About Wildlife|access-date=2013-04-22|archive-date=2019-11-18|archive-url=https://web.archive.org/web/20191118084847/http://www.allaboutwildlife.com/what-do-chimps-eat|url-status=dead}} The meat consumption includes predation on other primate species, such as the [[western red colobus]] monkey. The [[bonobo]] is an [[omnivorous]] [[frugivore]] – the majority of its diet is fruit, but it supplements this with leaves, meat from small [[vertebrate]]s, such as [[anomalure]]s, [[flying squirrel]]s and [[duiker]]s,{{cite journal |author=Ihobe H |title=Observations on the meat-eating behavior of wild bonobos (''Pan paniscus'') at Wamba, Republic of Zaire |journal=Primates |volume=33 |issue=2 |pages=247–250|year=1992 |doi=10.1007/BF02382754|s2cid=10063791 }} and [[invertebrate]]s.{{cite book |last1=Rafert |first1=J. |first2=E.O. |last2=Vineberg |year=1997 |chapter=Bonobo Nutrition – relation of captive diet to wild diet |chapter-url=http://www.nagonline.net/HUSBANDRY/Diets%20pdf/Bonobo%20Nutrition.pdf |url-status=dead |archive-url=https://web.archive.org/web/20120425232556/http://www.nagonline.net/HUSBANDRY/Diets%20pdf/Bonobo%20Nutrition.pdf |archive-date=2012-04-25 |title=Bonobo Husbandry Manual |publisher=American Association of Zoos and Aquariums}} In some instances, bonobos have been shown to consume lower-order primates.{{cite journal |last1=Surbeck |first1=M |last2=Fowler |first2=A |last3=Deimel |first3=C |last4=Hohmann |first4=G |title=Evidence for the consumption of arboreal, diurnal primates by bonobos (''Pan paniscus'') |journal=American Journal of Primatology |volume=71 |issue=2 |pages=171–4 |year=2008 |doi=10.1002/ajp.20634 |pmid=19058132|s2cid=32622605 }}{{cite journal |author=Surbeck M, Hohmann G |title=Primate hunting by bonobos at LuiKotale, Salonga National Park |journal=Current Biology |volume=18 |issue=19 |pages=R906–7 |date=14 October 2008 |doi=10.1016/j.cub.2008.08.040 |pmid=18957233 |last2=Hohmann|s2cid=6708310 |doi-access=free }} [322] => [323] => Until the development of agriculture approximately 10,000 years ago, ''Homo sapiens'' employed a hunter-gatherer method as their sole means of food collection. This involved combining stationary food sources (such as fruits, grains, tubers, and mushrooms, insect larvae and aquatic mollusks) with [[Game (food)|wild game]], which must be hunted and killed in order to be consumed.{{cite journal |author=Cordain L |title=Origins and evolution of the Western diet: health implications for the 21st century |journal=Am. J. Clin. Nutr. |volume=81 |issue=2 |pages=341–54 |date=February 2005 |pmid=15699220 |name-list-style=vanc |author2=Eaton SB |author3=Sebastian A |display-authors=3 |last4=Mann |first4=N |last5=Lindeberg |first5=S |last6=Watkins |first6=BA |last7=O'Keefe |first7=JH |last8=Brand-Miller |first8=J |doi=10.1093/ajcn.81.2.341|doi-access=free }} It has been proposed that humans have used fire to prepare and [[cooking|cook]] food since the time of ''[[Homo erectus]]''.{{cite journal |author=Ulijaszek SJ |title=Human eating behaviour in an evolutionary ecological context |journal=Proc Nutr Soc |volume=61 |issue=4 |pages=517–26 |date=November 2002 |pmid=12691181 |doi=10.1079/PNS2002180|doi-access=free }} Around ten thousand years ago, [[History of agriculture|humans developed agriculture]],[http://www.archaeology.org/9707/newsbriefs/squash.html Earliest agriculture in the Americas] {{webarchive|url=https://web.archive.org/web/20100603232246/http://www.archaeology.org/9707/newsbriefs/squash.html |date=3 June 2010 }} [324] => [http://sciencenow.sciencemag.org/cgi/content/full/2007/213/2 Earliest cultivation of barley] {{webarchive|url=https://web.archive.org/web/20070216093200/http://sciencenow.sciencemag.org/cgi/content/full/2007/213/2 |date=16 February 2007 }} [325] => [http://news.bbc.co.uk/2/hi/science/nature/5038116.stm Earliest cultivation of figs] {{webarchive|url=https://web.archive.org/web/20060602081110/http://news.bbc.co.uk/2/hi/science/nature/5038116.stm |date=2 June 2006 }}, retrieved 19 February 2007 which substantially altered their diet. This change in diet may also have altered human biology; with the spread of [[dairy farming]] providing a new and rich source of food, leading to the evolution of the ability to digest [[lactose]] in some adults.{{cite journal |author=Krebs JR |title=The gourmet ape: evolution and human food preferences |journal=Am. J. Clin. Nutr. |volume=90 |issue=3 |pages=707S–11S |date=September 2009 |pmid=19656837 |doi=10.3945/ajcn.2009.27462B|doi-access=free }}{{cite journal |vauthors=Holden C, Mace R |title=Phylogenetic analysis of the evolution of lactose digestion in adults |url=https://archive.org/details/sim_human-biology_1997-10_69_5/page/605 |journal=Hum. Biol. |volume=69 |issue=5 |pages=605–28 |date=October 1997 |pmid=9299882}} [326] => [327] => ===As prey=== [328] => Predators of primates include various species of [[carnivora]]ns, [[birds of prey]], [[reptile]]s, and other primates. Even gorillas have been recorded as prey. Predators of primates have diverse hunting strategies and as such, primates have evolved several different [[antipredator adaptation]]s including [[crypsis]], [[Alarm signal|alarm calls]] and [[Mobbing (animal behavior)|mobbing]]. Several species have separate alarm calls for different predators such as air-borne or ground-dwelling predators. Predation may have shaped group size in primates as species exposed to higher predation pressures appear to live in larger groups.{{cite book|author=Fichtel, Claudia|contribution=Predation|year=2012|title=The Evolution of Primate Societies|editor1=Mitani, John C. |editor2=Call, Josep |editor3=Kappeler, Peter M. |editor4=Palombit, Ryne A. |editor5=Silk, Joan B. |editor5-link=Joan Silk |publisher=University of Chicago Press|pages=169–84|isbn=978-0-226-53172-4}} [329] => [330] => ===Intelligence and cognition=== [331] => {{Main|Primate cognition}} [332] => Primates have advanced cognitive abilities: some make tools and use them to acquire food and for social displays;{{cite journal |last1=Boesch |first1=C. |last2=Boesch |first2=H. |title=Tool Use and Tool Making in Wild Chimpanzees |journal=[[Folia Primatologica]] |year=1990 |volume=54 |pages=86–99 |doi=10.1159/000156428 |pmid=2157651 |issue=1–2}}{{cite journal |last1=Westergaard |year=1998 |title=Why some capuchin monkeys (''Cebus apella'') use probing tools (and others do not) |url=https://archive.org/details/sim_journal-of-comparative-psychology_1998-06_112_2/page/207 |journal=Journal of Comparative Psychology |volume=112 |issue=2 |pages=207–211 |doi=10.1037/0735-7036.112.2.207 |pmid=9642788 |first1=G. C. |last2=Lundquist|first2=A. L.