The evolutionary history of the primates can be traced back 57-90 million years. [1] One of the oldest known primate-like mammal species, Plesiadapis , came from North America; [2] another, Archicebus , came from China. [3] Other similar basal primates were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and Eocene. Purgatorius is the genus of the four extinct species believed to be the earliest example of a primate or a proto-primate, a primatomorph precursor to the Plesiadapiformes, dating to as old as 66 million years ago.
The surviving tropical population of primates, which is seen most completely in the upper Eocene and lowermost Oligocene fossil beds of the Faiyum depression southwest of Cairo, gave rise to all living species—lemurs of Madagascar, lorises of Southeast Asia, galagos or "bush babies" of Africa, and the anthropoids: platyrrhine or New World monkeys, catarrhines or Old World monkeys, and the apes, including Homo sapiens .
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The origins and early evolution of primates is shrouded in mystery due to lack of fossil evidence. They are believed to have split from plesiadapiforms in Eurasia around the early Eocene or earlier. The first true primates so far found in the fossil record are fragmentary and already demonstrate the major split between strepsirrhines and haplorines.
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The earliest strepsirrhines are known as adapiforms, a diverse group that ranged throughout Eurasia and North America. An early branch of this clade gave rise to lemuriform primates, which includes lemurs and their kin.
David Begun has theorised that early primates flourished in Eurasia and that a lineage leading to the African apes and humans, including Dryopithecus , migrated south from Europe or Western Asia into Africa. [4] However, the early European fauna is exemplified by Darwinius , a basal strepsirrhine dated to 47 million years (early Eocene) [5]
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The earliest haplorrhine primates from the fossil record are the omomyids, which resembled modern day tarsiers. Like the strepsirrhine adapiforms, omomyids were diverse and ranged throughout Eurasia and North America. The phylogeny of omomyids, tarsiers, and simians is currently unknown.
For many years, it was assumed that primates had first evolved in Africa, and this assumption and the excavations that resulted from it yielded many early simian fossils that chronicled their evolution. Due to the lack of fossils linking simians to the earliest haplorrhines, a more recently discovered stem group called eosimiids found in Asia are thought to have dispersed to Africa and evolved into simians. Eosimiids were very small and similar to tarsiers, though their dentition more closely resembles that of simians.
Some of the primates' vertebrate ancestors were tetrachromats, but their nocturnal mammalian ancestors lost two of their four cones during the mesozoic. Most modern primates, however, have evolved to be trichromats. All old world monkeys and apes are trichromats, but new world monkeys are polymorphic trichromats, meaning that males and homozygous females are dichromats while heterozygous females are trichromats (with the exceptions of howler monkeys and night monkeys, who have more and less robust color vision respectively).
There are four prevailing theories as to what the evolutionary pressure was for primates to develop trichromatic vision. The Fruit Theory suggests that it was easier for trichromatic primates to find ripe fruit against a green background. While there is data supporting the Fruit Theory, there is some dispute about whether or not trichromacy was more advantageous for determining how ripe fruit was up close or spotting fruit from afar. The Young Leaf hypothesis suggests that primates with more advanced color vision could better spot younger and more nutritious leaves during fruit shortages, while there are also theories that suggest more advanced color vision was better for recognizing changes in skin tone, allowing primates to better determine the blood oxygen saturation of others. Still other theories suppose that primates' color vision evolved alongside their sense of smell, though research has shown no direct correlation between concentration of olfactory receptors and acquisition of color vision.
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Following the emergence of Simiformes in Africa, Platyrrhini split from Catarrhini during the Eocene when New World monkeys dispersed to South America, likely by rafting on mats of vegetation across the Atlantic Ocean. The Atlantic is estimated to have been possibly 1,000 km (600 mi) narrower, based on estimates from the expansion of the Atlantic mid-ocean ridge formation processes (25 cm/1 in per year.) It is also possible that during this rafting process, there were a number of islands between Africa and South America which have since been submerged.
