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| Britannica Concise Encyclopedia: human evolution |
For more information on human evolution, visit Britannica.com.
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| World of the Body: human evolution |
The human body is the end product of a long period of evolution, stretching back millions of years. In the case of some aspects of our body, the ancestry goes back not just a few million years, but hundreds of millions. The basic layout of the human body, for example, is that of the vertebrates (being bilaterally symmetrical, organized around the backbone) and of the reptiles and amphibians (in having a pair of hind limbs and forelimbs, each with five digits — fingers or toes — at the end of them). But like every species, humans have a shape that is unique to themselves. Among primates, our closest evolutionary relatives, humans have three features that stand out — upright posture and walking, a relatively large brain, and relative hairlessness.
Most primates live in trees, and they do so like all mammals by using all four limbs (or in the case of spider monkeys, five — their tails are also prehensile and can grasp things). Their hands and feet can both be used for grasping. In this sense, all non-human primates are quadrupedal. In the case of something like an orang utan, legs and arms, hands and feet are equally mobile and dextrous, and in a way all act more like arms than legs — for holding and grasping, rather than support. With baboons the forelimbs and hindlimbs are both rather leg-like, and support the animal as it moves quadrupedally over the ground, rather like a dog. For the gibbon, the only truly arm-swinging primate, the arms are long and flexible, and the legs, short and reduced — basically to get them out of the way as the owner brachiates through the trees.
Everything about the human body is either a retention of these basic characteristics, or else has been modified by evolution. The grasping hand, the relatively mobile shoulder, the eyes that look forward with stereoscopic vision, are all part of the human being's ancient primate heritage. Each evolved for some reason in our past, long before any movement towards the human condition, but has remained useful and has been built upon. The close-set eyes that look directly forward, with overlapping fields of vision, evolved among the earlier primates, to allow them to judge distances in three-dimensional space — an essential part of leaping perilously from one tree branch to another. The ability to co-ordinate this vision with dextrous hand movements is an old evolutionary heritage, but one that is used every time we catch a ball or calculate whether it is safe to overtake a car at 100 km per hour.
While our body is a cumulative and often messy mix of this ancient past, it is also the product of a unique evolutionary history shared with no other living primate. It is often said that humans are the most generalized of species, lacking all the specializations that characterize other animals such as giraffes, with their long necks, or elephants, with their trunks. In actual fact, as primates we are very specialized in one way — bipedalism. Unlike virtually all other primates, we are highly dedicated ground-dwellers, and indeed are fairly poor at climbing and clambering in trees. Our ability to walk upright habitually and easily is our most distinctive and in many ways most divergent characteristic. It has also shaped virtually all aspects of our body, from head to toe. Our foot is effectively a highly sprung platform, with arches in two directions to take the endless pounding of hitting the ground, and to push off into the next stride. It is heavily built compared with the feet of monkeys and apes, and has lost any ability to grasp. The knee is also built to take pressure, being large, and heavily constrained in sideways movement. The leg as a whole is very long, to ensure a large stride. The pelvis is perhaps the most modified part of the body, being turned from a long baton for connecting upper and lower parts of the body, to a large bowl to take all the weight of the upper body, which is now resting entirely on two legs. The vertebral column is also robust. Unlike the back of a quadruped, which is built with a single arch like a cantilevered bridge, the human spine is S-shaped. The head is also modified, being perched more vertically on the spine.
The overall impression of a human from an evolutionary perspective is a tall, cylindical shape, a linear design. There has been considerable debate as to the evolutionary pressures that have shaped the human body, and it looks as if there are two main factors involved. The first is that bipedalism is an energy-saving way of moving on the ground: since our ancestors had to cope with the disappearance of forests, and search widely for food in dry African environments, it was the most evolutionarily effective way, turning an arm-swinging, tree-dwelling ape into a terrestrial specialist. The other factor is temperature. The open savannas where the earliest bipeds evolved were hot, with little shade, and the effect of the sun would have been severe. One of the effects of an upright posture is to reduce the area of the body that receives direct sunlight, and to remove more of it away from the reflected heat of the ground. The human body, then, was forged by selection in the heat of the more open plains of Africa.