|display-authors=2 |last3=Haynie |first3=Michael Katherine |last4=Kuhn |first4=Heather E. |last5=Suomi |first5=Stephen J.}} some can perform tasks requiring cooperation, influence and rank;{{cite journal |last1=de Waal |first1=F. B. M. |last2=Davis |first2=J. M. |year=2003 |title=Capuchin cognitive ecology: cooperation based on projected returns |journal=Neuropsychologia |volume=41 |pages=221–228 |doi=10.1016/S0028-3932(02)00152-5 |pmid=12459220 |issue=2|s2cid=8190458 }} they are status conscious, manipulative and capable of deception;{{cite journal |last1=Paar |first1=L. A. |last2=Winslow |first2=J. T. |last3=Hopkins |first3=W. D. |last4=de Waal |first4=F. B. M. |year=2000 |title=Recognizing facial cues: Individual discrimination by chimpanzees (''Pan troglodytes'') and rhesus monkeys (''Macaca mulatta'') |url=https://archive.org/details/sim_journal-of-comparative-psychology_2000-03_114_1/page/47 |journal=Journal of Comparative Psychology |volume=114 |issue=1 |pages=47–60 |doi=10.1037/0735-7036.114.1.47 |pmid=10739311 |pmc=2018744}}{{cite journal |last=Byrne |first=Richard |author2=Corp, Nadia |title=Neocortex size predicts deception rate in primates |journal= Proceedings of the Royal Society of London. Series B: Biological Sciences|year=2004 |volume=271 |issue=1549 |pages=1693–1699 |doi=10.1098/rspb.2004.2780|pmid=15306289 |pmc=1691785}} they can recognise [[Kinship|kin]] and [[conspecific]]s;{{cite journal |last1=Paar |first1=L. A. |last2=de Waal |first2=F. B. M. |year=1999 |title=Visual kin recognition in chimpanzees |journal=Nature |volume=399 |pages=647–648 |doi=10.1038/21345 |pmid=10385114 |issue=6737|bibcode = 1999Natur.399..647P|s2cid=4424086 }}{{cite journal |year=1997 |title=Discrimination of macaques by macaques: The case of sulawesi species |journal=Primates |volume=38 |issue=3 |pages=233–245 |last1=Fujita |first1=K. |last2=Watanabe |first2=K. |last3=Widarto |first3=T. H. |last4=Suryobroto |first4=B. |doi=10.1007/BF02381612|s2cid=21042762 }} and they can learn to use symbols and understand aspects of human language including some relational syntax and concepts of number and numerical sequence.{{cite journal |last=Call |first=J. |year=2001 |title=Object permanence in orangutans (''Pongo pygmaeus''), chimpanzees (''Pan troglodytes''), and children (''Homo sapiens'') |url=https://archive.org/details/sim_journal-of-comparative-psychology_2001-06_115_2/page/159 |journal=Journal of Comparative Psychology |volume=115 |issue=2 |pages=159–171 |doi=10.1037/0735-7036.115.2.159 |pmid=11459163}}{{cite journal |last1=Itakura |first1=S. |last2=Tanaka |first2=M. |date=June 1998 |title=Use of experimenter-given cues during object-choice tasks by chimpanzees (''Pan troglodytes''), an orangutan (''Pongo pygmaeus''), and human infants (''Homo sapiens'') |url=https://archive.org/details/sim_journal-of-comparative-psychology_1998-06_112_2/page/119 |journal=Journal of Comparative Psychology |volume=112 |issue=2 |pages=119–126 |doi=10.1037/0735-7036.112.2.119 |pmid=9642782}}{{cite journal |last1=Gouteux |first1=S. |last2=Thinus-Blanc |first2=C. |last3=Vauclair |first3=J. |year=2001 |title=Rhesus monkeys use geometric and nongeometric information during a reorientation task |journal=Journal of Experimental Psychology: General |volume=130 |issue=3 |pages=505–519 |doi=10.1037/0096-3445.130.3.505 |pmid=11561924 |url=http://cogprints.org/3590/1/Gouteux_et_al_JEPGEN_01.pdf}} Research in primate cognition explores problem solving, memory, social interaction, a [[Theory of mind#Non-human|theory of mind]], and numerical, spatial, and abstract concepts.{{cite book |author1=Tomasello, M. |author2=Call, J. |name-list-style=amp |title=Primate Cognition |publisher=Oxford University Press US |year=1997 |isbn=978-0-19-510624-4}} 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.{{cite journal | last1 = Deaner | first1 = R. O. | last2 = van Schaik | first2 = C. P. | last3 = Johnson | first3 = V. E. | year = 2006 | title = Do some taxa have better domain-general cognition than others? A metaanalysis of nonhuman primate studies | journal = [[Evolutionary Psychology (journal)|Evolutionary Psychology]] | volume = 4 | pages = 149–196 | doi = 10.1177/147470490600400114| s2cid = 16702785 | doi-access = free }}{{Cite journal | last1 = Reader | first1 = S. M. | last2 = Hager | first2 = Y. | last3 = Laland | first3 = K. N. | title = The evolution of primate general and cultural intelligence | journal = [[Philosophical Transactions of the Royal Society B]] | volume = 366 | issue = 1567 | pages = 1017–1027 | date = 2011 | url = http://lalandlab.st-andrews.ac.uk/pdf/Publication163.pdf | doi = 10.1098/rstb.2010.0342 | pmid = 21357224 | access-date = 2011-07-04 | url-status=dead | archive-url = https://web.archive.org/web/20111003152030/http://lalandlab.st-andrews.ac.uk/pdf/Publication163.pdf | archive-date = 2011-10-03 | pmc = 3049098}} However, there is a great deal of variation in each group (e.g., among New World monkeys, both [[spider monkey|spider]] and [[capuchin monkey]]s have scored highly by some measures), as well as in the results of different studies. [333] => [334] => ====Tool use and manufacture==== [335] => {{Main|Tool use by animals}} [336] => [[File:Cultural-differences-in-ant-dipping-tool-length-between-neighbouring-chimpanzee-communities-at-srep12456-s2.ogv|thumb|right|[[Chimpanzee]]s using twigs to dip for ants]] [337] => [[File:Macaca fascicularis aurea stone tools - journal.pone.0072872.g002.png|thumb|[[Crab-eating macaque]]s with [[stone tool]]s]] [338] => 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.{{cite web|url=https://www.janegoodall.org.uk/chimpanzees/chimpanzee-central/15-chimpanzees/chimpanzee-central/19-toolmaking|title=Toolmaking|publisher=The Jane Goodall Institute|access-date=2013-08-01}} 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.