Human evolution – Wikipedia

Human evolution, also known as hominization, 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 involves the gradually loss of typical animal characteristics and the development of exclusively human properties.[1]

The study of human evolution involves many scientific disciplines, including physical anthropology, primatology, archaeology, paleontology, neurobiology, ethology, linguistics, evolutionary psychology, embryology and genetics.[2] 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.[3]

Within the Hominoidea (apes) superfamily, the Hominidae family diverged from the Hylobatidae (gibbon) family some 1520 million years ago; African great apes (subfamily Homininae) diverged from orangutans (Ponginae) about 14 million years ago; the Hominini tribe (humans, Australopithecines and other extinct biped genera, and chimpanzee) parted from the Gorillini tribe (gorillas) between 9 million years ago and 8 million years ago; and, in turn, the subtribes Hominina (humans and biped ancestors) and Panina (chimps) separated about 7.5 million years ago to 5.6 million years ago.[4]

Human evolution from its first separation from the last common ancestor of humans and chimpanzees is characterized by a number of morphological, developmental, physiological, and behavioral changes. The most significant of these adaptations are bipedalism, increased brain size, lengthened ontogeny (gestation and infancy), and decreased sexual dimorphism. The relationship between these changes is the subject of ongoing debate.[5][pageneeded] Other significant morphological changes included the evolution of a power and precision grip, a change first occurring in H. erectus.[6]

Bipedalism is the basic adaptation of the hominid and is considered the main cause behind a suite of skeletal changes shared by all bipedal hominids. The earliest hominin, of presumably primitive bipedalism, is considered to be either Sahelanthropus[7] or Orrorin, both of which arose some 6 to 7 million years ago. The non-bipedal knuckle-walkers, the gorilla and chimpanzee, diverged from the hominin line over a period covering the same time, so either of Sahelanthropus or Orrorin may be our last shared ancestor. Ardipithecus, a full biped, arose somewhat later.[citation needed]

The early bipeds eventually evolved into the australopithecines and still later into the genus Homo. There are several theories of the adaptation value of bipedalism. It is possible that bipedalism was favored because it freed the hands for reaching and carrying food, saved energy during locomotion,[8] enabled long distance running and hunting, provided an enhanced field of vision, and helped avoid hyperthermia by reducing the surface area exposed to direct sun; features all advantageous for thriving in the new savanna and woodland environment created as a result of the East African Rift Valley uplift versus the previous closed forest habitat.[9][8][10] A new study provides support for the hypothesis that walking on two legs, or bipedalism, evolved because it used less energy than quadrupedal knuckle-walking.[11][12] However, recent studies suggest that bipedality without the ability to use fire would not have allowed global dispersal.[13] This change in gait saw a lengthening of the legs proportionately when compared to the length of the arms, which were shortened through the removal of the need for brachiation. Another change is the shape of the big toe. Recent studies suggest that Australopithecines still lived part of the time in trees as a result of maintaining a grasping big toe. This was progressively lost in Habilines.

Anatomically, the evolution of bipedalism has been accompanied by a large number of skeletal changes, not just to the legs and pelvis, but also to the vertebral column, feet and ankles, and skull.[14] The femur evolved into a slightly more angular position to move the center of gravity toward the geometric center of the body. The knee and ankle joints became increasingly robust to better support increased weight. To support the increased weight on each vertebra in the upright position, the human vertebral column became S-shaped and the lumbar vertebrae became shorter and wider. In the feet the big toe moved into alignment with the other toes to help in forward locomotion. The arms and forearms shortened relative to the legs making it easier to run. The foramen magnum migrated under the skull and more anterior.[15]

The most significant changes occurred in the pelvic region, where the long downward facing iliac blade was shortened and widened as a requirement for keeping the center of gravity stable while walking;[16] bipedal hominids have a shorter but broader, bowl-like pelvis due to this. A drawback is that the birth canal of bipedal apes is smaller than in knuckle-walking apes, though there has been a widening of it in comparison to that of australopithecine and modern humans, permitting the passage of newborns due to the increase in cranial size but this is limited to the upper portion, since further increase can hinder normal bipedal movement.[17]