Evolution is the process of change over time, over thousands and millions of years. The fossil record has shown that the basics of bipedalism go right back to the roots of our evolutionary history, back to over four million years ago, soon (in evolutionary terms) after our ancestors diverged from the ancestors of the living chimpanzees, our closest relatives. The modern form of bipedalism, with the cylindrical, linear pattern, is probably about two million years old. With bipedalism would have come other changes. The hand, no longer needed to support the body in movement, became the highly dextrous and finely-tuned structure that we use today for so many activities.
The upright stance is such a universal and uniform human characteristic that it is taken totally for granted: it is the essence of humanity. Around the world, though, the human body comes in enormous variety — tall, short, fat, thin, hairy, smooth, dark, and light. Unlike the basic upright body plan, these variations are not millions of years old, but just a few tens of thousands or even less. But they are still the product of evolution. Once again the environment has played a major part. Although humans vary in the amount of hair cover they have, they are, by comparison with apes, largely hairless. This is again a response to heat. Humans have evolved a copious sweating system — we use the evaporation of moisture from the skin to cool our body, and this works more effectively where the air can move freely over the skin — that is, where there is no hair. As a whole, therefore, the species is ‘naked’ — not actually hairless, but with a miniaturized hair cover. And those people who have a long history of living in the hotter parts of the world are the most hairless. Skin colour follows this pattern, with darker skins, produced by higher levels of melanin, acting as a compensatory mechanism to reduce the effect of high levels of solar radiation on the skin. Body shape is also affected by the environment — larger, shorter-limbed bodies are better at keeping in heat, where thin, long-limbed individuals are better at dissipating heat. As a result, people who live at higher latitude have shorter limbs, and are often robustly built; people in the tropics are small, linear, and lean.
While the human body has evolved to suit the environment, especially the temperature, it has been affected by one other major factor — sex. Evolution is driven by selection — the survival of those best suited to the environment — but Darwin pointed out that there were two elements to this; natural selection and sexual selection. Most of the characteristics described so far have been the product of natural selection, but much of the human body is probably the result of how males and females have chosen their mates, and how well they are able to reproduce. Out of this has arisen the differences between the sexes. Some of these differences have a direct function — women have wider hips than men, compensating for the narrower birth outlet forced by bipedalism. Others are probably related to the preferences of men or women — larger breasts and curvaceous hips in women, for example. These secondary sexual characteristics may have their basis in some function, but are as much a signal and a symbol, and selected as such — in this case, a signal of fertility. Men also give signals with their bodies — simple ones related to strength and size, but also more subtle ones, such as grey hair or baldness as a sign of having lived a long time — and therefore being a successful male. Most characteristics, though, are a mixture of the sexual and functional. Men often prefer women who are more curvaceous, which is often related to fat deposition — women lay down fat more easily than men. This fat is also necessary for ensuring that a woman is well-nourished, and thus better able to withstand the costs of pregnancy and lactation. Women may prefer large, strong men, but such men may also be better at other things, such as hunting or fighting, and thus better adapted.
In the end, the evolution of the human body is a seamless mix of sex, reproduction, activity, and environment; it is also a mix of the very old and the very new, and over evolutionary time has changed and shifted. In some ways it is a sleek and efficient machine; in others, it is full of flaws. In this sense it is like any other evolutionary product, a compromise between all the demands placed on it during the course of the many different lives that humans have to live, have lived in the past, and will live in the future.