{{cite web|url=http://www.clemetzoo.com/apetag/Bonobos.html |title=Bonobos |publisher=ApeTag |year=2010 |access-date=2013-08-03 |url-status=dead |archive-url=https://web.archive.org/web/20131102050710/https://www.clemetzoo.com/apetag/Bonobos.html |archive-date=2013-11-02}} It has been reported that females, both chimpanzee and bonobo, use tools more avidly than males. [339] => {{cite journal|last1=Gruber |first1=T. |last2=Clay |first2=Z. |last3=Zuberbühler |first3=K.|year=2010|title= A comparison of bonobo and chimpanzee tool use: evidence for a female bias in the ''Pan'' lineage|journal= Animal Behaviour|volume= 80|issue=6|pages= 1023–1033|url=http://www.emory.edu/LIVING_LINKS/publications/articles/Gruber_etal_2010.pdf|doi=10.1016/j.anbehav.2010.09.005|s2cid=14923158 }} [[Orangutan]]s in [[Borneo]] scoop catfish out of small ponds. Over two years, anthropologist Anne Russon observed orangutans learning to jab sticks at catfish to scare them out of the ponds and in to their waiting hands.{{cite magazine|url=https://www.wired.com/wiredscience/2011/04/orangutan-tools-fishing/ |title=Orangutans use simple tools to catch fish |author=Bower, B. |magazine=[[Wired (website)|Wired]] |date=18 April 2011|access-date=2013-08-05}} There are few reports of [[gorilla]]s using tools in the wild. An adult female [[western lowland gorilla]] used a branch as a walking stick apparently to test water depth and to aid her in crossing a pool of water. Another adult female used a detached trunk from a small shrub as a stabilizer during food gathering, and another used a log as a bridge.{{cite journal|last1=Breuer |first1=T. |last2=Ndoundou-Hockemba |first2=M. |last3=Fishlock |first3=V.|year=2005|title= First observation of tool use in wild gorillas|journal= PLOS Biology |volume=3|issue=11|pages=e380 |doi=10.1371/journal.pbio.0030380|pmid=16187795|pmc=1236726 |doi-access=free }} [340] => [341] => The first direct observation of a non-ape primate using a tool in a wild environment occurred in 1988. Primatologist Sue Boinski watched an adult male white-faced capuchin beat a [[Bothrops asper|fer-de-lance]] snake to death with a dead branch.{{cite journal | title = Use of a club by a wild white-faced capuchin (''Cebus capucinus'') to attack a venomous snake (''Bothrops asper'')| year = 1988| last1 = Boinski| first1 = S.| journal = American Journal of Primatology| volume = 14| issue = 2| pages = 177–179| pmid = 31973450| s2cid = 84653622 |doi = 10.1002/ajp.1350140208}} The black-striped capuchin was the first non-ape primate for which routine tool use was documented in the wild; individuals were observed [[Primate archaeology|cracking nuts]] by placing them on a stone anvil and hitting them with another large stone.{{cite journal|last1=Fragaszy |first1=D. |last2=Izar |first2=P. |last3=Visalberghi |first3=E. |last4=Ottoni |first4=E.B. |last5=de Oliveira |first5=M.G.|year=2004|title= Wild capuchin monkeys (''Cebus libidinosus'') use anvils and stone pounding tools|journal= American Journal of Primatology|volume=64|issue=4|pages= 359–366|doi= 10.1002/ajp.20085|pmid=15580579|s2cid=16222308 }} In Thailand and Myanmar, [[crab-eating macaque]]s use stone tools to open nuts, oysters and other bivalves, and various types of sea snails.{{cite journal|last1=Gumert|first1=M.D.|last2=Kluck|first2=M.|last3=Malaivijitnond|first3=S.|title=The physical characteristics and usage patterns of stone axe and pounding hammers used by long-tailed macaques in the Andaman Sea region of Thailand|journal=American Journal of Primatology|year=2009|volume=71|issue=7|pages=594–608|doi=10.1002/ajp.20694|pmid=19405083|s2cid=22384150}} 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.{{cite journal |last1=Hamilton |first1=W.J. |last2=Buskirk |first2=R.E. |last3=Buskirk |first3=W.H.|year=1975|title= Defensive stoning by baboons|journal= Nature|volume= 256|pages= 488–489| doi=10.1038/256488a0|issue=5517|bibcode=1975Natur.256..488H|s2cid=4149862 }} [342] => [343] => 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.{{cite book | last1 = Fichtel | first1 = C. | last2 = Kappeler | first2 = P. M. | chapter = Chapter 19: Human universals and primate symplesiomorphies: Establishing the lemur baseline | editor1-last = Kappeler | editor1-first = P. M. | editor2-last = Silk | editor2-first = J. B. | title = Mind the Gap: Tracing the Origins of Human Universals | publisher = Springer | year = 2010 | isbn = 978-3-642-02724-6 | chapter-url = https://books.google.com/books?id=MFzxVH_OxjsC&pg=PA395}} [344] => [345] => 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 [[human]]s manufactured and used tools, and that this ability was what separated humans from other animals. Chimpanzees have also been observed making "sponges" out of leaves and moss that suck up water.{{cite journal | author=Sugiyama, Y. | year=1995 | title=Drinking tools of wild chimpanzees at Bossou | journal=American Journal of Primatology | volume=37 | issue=1 | pages=263–269 | doi=10.1002/ajp.1350370308| pmid=31936951| s2cid=86473603 }} 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.{{cite web|url=http://orangutanislands.com/sumatra-orangutans.htm |title=Sumatran orangutans |publisher=OrangutanIslands.com |access-date=2013-08-02 |url-status=dead |archive-url=https://web.archive.org/web/20131126074458/https://orangutanislands.com/sumatra-orangutans.htm |archive-date=2013-11-26}}{{cite journal|last1=van Schaik |first1=C. |last2=Fox |first2=E. |last3=Sitompul |first3=A.|year=1996|title= Manufacture and use of tools in wild Sumatran orangutans|url=https://archive.org/details/sim_naturwissenschaften_1996-04_83_4/page/186 |journal= Naturwissenschaften|volume= 83|issue=4|pages= 186–188|doi=10.1007/BF01143062|pmid=8643126 |bibcode=1996NW.....83..186V|s2cid=27180148 }} 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.{{cite web | url = http://www.bbc.co.uk/nature/14227783 | title = Mandrill monkey makes 'pedicuring' tool | first = Victoria | last = Gill | publisher = BBC | date = 22 July 2011 | access-date = 2013-08-11}} Captive gorillas have made a variety of tools.{{cite web|url=http://www.rozhlas.cz/therevealed/comments/_zprava/488947|author=Vancatova, M.