The shortening of the pelvis and smaller birth canal evolved as a requirement for bipedalism and had significant effects on the process of human birth which is much more difficult in modern humans than in other primates. During human birth, because of the variation in size of the pelvic region, the fetal head must be in a transverse position (compared to the mother) during entry into the birth canal and rotate about 90 degrees upon exit.[18] The smaller birth canal became a limiting factor to brain size increases in early humans and prompted a shorter gestation period leading to the relative immaturity of human offspring, who are unable to walk much before 12 months and have greater neoteny, compared to other primates, who are mobile at a much earlier age.[10] The increased brain growth after birth and the increased dependency of children on mothers had a big effect upon the female reproductive cycle,[19] and the more frequent appearance of alloparenting in humans when compared with other hominids.[20] Delayed human sexual maturity also led to the evolution of menopause with one explanation providing that elderly women could better pass on their genes by taking care of their daughter's offspring, as compared to having more children of their own.[21]

The human species eventually developed a much larger brain than that of other primatestypically 1,330 cm3 in modern humans, nearly three times the size of that of a chimpanzee or gorilla.[22] The pattern of encephalization started with Homo habilis, after a hiatus with Anamensis and Ardipithecus species which had smaller brains as a result of their bipedal locomotion[23] which at approximately 600cm3Homo habilis had a brain slightly larger than that of chimpanzees, and this evolution continued with Homo erectus (8001,100cm3), reaching a maximum in Neanderthals with an average size of (1,2001,900cm3), larger even than modern Homo sapiens. This pattern of brain increase happened through the pattern of human postnatal brain growth which differs from that of other apes (heterochrony). It also allows for extended periods of social learning and language acquisition in juvenile humans which may have begun 2 million years ago. However, the differences between the structure of human brains and those of other apes may be even more significant than differences in size.[24][25][26][27]

The increase in volume over time has affected areas within the brain unequallythe temporal lobes, which contain centers for language processing, have increased disproportionately, and seems to favor a belief that there was evolution after leaving Africa, as has the prefrontal cortex which has been related to complex decision-making and moderating social behavior.[22] Encephalization has been tied to an increasing emphasis on meat in the diet,[28][29][30] or with the development of cooking,[31] and it has been proposed that intelligence increased as a response to an increased necessity for solving social problems as human society became more complex.[32] The human brain was able to expand because of the changes in the morphology of smaller mandibles and mandible muscle attachments to the skull into allowing more room for the brain to grow.[33]

The increase in volume of the neocortex also included a rapid increase in size of the cerebellum. Traditionally the cerebellum has been associated with a paleocerebellum and archicerebellum as well as a neocerebellum. Its function has also traditionally been associated with balance, fine motor control but more recently speech and cognition. The great apes including humans and its antecessors had a more pronounced development of the cerebellum relative to the neocortex than other primates. It has been suggested that because of its function of sensory-motor control and assisting in learning complex muscular action sequences, the cerebellum may have underpinned the evolution of human's technological adaptations including the preadaptation of speech.[34][35][36][37]

The reason for this encephalization is difficult to discern, as the major changes from Homo erectus to Homo heidelbergensis were not associated with major changes in technology. It has been suggested that the changes have been associated with social changes, increased empathic abilities[38][39] and increases in size of social groupings[40][41][42]

The reduced degree of sexual dimorphism is visible primarily in the reduction of the male canine tooth relative to other ape species (except gibbons) and reduced brow ridges and general robustness of males. Another important physiological change related to sexuality in humans was the evolution of hidden estrus. Humans and bonobos are the only apes in which the female is fertile year round and in which no special signals of fertility are produced by the body (such as genital swelling during estrus).

Nonetheless, humans retain a degree of sexual dimorphism in the distribution of body hair and subcutaneous fat, and in the overall size, males being around 15% larger than females. These changes taken together have been interpreted as a result of an increased emphasis on pair bonding as a possible solution to the requirement for increased parental investment due to the prolonged infancy of offspring.

The ulnar opposition the contact between the thumb and the tip of the little finger of the same hand is unique to anatomically modern humans.[43][44] In other primates the thumb is short and unable to touch the little finger.[43] The ulnar opposition facilitates the precision grip and power grip of the human hand, underlying all the skilled manipulations.

A number of other changes have also characterized the evolution of humans, among them an increased importance on vision rather than smell; a smaller gut; loss of body hair; evolution of sweat glands; a change in the shape of the dental arcade from being u-shaped to being parabolic; development of a chin (found in Homo sapiens alone); development of styloid processes; and the development of a descended larynx.

