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Although the narratives of human evolution are often contentious, several discoveries since show that human evolution should not be seen as a simple linear or branched progression, but a mix of related species.

Evolutionary history of the primates can be traced back 65 million years. Begun [] 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 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 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.

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 at 20 million years ago include fragments attributed to Victoriapithecus , the earliest Old World monkey. 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 Middle Miocene 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 the line of 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.

Species close to the last common ancestor of gorillas, chimpanzees and humans may be represented by Nakalipithecus fossils found in Kenya and Ouranopithecus found in Greece.

Molecular evidence suggests that between 8 and 4 million years ago, first the gorillas, and then the chimpanzees genus Pan split off from the line leading to the humans.

Human DNA is approximately The fossil record, however, of gorillas and chimpanzees is limited; both poor preservation—rain forest soils tend to be acidic and dissolve bone—and sampling bias probably contribute to this problem.

Other hominins probably adapted to the drier environments outside the equatorial belt; and there they encountered antelope, hyenas, dogs, pigs, elephants, horses, and others.

The equatorial belt contracted after about 8 million years ago, and there is very little fossil evidence for the split—thought to have occurred around that time—of the hominin lineage from the lineages of gorillas and chimpanzees.

The earliest fossils argued by some to belong to the human lineage are Sahelanthropus tchadensis 7 Ma and Orrorin tugenensis 6 Ma , followed by Ardipithecus 5.

It has been argued in a study of the life history of Ar. It was also argued that the species provides support for the notion that very early hominins, akin to bonobos Pan paniscus the less aggressive species of the genus Pan , may have evolved via the process of self-domestication.

Consequently, arguing against the so-called "chimpanzee referential model" [] the authors suggest it is no longer tenable to use common chimpanzee Pan troglodytes social and mating behaviors in models of early hominin social evolution.

When commenting on the absence of aggressive canine morphology in Ar. However, the fact that Ar. In fact the trend towards increased maternal care, female mate selection and self-domestication may have been stronger and more refined in Ar.

The authors argue that many of the basic human adaptations evolved in the ancient forest and woodland ecosystems of late Miocene and early Pliocene Africa.

Consequently, they argue that humans may not represent evolution from a chimpanzee-like ancestor as has traditionally been supposed. This suggests many modern human adaptations represent phylogenetically deep traits and that the behavior and morphology of chimpanzees may have evolved subsequent to the split with the common ancestor they share with humans.

The genus Australopithecus evolved in eastern Africa around 4 million years ago before spreading throughout the continent and eventually becoming extinct 2 million years ago.

During this time period various forms of australopiths existed, including Australopithecus anamensis , Au. There is still some debate among academics whether certain African hominid species of this time, such as Au.

However, if these species do indeed constitute their own genus, then they may be given their own name, the Paranthropus.

A new proposed species Australopithecus deyiremeda is claimed to have been discovered living at the same time period of Au.

There is debate if Au. The earliest documented representative of the genus Homo is Homo habilis , which evolved around 2. The brains of these early hominins were about the same size as that of a chimpanzee , although it has been suggested that this was the time in which the human SRGAP2 gene doubled, producing a more rapid wiring of the frontal cortex.

It is believed that Homo erectus and Homo ergaster were the first to use fire and complex tools, and were the first of the hominin line to leave Africa, spreading throughout Africa, Asia, and Europe between 1.

According to the recent African origin of modern humans theory, modern humans evolved in Africa possibly from Homo heidelbergensis , Homo rhodesiensis or Homo antecessor and migrated out of the continent some 50, to , years ago, gradually replacing local populations of Homo erectus , Denisova hominins , Homo floresiensis and Homo neanderthalensis.

Homo sapiens is the only extant species of its genus, Homo. While some extinct Homo species might have been ancestors of Homo sapiens , many, perhaps most, were likely "cousins", having speciated away from the ancestral hominin line.

Based on archaeological and paleontological evidence, it has been possible to infer, to some extent, the ancient dietary practices [34] of various Homo species and to study the role of diet in physical and behavioral evolution within Homo.

Some anthropologists and archaeologists subscribe to the Toba catastrophe theory , which posits that the supereruption of Lake Toba on Sumatran island in Indonesia some 70, years ago caused global consequences, [] killing the majority of humans and creating a population bottleneck that affected the genetic inheritance of all humans today.

Homo habilis lived from about 2. Homo habilis had smaller molars and larger brains than the australopithecines, and made tools from stone and perhaps animal bones.

Some scientists have proposed moving this species out of Homo and into Australopithecus due to the morphology of its skeleton being more adapted to living on trees rather than to moving on two legs like Homo sapiens.

In May , a new species, Homo gautengensis , was discovered in South Africa. These are proposed species names for fossils from about 1. The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in on the Indonesian island of Java.