— Robert Foley
Bibliography
See also bipedalism; evolution; heredity, language and the brain.
| Columbia Encyclopedia: human evolution |
The Evolutionary Tree
Humans are mammals of the Primate order. The earliest primates evolved about 65 million years ago in the geological period known as the Paleocene epoch. They were small-brained, arboreal fruit eaters, similar to modern tree shrews. Primates of the Eocene epoch (55 to 38 million years ago) were similar and ancestral to contemporary tarsiers, lemurs, and tree shrews, and are classified as lower primates or prosimians. During the late Eocene, the higher primates, or anthropoids, developed from prosimian ancestors and, aided by continental drift, diverged into New World (or platyrrhine) and Old World (or catarrhine) monkeys. The branching of Old World monkeys and hominoids apparently occurred in the late Oligocene (38 to 25 million years ago) or early Miocene (25 to 8 million years ago), a time period poorly represented in the fossil record. The lesser apes (gibbons and siamangs) and other hominoid lines diverged about 20 million years ago, while the Asian great apes (the orangutan being the only surviving form) diverged from the African hominoids about 15 to 10 million years ago. Genetic evidence suggests that the ancestral lines of gorillas diverged about 8 million years ago and that chimpanzees and hominids diverged about 5 million years ago.
Hominid Evolution
The earliest known hominids are members of the genus Australopithecus, the earliest of which date to more than 4 million years ago. Unlike other primates, but like all hominids, australopithecines were bipedal. Their crania, however, were small and apelike, with an average cranial capacity of about 450 cc in the gracile species and 600 cc in the robust forms. Australopithecines that have been considered ancestral in the lineage leading to the human genus Homo include A. afarensis (an important skeleton of which is popularly known as Lucy) and A. africanus. The exact position of these and other early species on the hominid family tree continues to be disputed.
The first member of the genus Homo, a small gracile species known as H. habilis, was present in east Africa at least 2 million years ago. H. habilis was the first hominid to exhibit the marked expansion of the brain (with an average cranial capacity of about 750 cc) that would become a hallmark of subsequent hominid evolutionary history. By about 1.6 million years ago, H. habilis had evolved into a larger, more robust, and larger-brained species known as Homo erectus. Cranial capacities ranged from about 900 cc in early specimens to 1050 cc in later ones. H. erectus persisted for well over a million years and migrated off the African continent into Asia, Indonesia, and Europe.
Between 500,000 and 250,000 years ago, H. erectus evolved into H. sapiens. Transitional forms between H. erectus and H. sapiens are referred to as archaic H. sapiens. With the exception of H. sapiens neandertalensis (see Neanderthal man), no additional subspecies are recognized. Indeed, some scientists consider Neanderthal a separate species. Archaic H. sapiens changed gradually, becoming somewhat larger, more gracile and larger-brained through time. Cranial capacity, for example, increased from about 1150 cc in early transitional forms to the current world average of just over 1350 cc. By 150,000 years ago in Africa and Asia and 28,000 years ago in Europe (see Cro-Magnon man), the transition to H. sapiens was complete, and fully modern humans became the single surviving hominid species.
The Evolution of Culture
Among hominids, a parallel evolutionary process involving increased intelligence and cultural complexity is apparent in the material record. Evidence of greater behavioral flexibility and adaptability presumably reflects the decreased influence of genetically encoded behaviors and the increased importance of learning and social interaction in transmitting and maintaining behavioral adaptations (see culture). Because the organization of neural circuitry is more significant than overall cranial capacity in establishing mental capabilities, direct inferences from the fossil record are likely to be misleading. Contemporary humans, for example, exhibit considerable variability in cranial capacity (1150 cc to 1600 cc), none of which is related to intelligence.
Tool use was once thought to be the hallmark of members of the genus Homo, beginning with H. habilis, but is now known to be common among chimpanzees. The earliest stone tools of the lower Paleolithic, known as Oldowan tools and dating to about 2 to 2.5 million years ago, were once thought to have been manufactured by H. habilis. Recent finds suggest that Oldowan tools may also have been made by robust australopithecines. The simultaneous emergence of H. erectus and the more complex Achuelian tool tradition may indicate shifting adaptations as much as increased intelligence.