|title=Gorillas and Tools – Part I|year=2008|access-date=2013-08-04}} [346] => [347] => ==Ecology== [348] => {{See also|List of primates by population}} [349] => [[File:Macaque India 3.jpg|thumb|upright|right|[[Rhesus macaque]] at [[Agra Fort]], [[India]]]] [350] => Non-human primates primarily live in the [[Tropics|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 [[Honshū]] 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 and subtropical moist broadleaf forests|tropical rainforest]]s but are also found in [[temperate forest]]s, [[savanna]]s, [[desert]]s, mountains and coastal areas.{{Cite book |author=Cowlishaw, G. |author2=Clutton-Brock, T. |contribution=Primates |year=2009 |title=The Princeton Encyclopedia of Mammals |editor=MacDonald, D. |publisher=Princeton and Oxford University Press |pages=270–280 |isbn=978-0-691-14069-8}} The number of primate species within tropical areas has been shown to be positively [[correlation|correlated]] to the amount of rainfall and the amount of rain forest area.{{cite journal|title=Geographic and climatic control of primate diversity|last1=Reed |first1=K. |last2=Fleagle |first2=J.|journal=Proceedings of the National Academy of Sciences of the United States of America|volume=92|issue=17|pages=7874–7876|date=August 15, 1995|doi=10.1073/pnas.92.17.7874|pmid=7644506|pmc=41248|bibcode = 1995PNAS...92.7874R|doi-access=free }} Accounting for 25% to 40% of the [[frugivore|fruit-eating animals]] (by [[biomass|weight]]) within tropical rainforests, primates play an important [[Ecology|ecological]] role by dispersing seeds of many tree species.{{cite book|title=Primates in Perspective|author1=Chapman, C. |author2=Russo, S. |name-list-style=amp |chapter=Primate Seed Dispersal|year=2007|publisher=Oxford University Press|editor=Campbell, C. J. |editor2=Fuentes, A. |editor3=MacKinnon, K. C. |editor4=Panger, M. |editor5=Bearder, S. K.|isbn=978-0-19-517133-4|pages=510}} [351] => [352] => 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),{{cite journal | last1=Long |first1=Y. C. |last2=Kirkpatrick |first2=R. C. |last3=Zhong |first3=T. |last4=Xiao |first4=L. | date = April 1994 | title = Report on the distribution, population, and ecology of the Yunnan snub-nosed monkey (''Rhinopithecus bieti'') | journal = Primates | volume = 35 | issue = 2 | pages = 241–250 | doi = 10.1007/BF02382060|s2cid=23931368 }} the [[mountain gorilla]] can be found at 4,200 meters (13,200 ft) crossing the [[Virunga Mountains]],{{cite book | author = Schaller, G. B. | year = 1963 | title = The Mountain Gorilla: Ecology and Behavior | location = Chicago | publisher = University Chicago Press | isbn = 978-0-226-73635-8 | url = https://archive.org/details/mountaingorillae00scha}} and the gelada has been found at elevations of up to {{Convert|5,000|m|ft|abbr=on}} in the [[Ethiopian Highlands]].{{cite book |author=Stammbach, E. |year=1987 |contribution=Desert, Forest, and Montane Baboons: Multilevel-Societies |pages=112–120 |editor=Smuts, B |editor2=Cheney, D |editor3=Seyfarth, R |editor4=Wrangham, R |editor5=Struhsaker, T. |title=Primate Societies |publisher=The University of Chicago Press |isbn=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]].{{cite journal|author=Kemp, E.|year=2009|title=Patterns of Water Use in Primates|journal=Folia Primatologica|volume=80|issue=4|pages=275–294|doi=10.1159/000252586|pmid=19864919|s2cid=5108827}} Some primates, such as the [[rhesus macaque]] and gray langurs, can exploit human-modified environments and even live in cities.{{cite book|title=Primates in Perspective|author1=Wolfe, L. D. |author2=Fuentes, A. |name-list-style=amp |chapter=Ethnoprimatology|year=2007|publisher=Oxford University Press|editor=Campbell, C. J. |editor2=Fuentes, A. |editor3=MacKinnon, K. C. |editor4=Panger, M. |editor5=Bearder, S. K.|isbn=978-0-19-517133-4|pages=692}} [353] => [354] => ==Interactions between humans and other primates== [355] => [356] => ===Disease transmission=== [357] => Close interactions between humans and non-human primates (NHPs) can create pathways for the transmission of [[Zoonosis|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.{{cite journal |author1=Renquist, D. M. |author2=Whitney, R. A. |name-list-style=amp |year=1987 |title=Zoonoses Acquired from Pet Primates |journal=Veterinary Clinics of North America: Small Animal Practice |volume=17 |issue=1 |pages=219–240 |url=http://pin.primate.wisc.edu/aboutp/pets/zoonoses.html |access-date=2008-08-11 |doi=10.1016/s0195-5616(87)50614-3|pmid=3551307 }} [358] => [359] => ===Legal and social status=== [360] => {{further|Great ape personhood}} [361] => [[File:Slow Loris Female.jpg|thumb|left|[[Slow loris]]es are popular in the [[exotic pet]] trade, which threatens wild populations.]] [362] => Only [[human]]s 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.{{cite web|url=https://www.un.org/Overview/rights.html |year=1948 |access-date=2008-12-02 |title=The Universal Declaration of Human Rights |publisher=[[United Nations]] |url-status=dead |archive-url=https://web.archive.org/web/20141208080853/http://www.un.org/Overview/rights.html |archive-date=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 [[great ape personhood|award at least some of them legal rights]].{{cite web|author1=Cavalieri, P. |author2=Singer, P. |name-list-style=amp |url=http://www.greatapeproject.org/declaration.php |title=Declaration on Great Apes |publisher=[[Great Ape Project]] |access-date=2008-06-16 |url-status=dead |archive-url=https://web.archive.org/web/20080820011354/http://www.greatapeproject.org/declaration.php |archive-date=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 [[chimpanzee]]s, [[orangutan]]s and [[gorilla]]s are not to be used for animal experiments.{{cite news |author=Glendinning, L. |url=https://www.theguardian.com/world/2008/jun/26/humanrights.animalwelfare?gusrc=rss&feed=networkfront |title=Spanish parliament approves 'human rights' for apes |newspaper=The Guardian |date=26 June 2008 |access-date=2008-11-10}}{{cite news |author=Singer, P. |url=https://www.theguardian.com/commentisfree/2008/jul/18/animalwelfare.animalbehaviour |title=Of great apes and men |newspaper=The Guardian |date=18 July 2008 |access-date=2008-11-10}} [363] => [364] => 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.{{cite magazine|title=The Perils of Keeping Monkeys as Pets |url=http://news.nationalgeographic.co.uk/news/2003/09/0916_030916_primatepets.html |first=M. |last=Mott |magazine=National Geographic |date=16 September 2003 |access-date=2013-02-06 |url-status=dead |archive-url=https://web.archive.org/web/20130608041322/http://news.nationalgeographic.co.uk/news/2003/09/0916_030916_primatepets.html |archive-date=2013-06-08}} The expanding Chinese middle class has increased demand for NHPs as exotic pets in recent years.