He originally named the material Anthropopithecus erectus —, considered at this point as a chimpanzee-like fossil primate and Pithecanthropus erectus —, changing his mind as of based on its morphology, which he considered to be intermediate between that of humans and apes.

Weidenreich concluded in that because of their anatomical similarity with modern humans it was necessary to gather all these specimens of Java and China in a single species of the genus Homo , the species Homo erectus.

The early phase of Homo erectus , from 1. In Africa in the Early Pleistocene, 1. This species also may have used fire to cook meat.

Richard Wrangham suggests that the fact that Homo seems to have been ground dwelling, with reduced intestinal length, smaller dentition, "and swelled our brains to their current, horrendously fuel-inefficient size", [] suggest that control of fire and releasing increased nutritional value through cooking was the key adaptation that separated Homo from tree-sleeping Australopithecines.

Many paleoanthropologists now use the term Homo ergaster for the non-Asian forms of this group, and reserve Homo erectus only for those fossils that are found in Asia and meet certain skeletal and dental requirements which differ slightly from H.

These are proposed as species that may be intermediate between H. Also proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.

Homo neanderthalensis , alternatively designated as Homo sapiens neanderthalensis , [] lived in Europe and Asia from , [] to about 28, years ago.

Many of these relate to the superior adaptation to cold environments possessed by the Neanderthal populations. Their surface to volume ratio is an extreme version of that found amongst Inuit populations, indicating that they were less inclined to lose body heat than were AMH.

From brain Endocasts, Neanderthals also had significantly larger brains. This would seem to indicate that the intellectual superiority of AMH populations may be questionable.

Dunbar, however, have shown important differences in Brain architecture. For example, in both the orbital chamber size and in the size of the occipital lobe , the larger size suggests that the Neanderthal had a better visual acuity than modern humans.

This would give a superior vision in the inferior light conditions found in Glacial Europe. It also seems that the higher body mass of Neanderthals had a correspondingly larger brain mass required for body care and control.

The Neanderthal populations seem to have been physically superior to AMH populations. These differences may have been sufficient to give Neanderthal populations an environmental superiority to AMH populations from 75, to 45, years BP.

With these differences, Neanderthal brains show a smaller area was available for social functioning. Plotting group size possible from endocrainial volume, suggests that AMH populations minus occipital lobe size , had a Dunbars number of possible relationships.

Neanderthal populations seem to have been limited to about individuals. This would show up in a larger number of possible mates for AMH humans, with increased risks of inbreeding amongst Neanderthal populations.

It also suggests that humans had larger trade catchment areas than Neanderthals confirmed in the distribution of stone tools.

With larger populations, social and technological innovations were easier to fix in human populations, which may have all contributed to the fact that modern Homo sapiens replaced the Neanderthal populations by 28, BP.

Earlier evidence from sequencing mitochondrial DNA suggested that no significant gene flow occurred between H. Though this interbred Romanian population seems not to have been ancestral to modern humans, the finding indicates that interbreeding happened repeatedly.

In , archaeologists working at the site of Denisova Cave in the Altai Mountains of Siberia uncovered a small bone fragment from the fifth finger of a juvenile member of Denisovans.

While the divergence point of the mtDNA was unexpectedly deep in time, [] the full genomic sequence suggested the Denisovans belonged to the same lineage as Neanderthals, with the two diverging shortly after their line split from the lineage that gave rise to modern humans.

The existence of this distant branch creates a much more complex picture of humankind during the Late Pleistocene than previously thought.

Alleles thought to have originated in Neanderthals and Denisovans have been identified at several genetic loci in the genomes of modern humans outside of Africa.

HLA haplotypes from Denisovans and Neanderthal represent more than half the HLA alleles of modern Eurasians, [91] indicating strong positive selection for these introgressed alleles.

Corinne Simoneti at Vanderbilt University, in Nashville and her team have found from medical records of 28, people of European descent that the presence of Neanderthal DNA segments may be associated with a likelihood to suffer depression more frequently.

The flow of genes from Neanderthal populations to modern human was not all one way. Sergi Castellano of the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, has in reported that while Denisovan and Neanderthal genomes are more related to each other than they are to us, Siberian Neanderthal genomes show similarity to the modern human gene pool, more so than to European Neanderthal populations.

The evidence suggests that the Neanderthal populations interbred with modern humans possibly , years ago, probably somewhere in the Near East.

Studies of a Neanderthal child at Gibraltar show from brain development and teeth eruption that Neanderthal children may have matured more rapidly than is the case for Homo sapiens.

In other words, H. The main find was a skeleton believed to be a woman of about 30 years of age. Found in , it has been dated to approximately 18, years old.

However, there is an ongoing debate over whether H. This, coupled with pathological dwarfism, could have resulted in a significantly diminutive human.

The other major attack on H. The hypothesis of pathological dwarfism, however, fails to explain additional anatomical features that are unlike those of modern humans diseased or not but much like those of ancient members of our genus.