While it is clear that H. erectus was much more versatile than any of its predecessors, adapting its technologies and behaviors to diverse environmental conditions, the extent and limitations of its intellectual endowment remain a subject of heated debate. This is also the case for both archaic H. sapiens and Neanderthals, the latter associated with the more sophisticated technologies of the middle Paleolithic. However impressive the achievements of H. erectus and early H. sapiens, most material remains predating 40,000 years ago reflect utilitarian concerns. Nonetheless, there is now scattered African archaeological evidence from before that time (in one case as early as 90,000 years ago) of the production by H. sapiens of beads and other decorative work, perhaps indicating a gradual development of the aesthetic concerns and other symbolic thinking characteristic of later human societies. Whether the emergence of modern H. sapiens corresponds to the explosion of technological innovations and artistic activities associated with Cro-Magnon culture or was a more prolonged process of development is a subject of archaeological debate.
Bibliography
See R. Lewin, Human Evolution (2d ed. 1989) and, with R. Leakey, Origins Reconsidered (1992); I. Tattersall, The Fossil Trail: How We Know What We Think We Know about Human Evolution (1995); A. Walker and P. Shipman, The Wisdom of the Bones: In Search of Human Origins (1996); C. Stringer and R. McKie, African Exodus: The Origins of Modern Humanity (1997); L. R. Berger and B. Hilton-Barber, In the Footsteps of Eve: The Mystery of Human Origins (2000); I. Tattersall and J. H. Schwartz, Extinct Humans (2000).
| Wikipedia: Human evolution |
Human evolution, or anthropogenesis, is the origin and evolution of Homo sapiens as a distinct species from other hominids, great apes and placental mammals. The study of human evolution encompasses many scientific disciplines, including physical anthropology, primatology, archaeology, linguistics and genetics.[1]
The term "human" in the context of human evolution refers to the genus Homo, but studies of human evolution usually include other hominids, such as the Australopithecines. The genus Homo had diverged from the Australopithecines by about 2.3 to 2.4 million years ago in Africa.[2][3] Scientists have estimated that humans branched off from their common ancestor with chimpanzees - the only other living hominins - about 5–7 million years ago. Several species of Homo evolved and are now extinct. These include Homo erectus, which inhabited Asia, and Homo neanderthalensis, which inhabited Europe. Archaic Homo sapiens evolved between 400,000 and 250,000 years ago.
The dominant view among scientists concerning the origin of anatomically modern humans is the "Out of Africa" or recent African origin hypothesis,[4][5][6][7] which argues that H. sapiens arose in Africa and migrated out the continent around 50-100,000 years ago, replacing populations of H. erectus in Asia and H. neanderthalensis in Europe. Scientists supporting the alternative multiregional hypothesis argue that H. sapiens evolved as geographically separate but interbreeding populations stemming from a worldwide migration of H. erectus out of Africa nearly 2.5 million years ago.
The word homo, the name of the biological genus to which humans belong, is Latin for "human". It was chosen originally by Carolus 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".[8]
Carolus Linnaeus and other scientists of his time also considered the great apes to be the closest relatives of human beings due to morphological and anatomical similarities. The possibility of linking humans with earlier apes by descent only became clear after 1859 with the publication of Charles Darwin's On the Origin of Species. This 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".
The first debates about the nature of human evolution arose between Thomas 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 agree that the origin of the mental capacities and the moral sensibilities of humans could be explained by natural selection. Darwin applied the theory of evolution and sexual selection to humans when he published The Descent of Man in 1871.[9]
A major problem was the lack of fossil intermediaries. It was only in the 1920s that such fossils were discovered in Africa. In 1925, Raymond Dart described Australopithecus africanus. The type specimen was the Taung Child, an Australopithecine infant discovered in a cave. The child's remains were a remarkably well-preserved tiny skull and an endocranial cast of the individual's brain. Although the brain was small (410 cm³), 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 was evidence of bipedal locomotion. All of these traits convinced Dart that the Taung baby was a bipedal human ancestor, a transitional form between apes and humans.