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The word homo, the name of the biological genus to which humans belong, is Latin for "human". It was chosen originally by Carl Linnaeus in his classification system. The word "human" is from the Latin humanus, the adjectival form of homo. The Latin "homo" derives from the Indo-European root *dhghem, or "earth".[45] Linnaeus and other scientists of his time also considered the great apes to be the closest relatives of humans based on morphological and anatomical similarities.

The possibility of linking humans with earlier apes by descent became clear only after 1859 with the publication of Charles Darwin's On the Origin of Species, in which he argued for the idea of the evolution of new species from earlier ones. Darwin's book did not address the question of human evolution, saying only that "Light will be thrown on the origin of man and his history."[46]

The first debates about the nature of human evolution arose between Thomas Henry Huxley and Richard Owen. Huxley argued for human evolution from apes by illustrating many of the similarities and differences between humans and apes, and did so particularly in his 1863 book Evidence as to Man's Place in Nature. However, many of Darwin's early supporters (such as Alfred Russel Wallace and Charles Lyell) did not initially agree that the origin of the mental capacities and the moral sensibilities of humans could be explained by natural selection, though this later changed. Darwin applied the theory of evolution and sexual selection to humans when he published The Descent of Man in 1871.[47]

A major problem at that time was the lack of fossil intermediaries. Neanderthal remains were discovered in a limestone quarry in 1856, three years before the publication of On the Origin of Species, and Neanderthal fossils had been discovered in Gibraltar even earlier, but it was originally claimed that these were human remains of a creature suffering some kind of illness.[48] Despite the 1891 discovery by Eugne Dubois of what is now called Homo erectus at Trinil, Java, it was only in the 1920s when such fossils were discovered in Africa, that intermediate species began to accumulate.[citation needed] In 1925, Raymond Dart described Australopithecus africanus.[49] The type specimen was the Taung Child, an australopithecine infant which was discovered in a cave. The child's remains were a remarkably well-preserved tiny skull and an endocast of the brain.

Although the brain was small (410cm3), its shape was rounded, unlike that of chimpanzees and gorillas, and more like a modern human brain. Also, the specimen showed short canine teeth, and the position of the foramen magnum (the hole in the skull where the spine enters) was evidence of bipedal locomotion. All of these traits convinced Dart that the Taung Child was a bipedal human ancestor, a transitional form between apes and humans.

During the 1960s and 1970s, hundreds of fossils were found in East Africa in the regions of the Olduvai Gorge and Lake Turkana. The driving force of these searches was the Leakey family, with Louis Leakey and his wife Mary Leakey, and later their son Richard and daughter-in-law Meaveall successful and world-renowned fossil hunters and paleoanthropologists. From the fossil beds of Olduvai and Lake Turkana they amassed specimens of the early hominins: the australopithecines and Homo species, and even Homo erectus.

These finds cemented Africa as the cradle of humankind. In the late 1970s and the 1980s, Ethiopia emerged as the new hot spot of paleoanthropology after "Lucy", the most complete fossil member of the species Australopithecus afarensis, was found in 1974 by Donald Johanson near Hadar in the desertic Afar Triangle region of northern Ethiopia. Although the specimen had a small brain, the pelvis and leg bones were almost identical in function to those of modern humans, showing with certainty that these hominins had walked erect.[50] Lucy was classified as a new species, Australopithecus afarensis, which is thought to be more closely related to the genus Homo as a direct ancestor, or as a close relative of an unknown ancestor, than any other known hominid or hominin from this early time range; see terms "hominid" and "hominin".[51] (The specimen was nicknamed "Lucy" after the Beatles' song "Lucy in the Sky with Diamonds", which was played loudly and repeatedly in the camp during the excavations.[52]) The Afar Triangle area would later yield discovery of many more hominin fossils, particularly those uncovered or described by teams headed by Tim D. White in the 1990s, including Ardipithecus ramidus and Ardipithecus kadabba.[53]