Aside from cranial features, these features include the form of bones in the wrist, forearm, shoulder, knees, and feet. Additionally, this hypothesis fails to explain the find of multiple examples of individuals with these same characteristics, indicating they were common to a large population, and not limited to one individual.

The direct evidence suggests there was a migration of H. A subsequent migration both within and out of Africa eventually replaced the earlier dispersed H.

This migration and origin theory is usually referred to as the "recent single-origin hypothesis" or "out of Africa" theory.

The Toba catastrophe theory , which postulates a population bottleneck for H. The use of tools has been interpreted as a sign of intelligence, and it has been theorized that tool use may have stimulated certain aspects of human evolution, especially the continued expansion of the human brain.

Researchers have suggested that early hominins were thus under evolutionary pressure to increase their capacity to create and use tools.

Precisely when early humans started to use tools is difficult to determine, because the more primitive these tools are for example, sharp-edged stones the more difficult it is to decide whether they are natural objects or human artifacts.

Many species make and use tools , but it is the human genus that dominates the areas of making and using more complex tools. The oldest known tools are flakes from West Turkana, Kenya, which date to 3.

These tools date to about 2. It is a possibility but does not yet represent solid evidence. It allows humans the dexterity and strength to make and use complex tools.

This unique anatomical feature separates humans from apes and other nonhuman primates, and is not seen in human fossils older than 1.

Bernard Wood noted that Paranthropus co-existed with the early Homo species in the area of the "Oldowan Industrial Complex" over roughly the same span of time.

Although there is no direct evidence which identifies Paranthropus as the tool makers, their anatomy lends to indirect evidence of their capabilities in this area.

Most paleoanthropologists agree that the early Homo species were indeed responsible for most of the Oldowan tools found.

They argue that when most of the Oldowan tools were found in association with human fossils, Homo was always present, but Paranthropus was not.

In , Randall Susman used the anatomy of opposable thumbs as the basis for his argument that both the Homo and Paranthropus species were toolmakers.

He compared bones and muscles of human and chimpanzee thumbs, finding that humans have 3 muscles which are lacking in chimpanzees.

Humans also have thicker metacarpals with broader heads, allowing more precise grasping than the chimpanzee hand can perform. Susman posited that modern anatomy of the human opposable thumb is an evolutionary response to the requirements associated with making and handling tools and that both species were indeed toolmakers.

Stone tools are first attested around 2. Archaeologists working in the Great Rift Valley in Kenya have discovered the oldest known stone tools in the world.

Dated to around 3. The period from ,—, years ago is also known as the Acheulean , when H. After , BP the more refined so-called Levallois technique was developed, a series of consecutive strikes, by which scrapers, slicers "racloirs" , needles, and flattened needles were made.

In this period they also started to make tools out of bone. Until about 50,—40, years ago, the use of stone tools seems to have progressed stepwise.

Currently paleoanthropologists are debating whether these Homo species possessed some or many of the cultural and behavioral traits associated with modern humans such as language, complex symbolic thinking, technological creativity etc.

It seems that they were culturally conservative maintaining simple technologies and foraging patterns over very long periods.

Around 50, BP , modern human culture started to evolve more rapidly. The transition to behavioral modernity has been characterized by most as a Eurasian "Great Leap Forward", [] or as the "Upper Palaeolithic Revolution", [] due to the sudden appearance of distinctive signs of modern behavior and big game hunting [96] in the archaeological record.

Some other scholars consider the transition to have been more gradual, noting that some features had already appeared among archaic African Homo sapiens since , years ago.

Modern humans started burying their dead, using animal hides to make clothing, hunting with more sophisticated techniques such as using trapping pits or driving animals off cliffs , and engaging in cave painting.

Among concrete examples of modern human behavior , anthropologists include specialization of tools, use of jewellery and images such as cave drawings , organization of living space, rituals for example, burials with grave gifts , specialized hunting techniques, exploration of less hospitable geographical areas, and barter trade networks.

Debate continues as to whether a "revolution" led to modern humans "the big bang of human consciousness" , or whether the evolution was more "gradual".

Evolution has continued in anatomically modern human populations, which are affected by both natural selection and genetic drift.

Although selection pressure on some traits, such as resistance to smallpox, has decreased in modern human life, humans are still undergoing natural selection for many other traits.

Some of these are due to specific environmental pressures, while others are related to lifestyle changes since the development of agriculture 10, years ago , urban civilization 5, , and industrialization years ago.

It has been argued that human evolution has accelerated since the development of agriculture 10, years ago and civilization some 5, years ago, resulting, it is claimed, in substantial genetic differences between different current human populations.

Particularly conspicuous is variation in superficial characteristics, such as Afro-textured hair , or the recent evolution of light skin and blond hair in some populations, which are attributed to differences in climate.