The classification of humans and their relatives has changed considerably over time. The gracile Australopithecines are now thought to be ancestors of the genus Homo, the group to which modern humans belong. Both Australopithecines and Homo sapiens are part of the tribe Hominini. Recent data suggests Australopithecines were a diverse group and that A. africanus may not be a direct ancestor of modern humans. Reclassification of Australopithecines that originally were split into either gracile or robust varieties has put the latter into a family of its own, Paranthropus. Taxonomists place humans, Australopithecines and related species in the same family as other great apes, in the Hominidae.
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Note: 1e +06 years = 1 million years = 1 Ma.
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The evolutionary history of the primates can be traced back 65 million years, as one of the oldest of all surviving placental mammal groups. The oldest known primates come from North America, but they were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and Eocene.
With the beginning of modern climates, marked by the formation of the first Antarctic ice in the early Oligocene around 30 million years ago, primates went extinct everywhere but Africa and southern Asia.[citation needed] A primate from this time was Notharctus. Fossil evidence found in Germany in the 1980s was determined to be about 16.5 million years old, some 1.5 million years older than similar species from East Africa and challenging the original theory regarding human ancestry originating on the African continent.
Begun[10] says that these primates flourished in Eurasia and that the lineage leading to the African apes and humans— including Dryopithecus—migrated south from Europe or Western Asia into Africa. The surviving tropical population, which is seen most completely in the upper Eocene and lowermost Oligocene fossil beds of the Fayum depression southwest of Cairo, gave rise to all living primates—lemurs of Madagascar, lorises of Southeast Asia, galagos or "bush babies" of Africa, and the anthropoids; platyrrhines or New World monkeys, and catarrhines or Old World monkeys and the great apes and humans.
The earliest known catarrhine is Kamoyapithecus from uppermost Oligocene at Eragaleit in the northern Kenya Rift Valley, dated to 24 million years ago. Its ancestry is generally thought to be species related to Aegyptopithecus, Propliopithecus, and Parapithecus from the Fayum, at around 35 million years ago. There are no fossils from the intervening 11 million years.
In the early Miocene, after 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 age from sites far distant—Otavipithecus from cave deposits in Namibia, and Pierolapithecus and Dryopithecus from France, Spain and Austria—is evidence of a wide diversity of forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the early and middle Miocene. The youngest of the Miocene hominoids, Oreopithecus, is from 9 million year old coal beds in Italy.
Molecular evidence indicates that the lineage of gibbons (family Hylobatidae) became distinct from Great Apes between 18 and 12 million years ago, and that of orangutans (subfamily Ponginae) became distinct 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 Ramapithecus 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 98.4% identical to that of chimpanzees when comparing single nucleotide polymorphisms (see Human evolutionary genetics). The fossil record of gorillas and chimpanzees is quite limited. Both poor preservation (rain forest soils tend to be acidic and dissolve bone) and sampling bias probably contribute to this problem.
Other hominines likely adapted to the drier environments outside the equatorial belt, along with antelopes, hyenas, dogs, pigs, elephants, and horses. The equatorial belt contracted after about 8 million years ago. Fossils of these hominans - the species in the human lineage following divergence from the chimpanzees - are relatively well known. The earliest are Sahelanthropus tchadensis (7 Ma) and Orrorin tugenensis (6 Ma), followed by:
Homo sapiens is the only non-extinct species of its genus, Homo. There were other Homo species, all of which are now extinct. While some of these other species might have been ancestors of H. sapiens, many were likely our "cousins", having speciated away from our ancestral line.[11] There is not yet a consensus as to which of these groups should count as separate species and which as subspecies. In some cases this is due to the paucity of fossils, in other cases it is due to the slight differences used to classify species in the Homo genus. The Sahara pump theory (describing an occasionally passable "wet" Sahara Desert) provides an explanation of the early variation in the genus Homo.