{{cite journal |author=Workman, C. |date=June 2004 |title=Primate conservation in Vietnam: toward a holistic environmental narrative |url=https://archive.org/details/sim_american-anthropologist_2004-06_106_2/page/346 |journal=American Anthropologist |volume=106 |issue=2 |pages=346–352 |doi=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 [[United States dollar|US$]]3000 for monkeys to $30,000 for apes.{{cite web|publisher=[[International Primate Protection League]] |year=2003 |title=IPPL News: The US Pet Monkey Trade |url=http://www.aesop-project.org/US_Pet_Monkey_Trade.htm |access-date=2008-08-04 |url-status=dead |archive-url=https://web.archive.org/web/20080725190533/http://www.aesop-project.org/US_Pet_Monkey_Trade.htm |archive-date=2008-07-25}} [365] => [366] => Primates are used as [[model organism]]s in laboratories and have been used in [[Monkeys in space|space missions]].{{cite web |url=https://history.nasa.gov/afspbio/part1.htm |title=The Beginnings of Research in Space Biology at the Air Force Missile Development Center, 1946–1952 |access-date=2008-08-18 |author=Bushnell, D. |year=1958 |work=History of Research in Space Biology and Biodynamics |publisher=[[NASA]] |archive-date=2008-01-25 |archive-url=https://web.archive.org/web/20080125044753/https://history.nasa.gov/afspbio/part1.htm |url-status=dead }} They serve as [[service animal]]s for disabled humans. [[Capuchin monkey]]s can be [[Monkey helper|trained]] to assist [[quadriplegic]] humans; their intelligence, memory, and manual dexterity make them ideal helpers.{{cite news|title=Monkeys as Helpers To Quadriplegics At Home |author=Blumenthal, D.|date=1987-06-17|work=The New York Times |url=https://query.nytimes.com/gst/fullpage.html?sec=health&res=9B0DE5D81231F934A25755C0A961948260 |access-date=2008-10-08}} [367] => [368] => NHPs are kept in [[zoo]]s 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.{{cite book|author=Newman, James L.|year=2013|title=Encountering Gorillas: A Chronicle of Discovery, Exploitation, Understanding, and Survival|url=https://archive.org/details/encounteringgori0000newm|url-access=registration|publisher=Rowman and Littlefield|location=Plymouth, United Kingdom|isbn=978-1-4422-1957-1|page=[https://archive.org/details/encounteringgori0000newm/page/173 173]}} [369] => [370] => ===Role in scientific research=== [371] => {{Further|Animal testing on non-human primates|International trade in primates}} [372] => [[File:Monkey Sam Before The Flight On Little Joe 2.jpg|thumb|right|upright|Sam, a [[rhesus macaque]], was flown to the edge of space by [[NASA]] in the 1959 [[Little Joe 2]] flight of [[Project Mercury]].]] [373] => Thousands of non-human primates are used around the world in research because of their psychological and physiological similarity to humans.{{cite journal | last1=Chen |first1=F. C. |last2=Li |first2=W. H. | date = February 2001 | title = Genomic divergences between humans and other hominoids and the effective population size of the common ancestor of humans and chimpanzees | journal = [[American Journal of Human Genetics]] | volume = 68 | issue = 2 | pages = 444–456 | doi = 10.1086/318206 | pmid = 11170892 | pmc = 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 [[Clinical trial#Pre-clinical studies|preclinical trials]], [[neuroscience]], [[ophthalmology]] studies, and toxicity studies. [[Rhesus macaque]]s are often used, as are other [[macaque]]s, African [[Chlorocebus|green monkeys]], chimpanzees, [[baboon]]s, [[squirrel monkey]]s, and [[marmoset]]s, both wild-caught and purpose-bred.{{cite web|url=http://www.ebra.org/ebrabulletin-the-supply-and-use-of-primates-in-the-eu_17.htm |title=The supply and use of primates in the EU |publisher=European Biomedical Research Association |year=1996 |access-date=2008-08-18 |url-status=dead |archive-url=https://web.archive.org/web/20120117061036/http://www.ebra.org/ebrabulletin-the-supply-and-use-of-primates-in-the-eu_17.htm |archive-date=2012-01-17}}{{cite journal | last1=Conlee |first1=K. M. |last2=Hoffeld |first2=E. H. |last3=Stephens |first3=M. L. | year = 2004 | title = A Demographic Analysis of Primate Research in the United States | journal = Alternatives to Laboratory Animals | volume = 32 | issue = Sup 1 | pages = 315–322 |doi=10.1177/026119290403201s52 |pmid=23577480|s2cid=6743973 |doi-access=free }} [374] => [375] => 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.{{cite book |author= Presented to Parliament by the Secretary of State for the Home Department by Command of Her Majesty|title=Statistics of scientific procedures on living animals: Great Britain 2005 |date=July 2006 |publisher=[[The Stationery Office]] |isbn=0-10-168772-9 |url=http://www.official-documents.gov.uk/document/cm68/6877/6877.pdf |access-date=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.{{cite web|title=Nonhuman Primates: Research Animals |publisher=United States Department of Agriculture |work=Animal Welfare Information Center |url=http://awic.nal.usda.gov/nal_display/index.php?info_center=3&tax_level=3&tax_subject=169&topic_id=1078&level3_id=5345&level4_id=0&level5_id=0&placement_default=0 |access-date=2008-07-14 |url-status=dead |archive-url=https://web.archive.org/web/20081003004620/http://awic.nal.usda.gov/nal_display/index.php?info_center=3&tax_level=3&tax_subject=169&topic_id=1078&level3_id=5345&level4_id=0&level5_id=0&placement_default=0 |archive-date=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.{{cite web|url=http://www.eceae.org/a1_directive.php |title=Directive 86/609 |publisher=European Coalition to End Animal Experiments |access-date=2008-10-08 |url-status=dead |archive-url=https://web.archive.org/web/20080929214536/http://www.eceae.org/a1_directive.php |archive-date=2008-09-29}} [376] => [377] => ===Extinction threats=== [378] => [[File:Hunted Silky Sifakas.jpg|thumb|left|Humans are known to hunt other primates for food, called [[bushmeat]]. Pictured are two men who have killed a number of [[silky sifaka]] and [[white-headed lemur|white-headed brown]] lemurs.]] [379] => 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.