Particularly strong selective pressures have resulted in high-altitude adaptation in humans , with different ones in different isolated populations.

Studies of the genetic basis show that some developed very recently, with Tibetans evolving over 3, years to have high proportions of an allele of EPAS1 that is adaptive to high altitudes.

Other evolution is related to endemic diseases: For example, the population at risk of the severe debilitating disease kuru has significant over-representation of an immune variant of the prion protein gene GV versus non-immune alleles.

The frequency of this genetic variant is due to the survival of immune persons. Recent human evolution related to agriculture includes genetic resistance to infectious disease that has appeared in human populations by crossing the species barrier from domesticated animals, [] as well as changes in metabolism due to changes in diet, such as lactase persistence.

In contemporary times, since industrialization, some trends have been observed: This list is in chronological order across the table by genus.

Please see articles for more information. From Wikipedia, the free encyclopedia. Evolutionary process leading up to the appearance of anatomically modern humans.

Life timeline and Nature timeline. A global mapping model of human migration, based from divergence of the mitochondrial DNA which indicates the matrilineage.

A "trellis" as Milford H. Wolpoff called it that emphasizes back-and-forth gene flow among geographic regions.

Different models for the beginning of the present human species. Early human migrations , Recent African origin of modern humans , Multiregional origin of modern humans , and Early hominids in Southeast Asia.

Human evolutionary genetics and Human genetic variation. Interbreeding between archaic and modern humans.

For evolutionary history before primates, see Evolution of mammals , Evolutionary history of life , and Timeline of human evolution. It has been suggested that this section be split out into another article titled Homo.

Control of fire by early humans. Scientific American , May New Scientist , Volume , Issue , 30 July , pp. Brazilian Archives of Biology and Technology.

The Impact of Altitude and Activity Scheduling". Journal of Human Evolution. Evolutionary biologists consider menopause". Annual Review of Anthropology.

Josh; Robertson, Marcia L. Annual Review of Nutrition. University of California Press. The New York Times. Retrieved August 14, Computational Evidence for the Cognitive Costs of Sociality".

Proceedings of the Royal Society B. Archives of Psychiatry and Psychotherapy. Survival of the Nicest. Proceedings of the Royal Society of London B: Proceedings of the National Academy of Sciences.

American Journal of Physical Anthropology: Hand Preference and Hand Ability: Evidence from studies in Haptic Cognition. Retrieved 2 July Nature Full image view 1st ed.

The Man-Ape of South Africa". Explicit use of et al. Retrieved 10 September The lower limit of ka is due to optically stimulated luminescence dating of sediments with U-Th and palaeomagnetic analyses of flowstones; the upper limit of ka is due to U-series and electron spin resonance US-ESR dating of two H.

Full list of authors. Aiello, Leslie ; Dean, Christopher An Introduction to Human Evolutionary Anatomy. American Heritage Dictionaries editors More Word Histories and Mysteries: From Aardvark to Zombie.

The Anthropology of Medicine: From Culture to Method 3rd ed. Boyd, Robert ; Silk, Joan B. How Humans Evolved 3rd ed. In Siegel, Bernard J. Biennial Review of Anthropology Biennial Review of Anthropology.

Bryson, Bill [Originally published ]. A Short History of Nearly Everything. Hominid Adaptations and Extinctions. Cochran, Gregory ; Harpending, Henry The 10, Year Explosion: How Civilization Accelerated Human Evolution.

Darwin, Charles [Originally published ; London: A Pilgrimage to the Dawn of Evolution. DeSalle, Rob; Tattersall, Ian Guns, Germs, and Steel: The Fates of Human Societies.

Freeman, Scott; Herron, Jon C. Evolutionary Analysis 4th ed. Upper Saddle River, NJ: Johanson, Donald ; Edey, Maitland Lucy, the Beginnings of Humankind.

In Glick, Thomas F. The Comparative Reception of Darwinism. University of Chicago Press. American Journal of Physical Anthropology.

University of Tokyo Press. The Origin of Humankind. The Human Genome Diversity Project: An Ethnography of Scientific Practice.

Cambridge Studies in Society and the Life Sciences. In Smelser, Neil J. In Ruse, Michael ; Travis, Joseph. COM in 30 languages.

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All of these traits convinced Dart that the Taung Child was a bipedal human ancestor, a transitional form between apes and humans.

During the s and s, 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 Meave —all successful and world-renowned fossil hunters and paleoanthropologists.

From the fossil beds of Olduvai and Lake Turkana they amassed specimens of the early hominins: These finds cemented Africa as the cradle of humankind.

In the late s and the s, Ethiopia emerged as the new hot spot of paleoanthropology after "Lucy" , the most complete fossil member of the species Australopithecus afarensis , was found in 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.

White in the s, including Ardipithecus ramidus and Ardipithecus kadabba. 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 were dated close to , years ago, [66] and thus are not a direct ancestor but a contemporary with the first appearance of larger-brained anatomically modern humans.