Based on archaeological and paleontological evidence, it has been possible to infer the ancient dietary practices of various Homo species and to study the role of diet in physical and behavioral evolution within Homo.[12][13][14][15] [16]
H. habilis lived from about 2.4 to 1.4 Ma. H. habilis, the first species of the genus Homo, evolved in South and East Africa in the late Pliocene or early Pleistocene, 2.5–2 Ma, when it diverged from the Australopithecines. H. habilis had smaller molars and larger brains than the Australopithecines, and made tools from stone and perhaps animal bones. One of the first known hominids, it was nicknamed 'handy man' by its discoverer, Louis Leakey due to its association with stone tools. 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 H. sapiens.[17]
These are proposed species names for fossils from about 1.9–1.6 Ma, the relation of which with H. habilis is not yet clear.
The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in 1891 on the Indonesian island of Java. He originally gave the material the name Pithecanthropus erectus based on its morphology that he considered to be intermediate between that of humans and apes.[22] H. erectus lived from about 1.8 Ma to about 70,000 years ago (which would indicate that they were probably wiped out by the Toba catastrophe). Often the early phase, from 1.8 to 1.25 Ma, is considered to be a separate species, H. ergaster, or it is seen as a subspecies of H. erectus, Homo erectus ergaster.
In the early Pleistocene, 1.5–1 Ma, in Africa, Asia, and Europe, some populations of Homo habilis are thought to have evolved larger brains and made more elaborate stone tools; these differences and others are sufficient for anthropologists to classify them as a new species, H. erectus. In addition H. erectus was the first human ancestor to walk truly upright.[23] This was made possible by the evolution of locking knees and a different location of the foramen magnum (the hole in the skull where the spine enters). They may have used fire to cook their meat.
A famous example of Homo erectus is Peking Man; others were found in Asia (notably in Indonesia), Africa, and Europe. Many paleoanthropologists now use the term Homo ergaster for the non-Asian forms of this group, and reserving H. erectus only for those fossils found in the Asian region and meeting certain skeletal and dental requirements which differ slightly from H. ergaster.
These are proposed as species that may be intermediate between H. erectus and H. heidelbergensis.
H. heidelbergensis (Heidelberg Man) lived from about 800,000 to about 300,000 years ago. Also proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.[27]
H. neanderthalensis lived from 400,000 [29] or about 250,000 to as recent as 30,000[citation needed]years ago. Also proposed as Homo sapiens neanderthalensis: there is ongoing debate over whether the Neanderthal Man was a separate species, Homo neanderthalensis, or a subspecies of H. sapiens[30] While the debate remains unsettled, evidence from sequencing mitochondrial DNA indicates that no significant gene flow occurred between H. neanderthalensis and H. sapiens, and, therefore, the two were separate species that shared a common ancestor about 660,000 years ago.[31][32] In 1997, Mark Stoneking stated: "These results [based on mitochondrial DNA extracted from Neanderthal bone] indicate that Neanderthals did not contribute mitochondrial DNA to modern humans… Neanderthals are not our ancestors." Subsequent investigation of a second source of Neanderthal DNA supported these findings.[33] However, supporters of the multiregional hypothesis point to recent studies indicating non-African nuclear DNA heritage dating to one Ma,[34] although the reliability of these studies has been questioned.[35] Competition from Homo sapiens probably contributed to Neanderthal extinction.[36][37]
H. sapiens ("sapiens" is Latin for wise or intelligent) has lived from about 250,000 years ago to the present. Between 400,000 years ago and the second interglacial period in the Middle Pleistocene, around 250,000 years ago, the trend in skull expansion and the elaboration of stone tool technologies developed, providing evidence for a transition from H. erectus to H. sapiens. The direct evidence suggests there was a migration of H. erectus out of Africa, then a further speciation of H. sapiens from H. erectus in Africa. A subsequent migration within and out of Africa eventually replaced the earlier dispersed H. erectus. This migration and origin theory is usually referred to as the recent single origin or Out of Africa theory. Current evidence does not preclude some multiregional evolution or some admixture of the migrant H. sapiens with existing Homo populations. This is a hotly debated area of paleoanthropology.