{{Cite journal|last1=Estrada|first1=Alejandro|last2=Garber|first2=Paul A.|last3=Rylands|first3=Anthony B.|last4=Roos|first4=Christian|last5=Fernandez-Duque|first5=Eduardo|last6=Fiore|first6=Anthony Di|last7=Nekaris|first7=K. Anne-Isola|last8=Nijman|first8=Vincent|last9=Heymann|first9=Eckhard W.|date=2017-01-01|title=Impending extinction crisis of the world's primates: Why primates matter|journal=Science Advances|language=en|volume=3|issue=1|pages=e1600946|doi=10.1126/sciadv.1600946|issn=2375-2548|pmc=5242557|pmid=28116351|bibcode=2017SciA....3E0946E}} Additionally, 75% of primate species have decreasing populations. Trade is regulated, as all species are listed by [[CITES]] in [[CITES#Appendix II|Appendix II]], except 50 species and subspecies listed in [[CITES#Appendix I|Appendix I]], which gain full protection from trade.{{cite book|author=IFAW |title=Born to be wild: Primates are not pets |publisher=International Fund for Animal Welfare |year=2005 |url=http://www.ifaw.org/Publications/Program_Publications/Wildlife_Trade/Campaign_Scientific_Publications/asset_upload_file812_49478.pdf |access-date=2011-02-26 |author-link=International Fund for Animal Welfare |url-status=dead |archive-url=https://web.archive.org/web/20110726171340/http://www.ifaw.org/Publications/Program_Publications/Wildlife_Trade/Campaign_Scientific_Publications/asset_upload_file812_49478.pdf |archive-date=2011-07-26}}{{cite web | author = CITES | title = Appendices I, II and III | publisher = Convention on International Trade in Endangered Species of Wild Fauna and Flora | date = 2010-10-14 | url = http://www.cites.org/eng/app/index.php | access-date = 2012-04-02 |author-link=CITES}} [380] => [[File:Lemur poaching 004.jpg|alt=White-fronted Brown Lemur (Eulemur albifrons) killed in northeast Madagascar for bushmeat.|thumb|[[White-headed lemur]] (''Eulemur albifrons'') killed in northeast [[Madagascar]] for bushmeat.]] [381] => Common threats to primate species include [[deforestation]], [[forest fragmentation]], [[monkey drive]]s (resulting from primate crop raiding),{{cite book |author=Grubb, P. |year=1998 |chapter=The Sierra Leone monkey drives |title=Mammals of Ghana, Sierra Leone, and the Gambia |location=St. Ives |publisher=Trendrine |pages=214–219 |isbn=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.{{cite journal |last1=Chapman |first1=C. A. |last2=Peres |first2=C. A. |year=2001 |title=Primate conservation in the new millennium: the role of scientists |journal=Evolutionary Anthropology |volume=10 |issue=1 |pages=16–33 |doi=10.1002/1520-6505(2001)10:1<16::AID-EVAN1010>3.0.CO;2-O|s2cid=13494366 }}{{cite book |author1=Mittermeier, R. A. |author2=Cheney, D. L. |name-list-style=amp |year=1987 |chapter=Conservation of primates and their habitats |title=Primate Societies |editor=Smuts, B. B. |editor2=Cheney, D. L. |editor3=Seyfarth, R. M. |editor4=Wrangham, R. W. |editor5= Struhsaker, T. T. |publisher=University of Chicago Press |location=Chicago |pages=477–490}}{{cite journal|author1=Southwick, C. H. |author2=Siddiqi, M. F. |name-list-style=amp |year=2001 |title=Status, conservation and management of primates in India |journal=Envis Bulletin: Wildlife and Protected Areas |volume=1 |issue=1 |pages=81–91 |url=http://www.wii.gov.in/envis/primates/downloads/page81statusofprimates.pdf |archive-url=https://wayback.archive-it.org/all/20081001191112/http://wii.gov.in/envis/primates/downloads/page81statusofprimates.pdf |url-status=dead |archive-date=2008-10-01 |access-date=2008-08-04}} More than 90% of primate species occur in tropical forests.{{cite book |author1=Cowlishaw, G. |author2=Dunbar, R. |name-list-style=amp |year=2000 |title=Primate Conservation Biology |publisher=University of Chicago Press |location=Chicago |isbn=978-0-226-11637-2}} The main cause of forest loss is clearing for agriculture, although commercial logging, [[Subsistence agriculture|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 orangutan]]s per year in the [[Leuser Ecosystem]] alone.{{cite journal |last1=Van Schaik |first1=C. P. |last2=Monk |first2=K. A. |last3=Robertson |first3=J. M. Y. |year=2001 |title=Dramatic decline in orangutan numbers in the Leuser Ecosystem, northern Sumatra |journal=Oryx |volume=35 |issue=1 |pages=14–25 |doi=10.1046/j.1365-3008.2001.00150.x|doi-access=free }} [382] => [[File:Silky Sifaka Pink Face Closeup.JPG|thumb|right|The critically endangered [[silky sifaka]]]] [383] => Primates with a large body size (over 5 kg) are at increased extinction risk due to their greater profitability to [[poaching|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.{{cite journal |last1=Purvis |first1=A. |last2=Gittleman |first2=J. L. |last3=Cowlishaw |first3=G. |last4=Mace |first4=G. M. |year=2000 |title=Predicting extinction risk in declining species |journal=Proceedings of the Royal Society B |volume=267 |pages=1947–1952 |doi=10.1098/rspb.2000.1234 |pmid=11075706 |issue=1456 |pmc=1690772}} Data for some African cities show that half of all protein consumed in urban areas comes from the [[bushmeat]] trade.{{cite journal |last1=Fa |first1=J. E. |last2=Juste |first2=J. |last3=Perez de Val |first3=J. |last4=Castroviejo |first4=J. |year=1995 |title=Impact of market hunting on mammal species in Equatorial Guinea |url=https://archive.org/details/sim_conservation-biology_1995-10_9_5/page/1107 |journal=Conservation Biology |volume=9 |issue=5 |pages=1107–1115 |doi=10.1046/j.1523-1739.1995.9051107.x|pmid=34261280 |hdl=10261/49187 |hdl-access=free }} Endangered primates such as [[guenon]]s and the [[Drill (mammal)|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.{{cite journal |author=Hill, C. M. |year=1997 |title=Crop-raiding by wild vertebrates: The farmer's perspective in an agricultural community in western Uganda |journal=International Journal of Pest Management |volume=43 |issue=1 |pages=77–84 |doi=10.1080/096708797229022}} Primate crop raiding gives locals a negative impression of primates, hindering conservation efforts.{{cite journal |author=Hill, C. M. |year=2002 |title=Primate conservation and local communities: Ethical issues and debates |url=https://archive.org/details/sim_american-anthropologist_2002-12_104_4/page/1184 |journal=American Anthropologist |volume=104 |issue=4 |pages=1184–1194 |doi=10.1525/aa.2002.104.4.1184|doi-access=free }} [384] => [385] => [[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.