Bayesian estimates of divergence time using "conservative but realistic fossil constraints" have indicated the most recent common ancestor of new world monkeys to have existed between 27-31 million years ago. [6]
The earliest known catarrhine is Kamoyapithecus from uppermost Oligocene at Eragaleit in the northern Kenya Rift Valley, dated to 24 million years ago. [7] Its ancestry is thought to be species related to Aegyptopithecus , Propliopithecus , and Parapithecus from the Faiyum depression, at around 35 million years ago. [8] In 2010, Saadanius was described as a close relative of the last common ancestor of the crown catarrhines, and tentatively dated to 29–28 million years ago, helping to fill an 11-million-year gap in the fossil record. [9] Notable species also include Nsungwepithecusgunnelli and Rukwapithecus fleaglei of the Oligocene. [10]
In the early Miocene, about 22 million years ago, the many kinds of arboreally adapted primitive catarrhines from East Africa suggest a long history of prior diversification. Fossils dated to be 20 million years old include fragments attributed to Victoriapithecus , believed to be the earliest Old World monkey.[ citation needed ] Among the genera thought to be in the ape lineage leading up to 13 million years ago are Proconsul , Rangwapithecus , Dendropithecus , Limnopithecus , Nacholapithecus , Equatorius , Nyanzapithecus , Afropithecus , Heliopithecus, and Kenyapithecus , all from East Africa.
The presence of other generalized non-cercopithecids of the middle Miocene age from sites far distant— Otavipithecus from cave deposits in Namibia, and Pierolapithecus and Dryopithecus from France, Spain and Austria—is evidence of a wide diversity of forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the early and middle Miocene. The youngest of the Miocene hominoids, Oreopithecus , is from coal beds in Italy that have been dated to 9 million years ago.
Molecular evidence indicates that the lineage of gibbons (family Hylobatidae) diverged from Great Apes some 18–12 million years ago, and that of orangutans (subfamily Ponginae) diverged from the other Great Apes at about 12 million years; there are no fossils that clearly document the ancestry of gibbons, which may have originated in a so-far-unknown South East Asian hominoid population, but fossil proto-orangutans may be represented by Sivapithecus from India and Griphopithecus from Turkey, dated to around 10 million years ago. [11]
Human evolution is the evolutionary process that led to the emergence of anatomically modern humans, beginning with the evolutionary history of primates – in particular genus Homo – and leading to the emergence of Homo sapiens as a distinct species of the hominid family, the great apes. This process involved the gradual development of traits such as human bipedalism and language. [12]
The study of human evolution involves many scientific disciplines, including physical anthropology, primatology, archaeology, paleontology, neurobiology, ethology, linguistics, evolutionary psychology, embryology and genetics. [13] Genetic studies show that primates diverged from other mammals about 85 million years ago, in the Late Cretaceous period, and the earliest fossils appear in the Paleocene, around 55 million years ago. [14]
Within the superfamily Hominoidea (apes), the family Hominidae diverged from the family Hylobatidae (gibbons) some 15–20 million years ago; African great apes (subfamily Homininae) diverged from orangutans (Ponginae) about 14 million years ago; the tribe Hominini (humans, Australopithecines and other extinct biped genera, and chimpanzee) parted from the tribe Gorillini (gorillas) between 9 million years ago and 8 million years ago; and, in turn, the subtribes Hominina (humans and biped ancestors) and Panina (chimpanzees) separated about 7.5 million years ago to 5.6 million years ago. [15]
In primates, the pelvis consists of four parts—the left and the right hip bones which meet in the mid-line ventrally and are fixed to the sacrum dorsally and the coccyx. Each hip bone consists of three components, the ilium, the ischium, and the pubis, and at the time of sexual maturity these bones become fused together, though there is never any movement between them. In humans, the ventral joint of the pubic bones is closed.