Current research has established that human beings are genetically highly homogenous; that is, the DNA of individuals is more alike than usual for most species, which may have resulted from their relatively recent evolution or the possibility of a population bottleneck resulting from cataclysmic natural events such as the Toba catastrophe.[38][39][40] Distinctive genetic characteristics have arisen, however, primarily as the result of small groups of people moving into new environmental circumstances. These adapted traits are a very small component of the Homo sapiens genome, but include various characteristics such as skin color and nose form, in addition to internal characteristics such as the ability to breathe more efficiently in high altitudes.
H. sapiens idaltu, from Ethiopia, is a possible extinct sub-species who lived from about 160,000 years ago. It is the oldest known anatomically modern human.[citation needed]
H. floresiensis, which lived from approximately 100,000 to 12,000 before present, has been nicknamed hobbit for its small size, possibly a result of insular dwarfism.[41] H. floresiensis is intriguing both for its size and its age, being a concrete example of a recent species of the genus Homo that exhibits derived traits not shared with modern humans. In other words, H. floresiensis share a common ancestor with modern humans, but split from the modern human lineage and followed a distinct evolutionary path. The main find was a skeleton believed to be a woman of about 30 years of age. Found in 2003 it has been dated to approximately 18,000 years old. The living woman was estimated to be one meter in height, with a brain volume of just 380 cm3 (considered small for a chimpanzee and less than a third of the H. sapiens average of 1400 cm3).
However, there is an ongoing debate over whether H. floresiensis is indeed a separate species.[42] Some scientists presently believe that H. floresiensis was a modern H. sapiens suffering from pathological dwarfism.[43] This hypothesis is supported in part, because some modern humans who live on Flores, the island where the skeleton was found, are pygmies. This coupled with pathological dwarfism could indeed create a hobbit-like human. The other major attack on H. floresiensis is that it was found with tools only associated with H. sapiens.[43]
| Species | Lived when (Ma) | Lived where | Adult height | Adult mass | Brain volume (cm³) | Fossil record | Discovery / publication of name |
|---|---|---|---|---|---|---|---|
| H. habilis | 2.2 – 1.6 | Africa | 1.0–1.5 m (3.3–4.9 ft) | 33–55 kg (73–120 lb) | 660 | Many | 1960/1964 |
| H. erectus | 1.4 – 0.2 | Africa, Eurasia (Java, China, Caucasus) | 1.8 m (5.9 ft) | 60 kg (130 lb) | 850 (early) – 1,100 (late) | Many | 1891/1892 |
| H. rudolfensis | 1.9 | Kenya | 1 skull | 1972/1986 | |||
| H. georgicus | 1.8 | Georgia | 600 | Few | 1999/2002 | ||
| H. ergaster | 1.9 – 1.4 | Eastern and Southern Africa | 1.9 m (6.2 ft) | 700–850 | Many | 1975 | |
| H. antecessor | 1.2 – 0.8 | Spain | 1.75 m (5.7 ft) | 90 kg (200 lb) | 1,000 | 2 sites | 1997 |
| H. cepranensis | 0.9 – 0.8? | Italy | 1,000 | 1 skull cap | 1994/2003 | ||
| H. heidelbergensis | 0.6 – 0.35 | Europe, Africa, China | 1.8 m (5.9 ft) | 60 kg (130 lb) | 1,100–1,400 | Many | 1908 |
| H. neanderthalensis | 0.35 – 0.03 | Europe, Western Asia | 1.6 m (5.2 ft) | 55–70 kg (120–150 lb) (heavily built) | 1,200–1,900 | Many | (1829)/1864 |
| H. rhodesiensis | 0.3 – 0.12 | Zambia | 1,300 | Very few | 1921 | ||
| H. sapiens sapiens | 0.2 – present | Worldwide | 1.4–1.9 m (4.6–6.2 ft) | 50–100 kg (110–220 lb) | 1,000–1,850 | Still living | —/1758 |
| H. sapiens idaltu | 0.16 – 0.15 | Ethiopia | 1,450 | 3 craniums | 1997/2003 | ||
| H. floresiensis | 0.10 – 0.012 | Indonesia | 1.0 m (3.3 ft) | 25 kg (55 lb) | 400 | 7 individuals | 2003/2004 |
Using 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—most notably the continued expansion of the human brain. Paleontology has yet to explain the expansion of this organ over millions of years despite being extremely demanding in terms of energy consumption. The brain of a modern human consumes about 20 watts (400 kilocalories per day), which is one fifth of the energy consumption of a human body. Increased tool use would allow hunting for energy-rich meat products, and would enable processing more energy-rich plant products. Researchers have suggested that early hominids were thus under evolutionary pressure to increase their capacity to create and use tools.[44]
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. There is some evidence that the australopithecines (4 Ma) may have used broken bones as tools, but this is debated.