{{cite encyclopedia | title = Primate | encyclopedia = Encyclopædia Britannica Online | publisher = [[Encyclopædia Britannica, Inc.]] | year = 2008 | url=http://www.britannica.com/EBchecked/topic/476264/primate | access-date=2008-07-21}} Among the primates wiped out were ''[[Archaeoindris]]'' (a lemur larger than a silverback gorilla) and the families [[Palaeopropithecidae]] and [[Archaeolemuridae]]. [386] => [[File:Man of the woods.JPG|thumb|left|The [[critically endangered]] Sumatran orangutan]] [387] => In 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.{{cite journal|author=Choudhury, A. |year=2001 |title=Primates in Northeast India: an overview of their distribution and conservation status |journal=Envis Bulletin: Wildlife and Protected Areas |volume=1 |issue=1 |pages=92–101 |url=http://www.wii.gov.in/envis/primates/downloads/page92primatesne.pdf |archive-url=https://wayback.archive-it.org/all/20081001191106/http://wii.gov.in/envis/primates/downloads/page92primatesne.pdf |url-status=dead |archive-date=2008-10-01 |access-date=2008-08-04}}{{cite journal |last1=Kumara |first1=H. N. |last2=Singh |first2=M. |date=October 2004 |title=Distribution and abundance of primates in rainforests of the Western Ghats, Karnataka, India and the conservation of ''Macaca silenus'' |journal=International Journal of Primatology |volume=25 |issue=5 |pages=1001–1018 |doi=10.1023/B:IJOP.0000043348.06255.7f|s2cid=30384142 }} The pet trade and traditional medicine also increase demand for illegal hunting.{{cite journal |author=Nijman, V. |year=2004 |title=Conservation of the Javan gibbon ''Hylobates moloch'': population estimates, local extinction, and conservation priorities |journal=The Raffles Bulletin of Zoology |volume=52 |issue=1 |pages=271–280 |url=http://rmbr.nus.edu.sg/rbz/biblio/52/52rbz271-280.pdf |access-date=2008-08-04 |url-status=dead |archive-url=https://web.archive.org/web/20080910043814/http://rmbr.nus.edu.sg/rbz/biblio/52/52rbz271-280.pdf |archive-date=2008-09-10}}{{cite journal |last1=O'Brien |first1=T. G. |last2=Kinnaird |first2=M. F. |last3=Nurcahyo |first3=A. |last4=Iqbal |first4=M. |last5=Rusmanto |first5=M. |date=April 2004 |title=Abundance and distribution of sympatric gibbons in a threatened Sumatran rain forest |journal=International Journal of Primatology |volume=25 |issue=2 |pages=267–284 |doi=10.1023/B:IJOP.0000019152.83883.1c|s2cid=32472118 }} 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. [388] => [389] => 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.{{cite journal |last1=Estrada |first1=A. |last2=Coates-Estrada |first2=R. |last3=Meritt |first3=D. |date=September 1994 |title=Non-flying mammals and landscape changes in the tropical forest region of Los Tuxtlas, Mexico |journal=Ecography |volume=17 |pages=229–241 |doi=10.1111/j.1600-0587.1994.tb00098.x |issue=3}} This increases the amount of forest vulnerable to [[edge effect]]s such as farmland encroachment, lower levels of humidity and a change in plant life.{{cite book |author=Marsh, L. K. |year=2003 |chapter=The nature of fragmentation. |title=Primates in Fragments: Ecology and Conservation |editor=Marsh, L. K. |location=New York |publisher=[[Kluwer Academic]]/[[Plenum Publishers]] |pages=1–10 |isbn=0-306-47696-7}}{{cite journal |author=Turner, I. M. |year=1996 |title=Species loss in fragments of tropical rain forest: a review of the evidence |url=https://archive.org/details/sim_journal-of-applied-ecology_1996-04_33_2/page/200 |journal=Journal of Applied Ecology |volume=33 |pages=200–209 |doi=10.2307/2404743 |issue=2 |jstor=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.{{cite book |author=Chiarello, A.G. |year=2003 |chapter=Primates of the Brazilian Atlantic forest: the influence of forest fragmentation on survival |title=Primates in Fragments: Ecology and Conservation |editor=Marsh, L. K. |location=New York |publisher=Kluwer Academic/Plenum Publishers |pages=99–121 |isbn=978-0-306-47696-9}}{{cite book |author=Pope, T.R. |year=1996 |chapter=Socioecology, population fragmentation, and patterns of genetic loss in endangered primates |title=Conservation Genetics: Case Histories from Nature |url=https://archive.org/details/conservationgene0000unse |editor=Avise, J. |editor2=Hamrick, J. |location=Norwell |publisher=Kluwer Academic Publishers |pages=[https://archive.org/details/conservationgene0000unse/page/119 119]–159 |isbn=978-0-412-05581-2}} [390] => [391] => 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]].{{cite book | editor1-last = Mittermeier | editor1-first = R.A. | editor2-last = Wallis | editor2-first = J. | editor3-last = Rylands | editor3-first = A.B. | editor4-last = Ganzhorn | editor4-first = J.U. | editor5-last = Oates | editor5-first = J.F. | editor6-last = Williamson | editor6-first = E.A. | editor7-last = Palacios | editor7-first = E. | editor8-last = Heymann | editor8-first = E.W. | editor9-last = Kierulff | editor9-first = M.C.M. | editor10-last = Yongcheng | editor10-first = L. | editor11-last = Supriatna | editor11-first = J. | editor12-last = Roos | editor12-first = C. | editor13-last = Walker | editor13-first = S. | editor14-last = Cortés-Ortiz | editor14-first = L. | editor15-last = Schwitzer | editor15-first = C. | others = Illustrated by S.D. Nash | editor-link = Russell Mittermeier | year = 2009 | title = Primates in Peril: The World's 25 Most Endangered Primates 2008–2010 | publisher = IUCN/SSC Primate Specialist Group (PSG), International Primatological Society (IPS), and Conservation International (CI) | location = Arlington, VA. | pages = 23–26 | isbn = 978-1-934151-34-1 | url = http://www.primate-sg.org/storage/PDF/Primates.in.Peril.2008-2010.pdf}} [[Miss Waldron's red colobus]] was recently declared extinct when no trace of the subspecies could be found from 1993 to 1999.{{cite journal |last1=Oates |first1=J. F. |last2=Abedi-Lartey |first2=M. |last3=McGraw |first3=W. S. |last4=Struhsaker |first4=T. T. |last5=Whitesides |first5=G. H. | title=Extinction of a West African Red Colobus Monkey | journal=Conservation Biology | date=October 2000 | volume=14 | issue=5 | doi=10.1046/j.1523-1739.2000.99230.x | pages=1526–1532|s2cid=84336872 }} A few hunters have found and killed individuals since then, but the subspecies' prospects remain bleak.