In 2013, fossil skeletons of Homo naledi, an extinct species of hominin assigned (provisionally) to the genus Homo, were found in the Rising Star Cave system, a site in South Africa's Cradle of Humankind region in Gauteng province near Johannesburg.[54][55] As of September 2015[update], fossils of at least fifteen individuals, amounting to 1550 specimens, have been excavated from the cave.[55] The species is characterized by a body mass and stature similar to small-bodied human populations, a smaller endocranial volume similar to Australopithecus, and a cranial morphology (skull shape) similar to early Homo species. The skeletal anatomy combines primitive features known from australopithecines with features known from early hominins. The individuals show signs of having been deliberately disposed of within the cave near the time of death. The fossils have not yet been dated.[56]

The genetic revolution in studies of human evolution started when Vincent Sarich and Allan Wilson measured the strength of immunological cross-reactions of blood serum albumin between pairs of creatures, including humans and African apes (chimpanzees and gorillas).[57] The strength of the reaction could be expressed numerically as an immunological distance, which was in turn proportional to the number of amino acid differences between homologous proteins in different species. By constructing a calibration curve of the ID of species' pairs with known divergence times in the fossil record, the data could be used as a molecular clock to estimate the times of divergence of pairs with poorer or unknown fossil records.

In their seminal 1967 paper in Science, Sarich and Wilson estimated the divergence time of humans and apes as four to five million years ago,[57] at a time when standard interpretations of the fossil record gave this divergence as at least 10 to as much as 30 million years. Subsequent fossil discoveries, notably "Lucy", and reinterpretation of older fossil materials, notably Ramapithecus, showed the younger estimates to be correct and validated the albumin method.

Progress in DNA sequencing, specifically mitochondrial DNA (mtDNA) and then Y-chromosome DNA (Y-DNA) advanced the understanding of human origins.[58][9][59] Application of the molecular clock principle revolutionized the study of molecular evolution.

On the basis of a separation from the orangutan between 10 and 20 million years ago, earlier studies of the molecular clock suggested that there were about 76 mutations per generation that were not inherited by human children from their parents; this evidence supported the divergence time between hominins and chimps noted above. However, a 2012 study in Iceland of 78 children and their parents suggests a mutation rate of only 36 mutations per generation; this datum extends the separation between humans and chimps to an earlier period greater than 7 million years ago (Ma). Additional research with 226 offspring of wild chimp populations in 8 locations suggests that chimps reproduce at age 26.5 years, on average; which suggests the human divergence from chimps occurred between 7 and 13 million years ago. And these data suggest that Ardipithecus (4.5 Ma), Orrorin (6 Ma) and Sahelanthropus (7 Ma) all may be on the hominid lineage, and even that the separation may have occurred outside the East African Rift region.

Furthermore, analysis of the two species' genes in 2006 provides evidence that after human ancestors had started to diverge from chimpanzees, interspecies mating between "proto-human" and "proto-chimps" nonetheless occurred regularly enough to change certain genes in the new gene pool:

The research suggests:

In the 1990s, several teams of paleoanthropologists were working throughout Africa looking for evidence of the earliest divergence of the hominin lineage from the great apes. In 1994, Meave Leakey discovered Australopithecus anamensis. The find was overshadowed by Tim D. White's 1995 discovery of Ardipithecus ramidus, which pushed back the fossil record to 4.2 million years ago.

In 2000, Martin Pickford and Brigitte Senut discovered, in the Tugen Hills of Kenya, a 6-million-year-old bipedal hominin which they named Orrorin tugenensis. And in 2001, a team led by Michel Brunet discovered the skull of Sahelanthropus tchadensis which was dated as 7.2 million years ago, and which Brunet argued was a bipedal, and therefore a hominidthat is, a hominin (cf Hominidae; terms "hominids" and hominins).

Different models for the beginning of the present human species.

Anthropologists in the 1980s were divided regarding some details of reproductive barriers and migratory dispersals of the Homo genus. Subsequently, genetics has been used to investigate and resolve these issues. According to the Sahara pump theory evidence suggests that genus Homo have migrated out of Africa at least three and possibly four times (e.g. Homo erectus, Homo heidelbergensis and two or three times for Homo sapiens). Recent evidence suggests these dispersals are closely related to fluctuating periods of climate change.[63]

Recent evidence suggests that humans may have left Africa half a million years earlier than previously thought. A joint Franco-Indian team has found human artefacts in the Siwalk Hills north of New Delhi dating back at least 2.6 million years. This is earlier than the previous earliest finding of genus Homo at Dmanisi, in Georgia, dating to 1.85 million years. Although controversial, tools found at a Chinese cave strengthen the case that humans used tools as far back as 2.48 million years ago. [64] This suggests that the Asian "Chopper" tool tradition, found in Java and northern China may have left Africa before the appearance of the Acheulian hand axe.