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.

In their seminal paper in Science , Sarich and Wilson estimated the divergence time of humans and apes as four to five million years ago, [68] 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.

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 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 offspring of wild chimp populations in 8 locations suggests that chimps reproduce at age And these data suggest that Ardipithecus 4.

In the s, several teams of paleoanthropologists were working throughout Africa looking for evidence of the earliest divergence of the hominin lineage from the great apes.

In , Meave Leakey discovered Australopithecus anamensis. The find was overshadowed by Tim D. In , 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 , a team led by Michel Brunet discovered the skull of Sahelanthropus tchadensis which was dated as 7. Anthropologists in the s were divided regarding some details of reproductive barriers and migratory dispersals of the genus Homo.

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.

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.

Recent evidence suggests that humans may have left Africa half a million years earlier than previously thought. This is earlier than the previous earliest finding of genus Homo at Dmanisi , in Georgia , dating to 1.

Although controversial, tools found at a Chinese cave strengthen the case that humans used tools as far back as 2. Up until the genetic evidence became available there were two dominant models for the dispersal of modern humans.

The multiregional hypothesis proposed that the genus Homo 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 of million years.

This model was proposed in by Milford H. This model has been developed by Chris B. Stringer and Peter Andrews. 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.

After analysing genealogy trees constructed using types of mtDNA, researchers concluded that all were descended from a female African progenitor, dubbed Mitochondrial Eve.

A broad study of African genetic diversity, headed by Sarah Tishkoff , found the San people had the greatest genetic diversity among the 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. However, evidence for archaic admixture in modern humans , both in Africa and later, throughout Eurasia has recently been suggested by a number of studies.

Recent sequencing of Neanderthal [88] and Denisovan [89] genomes shows that some admixture with these populations has occurred. 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 some researchers speculate might be linked to the Toba supervolcano catastrophe , a fairly small group left Africa and interbred 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.

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, [94] 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, 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.

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, 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.

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, 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.

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 s, 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.

The closest living relatives of humans are bonobos and chimpanzees both genus Pan and gorillas genus Gorilla. The gibbons family Hylobatidae and then orangutans genus Pongo were the first groups to split from the line leading to the hominins, including humans—followed by gorillas, and, ultimately, by the chimpanzees genus Pan.

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 , 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. 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.

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. The earliest member of the genus Homo is Homo habilis which evolved around 2.

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.

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. Homo erectus were the first of the hominins to emigrate from Africa, and, from 1.

One population of H. It is believed that these species, H. The earliest transitional fossils between H. These descendants of African H. The earliest fossils of anatomically modern humans are from the Middle Paleolithic, about , years ago such as the Omo remains of Ethiopia; later fossils from Es Skhul cave in Israel and Southern Europe begin around 90, years ago 0.

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.

This migration out of Africa is estimated to have begun about 70, years BP and modern humans subsequently spread globally, replacing earlier hominins either through competition or hybridization.

The hypothesis of interbreeding, also known as hybridization, admixture or hybrid-origin theory, has been discussed ever since the discovery of Neanderthal remains in the 19th century.

In the 21st century with the advent of molecular biology techniques and computerization, whole-genome sequencing of Neanderthal and human genome were performed, confirming recent admixture between different human species.

It has been demonstrated that interbreeding happened in several independent events that included Neanderthals and Denisovans, as well as several unidentified hominins.

For example, comparative studies in the mids found several traits related to neurological, immunological, [] developmental, and metabolic phenotypes, that were developed by archaic humans to European and Asian environments and inherited to modern humans through admixture with local hominins.

Although the narratives of human evolution are often contentious, several discoveries since show that human evolution should not be seen as a simple linear or branched progression, but a mix of related species.

Evolutionary history of the primates can be traced back 65 million years. Begun [] 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 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 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.

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 at 20 million years ago include fragments attributed to Victoriapithecus , the earliest Old World monkey. 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 Middle Miocene 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 the line of 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.

Species close to the last common ancestor of gorillas, chimpanzees and humans may be represented by Nakalipithecus fossils found in Kenya and Ouranopithecus found in Greece.

Molecular evidence suggests that between 8 and 4 million years ago, first the gorillas, and then the chimpanzees genus Pan split off from the line leading to the humans.

Human DNA is approximately The fossil record, however, of gorillas and chimpanzees is limited; both poor preservation—rain forest soils tend to be acidic and dissolve bone—and sampling bias probably contribute to this problem.

Other hominins probably adapted to the drier environments outside the equatorial belt; and there they encountered antelope, hyenas, dogs, pigs, elephants, horses, and others.

The equatorial belt contracted after about 8 million years ago, and there is very little fossil evidence for the split—thought to have occurred around that time—of the hominin lineage from the lineages of gorillas and chimpanzees.