{{cite journal |title=Update on the Search for Miss Waldron's Red Colobus Monkey |author=McGraw, W. S. |journal=International Journal of Primatology |volume=26 |issue=3 |date=June 2005 |pages=605–619 |doi=10.1007/s10764-005-4368-9|s2cid=861418 }} [392] => [393] => ==See also== [394] => {{Portal|Animals|Mammals|Primates}} [395] => {{div col|colwidth=18em}} [396] => * [[Arboreal theory]] [397] => * [[Great Ape Project]] [398] => * [[Human evolution]] [399] => * [[International Primate Day]] [400] => * [[List of primates]] [401] => * [[List of fossil primates]] [402] => * [[Monkey Day]] [403] => * [[Primatology]] [404] => {{div col end}} [405] => [406] => ==Footnotes== [407] => {{Reflist|group=lower-alpha}} [408] => [409] => ==References== [410] => {{Reflist}} [411] => [412] => ===Literature cited=== [413] => * {{cite book | last = Benton| first = Michael J. | chapter = Chapter 3: Primate evolution | title = Vertebrate palaeontology | publisher = Wiley-Blackwell | year = 2005 | isbn = 978-0-632-05637-8 | chapter-url=https://books.google.com/books?id=SyJO3vpCk8AC |access-date=2011-07-10 }} [414] => * {{cite book | last1 = Cartmill | first1 = M. | year = 2010 | title = Primate Neuroethology | editor1-last = Platt | editor1-first = M. | editor2-last = Ghazanfar | editor2-first = A | chapter = Primate Classification and Diversity | pages = 10–30 | publisher = Oxford University Press | isbn = 978-0-19-532659-8 | chapter-url = https://books.google.com/books?id=hv28p1tCnnEC&pg=PA15 }} [415] => * {{cite book | last1 = Hartwig | first1 = W. | editor1-last = Campbell | chapter = Chapter 3: Primate evolution | pages = 19–31 | editor1-first = C. J. | editor2-last = Fuentes | editor2-first = A. | editor3-last = MacKinnon | editor3-first = K. C. | editor4-last = Bearder | editor4-first = S. K. | editor5-last=Stumpf | editor5-first = R. M | title = Primates in Perspective | edition = 2nd | publisher = Oxford University Press | year = 2011 | isbn = 978-0-19-539043-8 }} [416] => * {{cite book | last1 = Szalay | first1 = F.S. | last2 = Delson | first2 = E. | title = Evolutionary History of the Primates | year = 1980 | publisher = [[Academic Press]] | isbn = 978-0126801507 | oclc = 893740473 | url = https://books.google.com/books?id=jE7gBAAAQBAJ&pg=PA149 }} [417] => [418] => ==Further reading== [419] => {{Library resources box|onlinebooks=yes|by=no|lcheading= Primates|label=Primates}} [420] => * {{Cite book |year=1984 |editor1=David J. Chivers |editor2=Bernard A. Wood |editor3=Alan Bilsborough |title=Food Acquisition and Processing in Primates |place=New York & London |publisher=Plenum Press |isbn=0-306-41701-4}} [421] => [422] => ==External links== [423] => {{EB1911 Poster|Primates}} [424] => *{{Curlie|Science/Biology/Flora_and_Fauna/Animalia/Chordata/Mammalia/Primates/}} [425] => * [https://primate.wisc.edu/primate-info-net/ Primate Info Net] [426] => * [https://animaldiversity.org/site/accounts/information/Primates.html Primates] at [[Animal Diversity Web]] [427] => * [http://www.pri.kyoto-u.ac.jp/ Primate Research Institute], [[Kyoto University]] [428] => * [https://web.archive.org/web/20191011040440/http://primate-brain.org/ High-Resolution Cytoarchitectural Primate Brain Atlases] [429] => * [http://www.euprim-net.eu EUPRIM-Net: European Primate Network] [430] => * [https://search.library.wisc.edu/digital/APCLNatHist PrimateImages: Natural History Collection] [431] => * [http://eskeletons.org/ Interactive views] of various primate skeletons at eSkeletons.org (associated with the [[University of Texas at Austin]]) [432] => * [http://tolweb.org/Primates/15963 Tree of Life web project] [433] => [434] => {{Mammals}} [435] => {{Primates}} [436] => {{Apes}} [437] => {{Taxonbar|from=Q7380}} [438] => {{Authority control}} [439] => [440] => {{Featured article}} [441] => [442] => [[Category:Primates| ]] [443] => [[Category:Extant Thanetian first appearances]] [444] => [[Category:Taxa named by Carl Linnaeus]] [] => )
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Primate

The Wikipedia page for "Primate" provides a comprehensive overview of the order of mammals known as primates. Primates are characterized by their advanced cognitive abilities, grasping hands and feet, and forward-facing eyes.

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Primates are characterized by their advanced cognitive abilities, grasping hands and feet, and forward-facing eyes. They are widely distributed across the globe, including regions in Africa, Asia, and the Americas. The page begins by discussing the classification of primates and provides information on the various suborders, families, and genera within this order. It also highlights the three main groups of primates based on their anatomical and genetic distinctions: prosimians, monkeys, and apes. The article covers the evolutionary history of primates, tracing their origins back to about 65 million years ago. It explores the ancestral primates, including the common ancestors of humans and other primates, and delves into their adaptations and diversification over time. Furthermore, the page discusses the key characteristics of primates, such as their complex social structures, omnivorous diets, and reproductive strategies. It also explores their communication systems, including vocalizations and various forms of non-verbal communication. The article further provides an in-depth exploration of primate behavior, including their tool use, problem-solving abilities, and culture. It examines their cognitive capacities and highlights the research and studies conducted on primate intelligence. Additionally, the page covers various aspects of primate conservation, including the threats they face due to habitat destruction, hunting, and the illegal pet trade. It discusses conservation efforts, such as protected areas and captive breeding programs, which aim to preserve the vast diversity of primate species. The Wikipedia page for "Primate" is a valuable resource for those interested in the scientific study of primates, their characteristics, behavior, and conservation. It is well-referenced, providing a wealth of information on this fascinating and diverse group of animals.

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