Up until the genetic evidence became available there were two dominant models for the dispersal of modern humans. The multiregional hypothesis proposed that Homo genus contained only a single interconnected population as it does today (not separate species), and that its evolution took place worldwide continuously over the last couple million years. This model was proposed in 1988 by Milford H. Wolpoff.[65][66] In contrast the "out of Africa" model proposed that modern H. sapiens speciated in Africa recently (that is, approximately 200,000 years ago) and the subsequent migration through Eurasia resulted in nearly complete replacement of other Homo species. This model has been developed by Chris B. Stringer and Peter Andrews.[67][68]

Sequencing mtDNA and Y-DNA sampled from a wide range of indigenous populations revealed ancestral information relating to both male and female genetic heritage, and strengthened the Out of Africa theory and weakened the views of Multiregional Evolutionism.[69] Aligned in genetic tree differences were interpreted as supportive of a recent single origin.[70] Analyses have shown a greater diversity of DNA patterns throughout Africa, consistent with the idea that Africa is the ancestral home of mitochondrial Eve and Y-chromosomal Adam, and that modern human dispersal out of Africa has only occurred over the last 55,000 years.[71]

"Out of Africa" has thus gained much support from research using female mitochondrial DNA and the male Y chromosome. After analysing genealogy trees constructed using 133 types of mtDNA, researchers concluded that all were descended from a female African progenitor, dubbed Mitochondrial Eve. "Out of Africa" is also supported by the fact that mitochondrial genetic diversity is highest among African populations.[72]

A broad study of African genetic diversity, headed by Sarah Tishkoff, found the San people had the greatest genetic diversity among the 113 distinct populations sampled, making them one of 14 "ancestral population clusters". The research also located a possible origin of modern human migration in south-western Africa, near the coastal border of Namibia and Angola.[73] The fossil evidence was insufficient for archaeologist Richard Leakey to resolve the debate about exactly where in Africa modern humans first appeared.[74] Studies of haplogroups in Y-chromosomal DNA and mitochondrial DNA have largely supported a recent African origin.[75] All the evidence from autosomal DNA also predominantly supports a Recent African origin. However, evidence for archaic admixture in modern humans, both in Africa and later, throughout Eurasia has recently been suggested by a number of studies.[76]

Recent sequencing of Neanderthal[77] and Denisovan[78] genomes shows that some admixture with these populations has occurred. Modern humans outside Africa have 24% Neanderthal alleles in their genome, and some Melanesians have an additional 46% of Denisovan alleles. These new results do not contradict the "out of Africa" model, except in its strictest interpretation, although they make the situation more complex. After recovery from a genetic bottleneck that could possibly be due to the Toba supervolcano catastrophe, a fairly small group left Africa and later briefly interbred on three separate occasions with Neanderthals, probably in the middle-east, on the Eurasian steppe or even in North Africa before their departure. Their still predominantly African descendants spread to populate the world. A fraction in turn interbred with Denisovans, probably in south-east Asia, before populating Melanesia.[79]HLA haplotypes of Neanderthal and Denisova origin have been identified in modern Eurasian and Oceanian populations.[80] The Denisovan EPAS1 gene has also been found in Tibetan populations.[81]

There are still differing theories on whether there was a single exodus from Africa or several. A multiple dispersal model involves the Southern Dispersal theory,[82] which has gained support in recent years from genetic, linguistic and archaeological evidence. In this theory, there was a coastal dispersal of modern humans from the Horn of Africa crossing the Bab el Mandib to Yemen at a lower sea level around 70,000 years ago. This group helped to populate Southeast Asia and Oceania, explaining the discovery of early human sites in these areas much earlier than those in the Levant.[82] This group seems to have been dependent upon marine resources for their survival.