The earliest fossils argued by some to belong to the human lineage are Sahelanthropus tchadensis 7 Ma and Orrorin tugenensis 6 Ma , followed by Ardipithecus 5.

It has been argued in a study of the life history of Ar. It was also argued that the species provides support for the notion that very early hominins, akin to bonobos Pan paniscus the less aggressive species of the genus Pan , may have evolved via the process of self-domestication.

Consequently, arguing against the so-called "chimpanzee referential model" [] the authors suggest it is no longer tenable to use common chimpanzee Pan troglodytes social and mating behaviors in models of early hominin social evolution.

When commenting on the absence of aggressive canine morphology in Ar. However, the fact that Ar. In fact the trend towards increased maternal care, female mate selection and self-domestication may have been stronger and more refined in Ar.

The authors argue that many of the basic human adaptations evolved in the ancient forest and woodland ecosystems of late Miocene and early Pliocene Africa.

Consequently, they argue that humans may not represent evolution from a chimpanzee-like ancestor as has traditionally been supposed.

This suggests many modern human adaptations represent phylogenetically deep traits and that the behavior and morphology of chimpanzees may have evolved subsequent to the split with the common ancestor they share with humans.

The genus Australopithecus evolved in eastern Africa around 4 million years ago before spreading throughout the continent and eventually becoming extinct 2 million years ago.

During this time period various forms of australopiths existed, including Australopithecus anamensis , Au. There is still some debate among academics whether certain African hominid species of this time, such as Au.

However, if these species do indeed constitute their own genus, then they may be given their own name, the Paranthropus. A new proposed species Australopithecus deyiremeda is claimed to have been discovered living at the same time period of Au.

There is debate if Au. The earliest documented representative of the genus Homo is Homo habilis , which evolved around 2. The brains of these early hominins were about the same size as that of a chimpanzee , although it has been suggested that this was the time in which the human SRGAP2 gene doubled, producing a more rapid wiring of the frontal cortex.

It is believed that Homo erectus and Homo ergaster were the first to use fire and complex tools, and were the first of the hominin line to leave Africa, spreading throughout Africa, Asia, and Europe between 1.

According to the recent African origin of modern humans theory, modern humans evolved in Africa possibly from Homo heidelbergensis , Homo rhodesiensis or Homo antecessor and migrated out of the continent some 50, to , years ago, gradually replacing local populations of Homo erectus , Denisova hominins , Homo floresiensis and Homo neanderthalensis.

Homo sapiens is the only extant species of its genus, Homo. While some extinct Homo species might have been ancestors of Homo sapiens , many, perhaps most, were likely "cousins", having speciated away from the ancestral hominin line.

Based on archaeological and paleontological evidence, it has been possible to infer, to some extent, the ancient dietary practices [34] of various Homo species and to study the role of diet in physical and behavioral evolution within Homo.

Some anthropologists and archaeologists subscribe to the Toba catastrophe theory , which posits that the supereruption of Lake Toba on Sumatran island in Indonesia some 70, years ago caused global consequences, [] killing the majority of humans and creating a population bottleneck that affected the genetic inheritance of all humans today.

Homo habilis lived from about 2. Homo habilis had smaller molars and larger brains than the australopithecines, and made tools from stone and perhaps animal bones.

Some scientists have proposed moving this species out of Homo and into Australopithecus due to the morphology of its skeleton being more adapted to living on trees rather than to moving on two legs like Homo sapiens.

In May , a new species, Homo gautengensis , was discovered in South Africa. These are proposed species names for fossils from about 1. The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in on the Indonesian island of Java.

He originally named the material Anthropopithecus erectus —, considered at this point as a chimpanzee-like fossil primate and Pithecanthropus erectus —, changing his mind as of based on its morphology, which he considered to be intermediate between that of humans and apes.

Weidenreich concluded in that because of their anatomical similarity with modern humans it was necessary to gather all these specimens of Java and China in a single species of the genus Homo , the species Homo erectus.

The early phase of Homo erectus , from 1. In Africa in the Early Pleistocene, 1. This species also may have used fire to cook meat. Richard Wrangham suggests that the fact that Homo seems to have been ground dwelling, with reduced intestinal length, smaller dentition, "and swelled our brains to their current, horrendously fuel-inefficient size", [] suggest that control of fire and releasing increased nutritional value through cooking was the key adaptation that separated Homo from tree-sleeping Australopithecines.

Many paleoanthropologists now use the term Homo ergaster for the non-Asian forms of this group, and reserve Homo erectus only for those fossils that are found in Asia and meet certain skeletal and dental requirements which differ slightly from H.

These are proposed as species that may be intermediate between H. Also proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.

Homo neanderthalensis , alternatively designated as Homo sapiens neanderthalensis , [] lived in Europe and Asia from , [] to about 28, years ago.