The most striking feature of evolution of the pelvis in primates is the widening and the shortening of the blade called the ilium. Because of the stresses involved in bipedal locomotion, the muscles of the thigh move the thigh forward and backward, providing the power for bi-pedal and quadrupedal locomotion. [16]
Apes are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia. Apes are more closely related to Old World monkeys than to the New World monkeys (Platyrrhini) with both Old World monkeys and apes placed in the clade Catarrhini. Apes do not have tails due to a mutation of the TBXT gene. In traditional and non-scientific use, the term ape can include tailless primates taxonomically considered Cercopithecidae, and is thus not equivalent to the scientific taxon Hominoidea. There are two extant branches of the superfamily Hominoidea: the gibbons, or lesser apes; and the hominids, or great apes.
Old World monkeys are primates in the family Cercopithecidae. Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboons, red colobus, and macaques. Common names for other Old World monkeys include the talapoin, guenon, colobus, douc, vervet, gelada, mangabey, langur, mandrill, drill, surili (Presbytis), patas, and proboscis monkey.
The parvorder Catarrhini consists of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name Catarrhini, "Old World monkeys",. Its sister in the infraorder Simiiformes is the parvorder Platyrrhini. There has been some resistance to directly designate apes as monkeys despite the scientific evidence, so "Old World monkey" may be taken to mean the Cercopithecoidea or the Catarrhini. That apes are monkeys was already realized by Georges-Louis Leclerc, Comte de Buffon in the 18th century. Linnaeus placed this group in 1758 together with what we now recognise as the tarsiers and the New World monkeys, in a single genus "Simia". The Catarrhini are all native to Africa and Asia. Members of this parvorder are called catarrhines.
Strepsirrhini or Strepsirhini is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Collectively they are referred to as strepsirrhines. Also belonging to the suborder are the extinct adapiform primates which thrived during the Eocene in Europe, North America, and Asia, but disappeared from most of the Northern Hemisphere as the climate cooled. Adapiforms are sometimes referred to as being "lemur-like", although the diversity of both lemurs and adapiforms does not support this comparison.
Haplorhini, the haplorhines or the "dry-nosed" primates is a suborder of primates containing the tarsiers and the simians, as sister of the Strepsirrhini ("moist-nosed"). The name is sometimes spelled Haplorrhini. The simians include catarrhines, and the platyrrhines.
Tarsiiformes are a group of primates that once ranged across Europe, northern Africa, Asia, and North America, but whose extant species are all found in the islands of Southeast Asia. Tarsiers are the only living members of the infraorder; other members of Tarsiidae include the extinct Tarsius eocaenus from the Eocene, and Tarsius thailandicus from the Miocene. Two extinct genera, Xanthorhysis and Afrotarsius, are considered to be close relatives of the living tarsiers, and are generally classified within Tarsiiformes, with the former grouped within family Tarsiidae, and the latter listed as incertae sedis (undefined). Omomyids are generally considered to be extinct relatives, or even ancestors, of the living tarsiers, and are often classified within Tarsiiformes.
Lemuriformes is the sole extant infraorder of primate that falls under the suborder Strepsirrhini. It includes the lemurs of Madagascar, as well as the galagos and lorisids of Africa and Asia, although a popular alternative taxonomy places the lorisoids in their own infraorder, Lorisiformes.
The simians, anthropoids, or higher primates are an infraorder of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders Platyrrhini and Catarrhini, the latter of which consists of the family Cercopithecidae and the superfamily Hominoidea.
Aegyptopithecus is an early fossil catarrhine that predates the divergence between hominoids (apes) and cercopithecids. It is known from a single species, Aegyptopithecus zeuxis, which lived around 38-29.5 million years ago in the early part of the Oligocene epoch. It likely resembled modern-day New World monkeys, and was about the same size as a modern howler monkey, which is about 56 to 92 cm long. Aegyptopithecus fossils have been found in the Jebel Qatrani Formation of modern-day Egypt. Aegyptopithecus is believed to be a stem-catarrhine, a crucial link between Eocene and Miocene fossils.
Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, constitute an incomplete paraphyletic grouping; however, in the broader sense based on cladistics, apes (Hominoidea) are also included, making the terms monkeys and simians synonyms in regard to their scope.
The evolution of color vision in primates is highly unusual compared to most eutherian mammals. A remote vertebrate ancestor of primates possessed tetrachromacy, but nocturnal, warm-blooded, mammalian ancestors lost two of four cones in the retina at the time of dinosaurs. Most teleost fish, reptiles and birds are therefore tetrachromatic while most mammals are strictly dichromats, the exceptions being some primates and marsupials, who are trichromats, and many marine mammals, who are monochromats.
Altiatlasius is an extinct genus of mammal, which may have been the oldest known primate, dating to the Late Paleocene from Morocco. The only species, Altiatlasius koulchii, was described in 1990.
Algeripithecus is an extinct genus of early fossil primate, weighing approximately 65 to 85 grams. Fossils have been found in Algeria dating from 50 to 46 million years ago.
Lemurs, primates belonging to the suborder Strepsirrhini which branched off from other primates less than 63 million years ago, evolved on the island of Madagascar, for at least 40 million years. They share some traits with the most basal primates, and thus are often confused as being ancestral to modern monkeys, apes, and humans. Instead, they merely resemble ancestral primates.
Saadanius is a genus of fossil primates dating to the Oligocene that is closely related to the common ancestor of the Old World monkeys and apes, collectively known as catarrhines. It is represented by a single species, Saadanius hijazensis, which is known only from a single partial skull tentatively dated between 29 and 28 million years ago. It was discovered in 2009 in western Saudi Arabia near Mecca and was first described in 2010 after comparison with both living and fossil catarrhines.
Archicebus is a genus of fossil primates that lived in the early Eocene forests of what is now Jingzhou in the Hubei Province in central China, discovered in 2003. The only known species, A. achilles, was a small primate, estimated to weigh about 20–30 grams (0.7–1.1 oz), and is the only known member of the family Archicebidae. When discovered, it was the oldest fossil haplorhine primate skeleton found, appearing to be most closely related to tarsiers and the fossil omomyids, although A. achilles is suggested to have been diurnal, whereas tarsiers are nocturnal. Resembling tarsiers and simians, it was a haplorhine primate, and it also may have resembled the last common ancestor of all haplorhines.
Anapithecus is a late Miocene primate known from fossil locations in Hungary and Austria. Many Anapithecus fossils come from the site of Rudabánya, in northern Hungary, where Anapithecus lived alongside the ape Rudapithecus. The only species in the genus, Anapithecus hernyaki, is named after Gabor Hernyák, chief geologist of the Iron Ore Works of Rudabánya.
Amphipithecus mogaungensis was a primate that lived in Late Eocene Myanmar. Along with another primate Pondaungia cotteri, both are difficult to categorise within the order Primates. What little is known suggests that they are neither adapiform nor omomyid primates, two of the earliest primate groups to appear in the fossil record. Deep mandibles and mandibular molars with low, broad crowns suggest they are both simians, a group that includes monkeys, apes, and humans, though more material is needed for further comparison. The teeth also suggest that these were frugivore primates, with a body mass of 6–10 kg (13–22 lb).
Propliopithecoidea is a superfamily of catarrhine primates that inhabited Africa and the Arabian Peninsula during the Early Oligocene about 32 to 29 million years ago. Fossils have been found in Egypt, Oman and Angola. They are one of the earliest known families of catarrhines. They have a number of features in common with extant catarrhines, but also features that are primitive and not found in later catarrhine families.
The Amphipithecidae were simian primates that lived in Late Eocene and Early Oligocene. Fossils have been found in Myanmar, Thailand, and Pakistan. The limited fossil evidence is consistent with, but not exclusive to, arboreal quadrupedalism. In other words, the species may have moved about in trees on four legs, but not with regular leaping as seen in later simians.