Stephen Oppenheimer has proposed a second wave of humans may have later dispersed through the Persian Gulf oases, and the Zagros mountains into the Middle East. Alternatively it may have come across the Sinai Peninsula into Asia, from shortly after 50,000 yrs BP, resulting in the bulk of the human populations of Eurasia. It has been suggested that this second group possibly possessed a more sophisticated "big game hunting" tool technology and was less dependent on coastal food sources than the original group. Much of the evidence for the first group's expansion would have been destroyed by the rising sea levels at the end of each glacial maximum.[82] The multiple dispersal model is contradicted by studies indicating that the populations of Eurasia and the populations of Southeast Asia and Oceania are all descended from the same mitochondrial DNA L3 lineages, which support a single migration out of Africa that gave rise to all non-African populations.[83]

Stephen Oppenheimer, on the basis of the early date of Badoshan Iranian Aurignacian, suggests that this second dispersal, may have occurred with a pluvial period about 50,000 years before the present, with modern human big-game hunting cultures spreading up the Zagros Mountains, carrying modern human genomes from Oman, throughout the Persian Gulf, northward into Armenia and Anatolia, with a variant travelling south into Israel and to Cyrenicia.[84]

The evidence on which scientific accounts of human evolution are based comes from many fields of natural science. The main source of knowledge about the evolutionary process has traditionally been the fossil record, but since the development of genetics beginning in the 1970s, DNA analysis has come to occupy a place of comparable importance. The studies of ontogeny, phylogeny and especially evolutionary developmental biology of both vertebrates and invertebrates offer considerable insight into the evolution of all life, including how humans evolved. The specific study of the origin and life of humans is anthropology, particularly paleoanthropology which focuses on the study of human prehistory.[85]

The closest living relatives of humans are bonobos and chimpanzees (both genus Pan) and gorillas (genus Gorilla).[86] With the sequencing of both the human and chimpanzee genome, current estimates of the similarity between their DNA sequences range between 95% and 99%.[86][87][88] By using the technique called the molecular clock which estimates the time required for the number of divergent mutations to accumulate between two lineages, the approximate date for the split between lineages can be calculated.

The gibbons (family Hylobatidae) and then orangutans (genus Pongo) were the first groups to split from the line leading to the hominins, including humansfollowed by gorillas, and, ultimately, by the chimpanzees (genus Pan). The splitting date between hominin and chimpanzee lineages is placed by some between 4to8 million years ago, that is, during the Late Miocene.[4][89][90]Speciation, however, appears to have been unusually drawn-out. Initial divergence occurred sometime between 7to13 million years ago, but ongoing hybridization blurred the separation and delayed complete separation during several millions of years. Patterson (2006) dated the final divergence at 5to6 million years ago.[91]

Genetic evidence has also been employed to resolve the question of whether there was any gene flow between early modern humans and Neanderthals, and to enhance our understanding of the early human migration patterns and splitting dates. By comparing the parts of the genome that are not under natural selection and which therefore accumulate mutations at a fairly steady rate, it is possible to reconstruct a genetic tree incorporating the entire human species since the last shared ancestor.

Each time a certain mutation (Single-nucleotide polymorphism) appears in an individual and is passed on to his or her descendants a haplogroup is formed including all of the descendants of the individual who will also carry that mutation. By comparing mitochondrial DNA which is inherited only from the mother, geneticists have concluded that the last female common ancestor whose genetic marker is found in all modern humans, the so-called mitochondrial Eve, must have lived around 200,000 years ago.

Human evolutionary genetics studies how one human genome differs from the other, the evolutionary past that gave rise to it, and its current effects. Differences between genomes have anthropological, medical and forensic implications and applications. Genetic data can provide important insight into human evolution.

There is little fossil evidence for the divergence of the gorilla, chimpanzee and hominin lineages.[92] The earliest fossils that have been proposed as members of the hominin lineage are Sahelanthropus tchadensis dating from 7 million years ago, Orrorin tugenensis dating from 5.7 million years ago, and Ardipithecus kadabba dating to 5.6 million years ago. Each of these have been argued to be a bipedal ancestor of later hominins but, in each case, the claims have been contested. It is also possible that one or more of these species are ancestors of another branch of African apes, or that they represent a shared ancestor between hominins and other apes.