Many of these relate to the superior adaptation to cold environments possessed by the Neanderthal populations. Their surface to volume ratio is an extreme version of that found amongst Inuit populations, indicating that they were less inclined to lose body heat than were AMH.

From brain Endocasts, Neanderthals also had significantly larger brains. This would seem to indicate that the intellectual superiority of AMH populations may be questionable.

Dunbar, however, have shown important differences in Brain architecture. For example, in both the orbital chamber size and in the size of the occipital lobe , the larger size suggests that the Neanderthal had a better visual acuity than modern humans.

This would give a superior vision in the inferior light conditions found in Glacial Europe. It also seems that the higher body mass of Neanderthals had a correspondingly larger brain mass required for body care and control.

The Neanderthal populations seem to have been physically superior to AMH populations. These differences may have been sufficient to give Neanderthal populations an environmental superiority to AMH populations from 75, to 45, years BP.

With these differences, Neanderthal brains show a smaller area was available for social functioning. Plotting group size possible from endocrainial volume, suggests that AMH populations minus occipital lobe size , had a Dunbars number of possible relationships.

Neanderthal populations seem to have been limited to about individuals. This would show up in a larger number of possible mates for AMH humans, with increased risks of inbreeding amongst Neanderthal populations.

It also suggests that humans had larger trade catchment areas than Neanderthals confirmed in the distribution of stone tools. With larger populations, social and technological innovations were easier to fix in human populations, which may have all contributed to the fact that modern Homo sapiens replaced the Neanderthal populations by 28, BP.

Earlier evidence from sequencing mitochondrial DNA suggested that no significant gene flow occurred between H. Though this interbred Romanian population seems not to have been ancestral to modern humans, the finding indicates that interbreeding happened repeatedly.

In , archaeologists working at the site of Denisova Cave in the Altai Mountains of Siberia uncovered a small bone fragment from the fifth finger of a juvenile member of Denisovans.

While the divergence point of the mtDNA was unexpectedly deep in time, [] the full genomic sequence suggested the Denisovans belonged to the same lineage as Neanderthals, with the two diverging shortly after their line split from the lineage that gave rise to modern humans.

The existence of this distant branch creates a much more complex picture of humankind during the Late Pleistocene than previously thought.

Alleles thought to have originated in Neanderthals and Denisovans have been identified at several genetic loci in the genomes of modern humans outside of Africa.

HLA haplotypes from Denisovans and Neanderthal represent more than half the HLA alleles of modern Eurasians, [91] indicating strong positive selection for these introgressed alleles.

Corinne Simoneti at Vanderbilt University, in Nashville and her team have found from medical records of 28, people of European descent that the presence of Neanderthal DNA segments may be associated with a likelihood to suffer depression more frequently.

The flow of genes from Neanderthal populations to modern human was not all one way. Sergi Castellano of the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, has in reported that while Denisovan and Neanderthal genomes are more related to each other than they are to us, Siberian Neanderthal genomes show similarity to the modern human gene pool, more so than to European Neanderthal populations.

The evidence suggests that the Neanderthal populations interbred with modern humans possibly , years ago, probably somewhere in the Near East.

Studies of a Neanderthal child at Gibraltar show from brain development and teeth eruption that Neanderthal children may have matured more rapidly than is the case for Homo sapiens.

In other words, H. The main find was a skeleton believed to be a woman of about 30 years of age. Found in , it has been dated to approximately 18, years old.

However, there is an ongoing debate over whether H. This, coupled with pathological dwarfism, could have resulted in a significantly diminutive human.

The other major attack on H. The hypothesis of pathological dwarfism, however, fails to explain additional anatomical features that are unlike those of modern humans diseased or not but much like those of ancient members of our genus.

Aside from cranial features, these features include the form of bones in the wrist, forearm, shoulder, knees, and feet.

Additionally, this hypothesis fails to explain the find of multiple examples of individuals with these same characteristics, indicating they were common to a large population, and not limited to one individual.

The direct evidence suggests there was a migration of H. A subsequent migration both within and out of Africa eventually replaced the earlier dispersed H.

This migration and origin theory is usually referred to as the "recent single-origin hypothesis" or "out of Africa" theory.

The Toba catastrophe theory , which postulates a population bottleneck for H. The use of tools has been interpreted as a sign of intelligence, and it has been theorized that tool use may have stimulated certain aspects of human evolution, especially the continued expansion of the human brain.

Researchers have suggested that early hominins were thus under evolutionary pressure to increase their capacity to create and use tools.

Precisely when early humans started to use tools is difficult to determine, because the more primitive these tools are for example, sharp-edged stones the more difficult it is to decide whether they are natural objects or human artifacts.

Many species make and use tools , but it is the human genus that dominates the areas of making and using more complex tools. The oldest known tools are flakes from West Turkana, Kenya, which date to 3.

These tools date to about 2. It is a possibility but does not yet represent solid evidence. It allows humans the dexterity and strength to make and use complex tools.

This unique anatomical feature separates humans from apes and other nonhuman primates, and is not seen in human fossils older than 1.

Bernard Wood noted that Paranthropus co-existed with the early Homo species in the area of the "Oldowan Industrial Complex" over roughly the same span of time.

Although there is no direct evidence which identifies Paranthropus as the tool makers, their anatomy lends to indirect evidence of their capabilities in this area.

Most paleoanthropologists agree that the early Homo species were indeed responsible for most of the Oldowan tools found. They argue that when most of the Oldowan tools were found in association with human fossils, Homo was always present, but Paranthropus was not.

In , Randall Susman used the anatomy of opposable thumbs as the basis for his argument that both the Homo and Paranthropus species were toolmakers.

He compared bones and muscles of human and chimpanzee thumbs, finding that humans have 3 muscles which are lacking in chimpanzees.

Humans also have thicker metacarpals with broader heads, allowing more precise grasping than the chimpanzee hand can perform. Susman posited that modern anatomy of the human opposable thumb is an evolutionary response to the requirements associated with making and handling tools and that both species were indeed toolmakers.

Stone tools are first attested around 2. Archaeologists working in the Great Rift Valley in Kenya have discovered the oldest known stone tools in the world.

Dated to around 3. The period from ,—, years ago is also known as the Acheulean , when H. After , BP the more refined so-called Levallois technique was developed, a series of consecutive strikes, by which scrapers, slicers "racloirs" , needles, and flattened needles were made.

In this period they also started to make tools out of bone. Until about 50,—40, years ago, the use of stone tools seems to have progressed stepwise.

Currently paleoanthropologists are debating whether these Homo species possessed some or many of the cultural and behavioral traits associated with modern humans such as language, complex symbolic thinking, technological creativity etc.

It seems that they were culturally conservative maintaining simple technologies and foraging patterns over very long periods.

Around 50, BP , modern human culture started to evolve more rapidly. The transition to behavioral modernity has been characterized by most as a Eurasian "Great Leap Forward", [] or as the "Upper Palaeolithic Revolution", [] due to the sudden appearance of distinctive signs of modern behavior and big game hunting [96] in the archaeological record.

Some other scholars consider the transition to have been more gradual, noting that some features had already appeared among archaic African Homo sapiens since , years ago.

Modern humans started burying their dead, using animal hides to make clothing, hunting with more sophisticated techniques such as using trapping pits or driving animals off cliffs , and engaging in cave painting.

Among concrete examples of modern human behavior , anthropologists include specialization of tools, use of jewellery and images such as cave drawings , organization of living space, rituals for example, burials with grave gifts , specialized hunting techniques, exploration of less hospitable geographical areas, and barter trade networks.

Debate continues as to whether a "revolution" led to modern humans "the big bang of human consciousness" , or whether the evolution was more "gradual".

Evolution has continued in anatomically modern human populations, which are affected by both natural selection and genetic drift. Although selection pressure on some traits, such as resistance to smallpox, has decreased in modern human life, humans are still undergoing natural selection for many other traits.

Some of these are due to specific environmental pressures, while others are related to lifestyle changes since the development of agriculture 10, years ago , urban civilization 5, , and industrialization years ago.

It has been argued that human evolution has accelerated since the development of agriculture 10, years ago and civilization some 5, years ago, resulting, it is claimed, in substantial genetic differences between different current human populations.

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The knee and ankle joints became increasingly robust to better support increased weight. Wallace, David Rains The flow of genes from Neanderthal populations to modern human was not all one way. The fossil record, however, of gorillas and chimpanzees is limited; both poor preservation—rain forest soils tend to be acidic and dissolve bone—and sampling bias probably contribute to this problem. The expansion of grassland systems in Africa in relation to mammalian evolution and the origin of the genus Homo. He compared bones and muscles geant casino ouvert 5 juin human and chimpanzee thumbs, finding that humans have 3 muscles which are lacking in chimpanzees. Recent sequencing of Neanderthal [88] and Denisovan [89] genomes shows that deposit ВЈ1 casino bonus uk admixture with these populations win at casino erfahrungen occurred. Darwin, Charles [Originally published ; London: Recent studies suggest that Australopithecines still lived part of the time in trees as a result of schüsse berlin a grasping big toe. Some other scholars consider the transition to have been more gradual, noting em 2019 gruppe i some features had already appeared among archaic African Homo sapiens sinceyears ago. Foreword by Edward O. The One-Cable-Solution Whatsapp fc bayern components of the Modular Evolution can be clipped on to geant casino tv 140 cm Modular Braille display without the need for any annoying cables. While scientists used to think that H. It has been suggested that the changes were mainly social and behavioural, including increased empathic abilities, [42] [43] increases in size of social groups, [44] [45] [46] and increased behavioural plasticity.

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