The question then of the relationship between these early fossil species and the hominin lineage is still to be resolved. From these early species, the australopithecines arose around 4 million years ago and diverged into robust (also called Paranthropus) and gracile branches, one of which (possibly A. garhi) probably went on to become ancestors of the genus Homo. The australopithecine species that is best represented in the fossil record is Australopithecus afarensis with more than one hundred fossil individuals represented, found from Northern Ethiopia (such as the famous "Lucy"), to Kenya, and South Africa. Fossils of robust australopithecines such as Au. robustus (or alternatively Paranthropus robustus) and Au./P. boisei are particularly abundant in South Africa at sites such as Kromdraai and Swartkrans, and around Lake Turkana in Kenya.

The earliest member of the genus Homo is Homo habilis which evolved around 2.8 million years ago.[93]Homo habilis is the first species for which we have positive evidence of the use of stone tools. They developed the Oldowan lithic technology, named after the Olduvai Gorge in which the first specimens were found. Some scientists consider Homo rudolfensis, a larger bodied group of fossils with similar morphology to the original H. habilis fossils, to be a separate species while others consider them to be part of H. habilissimply representing intraspecies variation, or perhaps even sexual dimorphism. The brains of these early hominins were about the same size as that of a chimpanzee, and their main adaptation was bipedalism as an adaptation to terrestrial living.

During the next million years, a process of encephalization began and, by the arrival (about 1.9 million years ago) of Homo erectus in the fossil record, cranial capacity had doubled. Homo erectus were the first of the hominins to emigrate from Africa, and, from 1.8to1.3 million years ago, this species spread through Africa, Asia, and Europe. One population of H. erectus, also sometimes classified as a separate species Homo ergaster, remained in Africa and evolved into Homo sapiens. It is believed that these species, H. erectus and H. ergaster, were the first to use fire and complex tools.

The earliest transitional fossils between H. ergaster/erectus and archaic H. sapiens are from Africa, such as Homo rhodesiensis, but seemingly transitional forms were also found at Dmanisi, Georgia. These descendants of African H. erectus spread through Eurasia from ca. 500,000 years ago evolving into H. antecessor, H. heidelbergensis and H. neanderthalensis. The earliest fossils of anatomically modern humans are from the Middle Paleolithic, about 200,000 years ago such as the Omo remains of Ethiopia; later fossils from Es Skhul cave in Israel and Southern Europe begin around 90,000 years ago (0.09 million years ago).

As modern humans spread out from Africa, they encountered other hominins such as Homo neanderthalensis and the so-called Denisovans, who may have evolved from populations of Homo erectus that had left Africa around 2 million years ago. The nature of interaction between early humans and these sister species has been a long-standing source of controversy, the question being whether humans replaced these earlier species or whether they were in fact similar enough to interbreed, in which case these earlier populations may have contributed genetic material to modern humans.[94][95]

This migration out of Africa is estimated to have begun about 70,000 years BP (Before Present) and modern humans subsequently spread globally, replacing earlier hominins either through competition or hybridization. They inhabited Eurasia and Oceania by 40,000 years BP, and the Americas by at least 14,500 years BP.[96]

Evolutionary history of the primates can be traced back 65 million years.[97] One of the oldest known primate-like mammal species, the Plesiadapis, came from North America;[98] another, Archicebus, came from China.[99] Other similar basal primates were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and Eocene.

David R. Begun [100] concluded that early primates flourished in Eurasia and that a lineage leading to the African apes and humans, including to Dryopithecus, migrated south from Europe or Western Asia into Africa. The surviving tropical population of primateswhich is seen most completely in the Upper Eocene and lowermost Oligocene fossil beds of the Faiyum depression southwest of Cairogave rise to all extant primate species, including the lemurs of Madagascar, lorises of Southeast Asia, galagos or "bush babies" of Africa, and to the anthropoids, which are the Platyrrhines or New World monkeys, the Catarrhines or Old World monkeys, and the great apes, including humans and other hominids.

The earliest known catarrhine is Kamoyapithecus from uppermost Oligocene at Eragaleit in the northern Great Rift Valley in Kenya, dated to 24 million years ago.[101] Its ancestry is thought to be species related to Aegyptopithecus, Propliopithecus, and Parapithecus from the Faiyum, at around 35 million years ago.[102] In 2010, Saadanius was described as a close relative of the last common ancestor of the crown catarrhines, and tentatively dated to 2928 million years ago, helping to fill an 11-million-year gap in the fossil record.[103]

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