It should be noted that many species make and use tools, but it is the human species that dominates the areas of making and using more complex tools. A good question is, what species made and used the first tools? The oldest known tools are the "Oldowan stone tools" from Ethiopia. It was discovered that these tools are from 2.5 to 2.6 million years old, which predates the earliest known "Homo" species. There is no known evidence that any "Homo" specimens appeared by 2.5 Ma. A Homo fossil was found near some Oldowan tools, and its age was noted at 2.3 million years old, suggesting that maybe the Homo species did indeed create and use these tools. It is surely possible, but not solid evidence. Bernard Wood noted that "Paranthropus" coexisted 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 that points to 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.[45]
In 1994, 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 that chimps lack. Humans also have thicker metacarpals with broader heads, making the human hand more successful at precision grasping than the chimpanzee hand. Susman defended that modern anatomy of the human thumb is an evolutionary response to the requirements associated with making and handling tools and that both species were indeed toolmakers.[45]
Stone tools are first attested around 2.6 Ma, when H. habilis in Eastern Africa used so-called pebble tools, choppers made out of round pebbles that had been split by simple strikes.[46] This marks the beginning of the Paleolithic, or Old Stone Age; its end is taken to be the end of the last Ice Age, around 10,000 years ago. The Paleolithic is subdivided into the Lower Paleolithic (Early Stone Age, ending around 350,000–300,000 years ago), the Middle Paleolithic (Middle Stone Age, until 50,000–30,000 years ago), and the Upper Paleolithic.
The period from 700,000–300,000 years ago is also known as the Acheulean, when H. ergaster (or erectus) made large stone hand-axes out of flint and quartzite, at first quite rough (Early Acheulian), later "retouched" by additional, more subtle strikes at the sides of the flakes. After 350,000 BP (Before Present) the more refined so-called Levallois technique was developed. It consisted of a series of consecutive strikes, by which scrapers, slicers ("racloirs"), needles, and flattened needles were made.[46] Finally, after about 50,000 BP, ever more refined and specialized flint tools were made by the Neanderthals and the immigrant Cro-Magnons (knives, blades, skimmers). In this period they also started to make tools out of bone.
Until about 50,000–40,000 years ago the use of stone tools seems to have progressed stepwise. Each phase (H. habilis, H. ergaster, H. neanderthalensis) started at a higher level than the previous one, but once that phase started further development was slow. These Homo species were culturally conservative, but after 50,000 BP modern human culture started to change at a much greater speed. Jared Diamond, author of The Third Chimpanzee, and other anthropologists characterize this as a "Great Leap Forward."
Modern humans started burying their dead, making clothing out of hides, developing sophisticated hunting techniques (such as using trapping pits or driving animals off cliffs)[citation needed], and engaging in cave painting.[47] As human culture advanced, different populations of humans introduced novelty to existing technologies: artifacts such as fish hooks, buttons and bone needles show signs of variation among different populations of humans, something that had not been seen in human cultures prior to 50,000 BP. Typically, H. neanderthalensis populations do not vary in their technologies.
Modern human behavior includes four aspects: abstract thinking (concepts free from specific examples), planning (taking steps to achieve a further goal), innovation (finding new solutions), and symbolic behaviour (such as images and rituals)[citation needed]. Among concrete examples of modern human behavior, anthropologists include specialization of tools, use of jewelery 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.[48]
Today, all humans belong to one, undivided by species barrier, population of Homo sapiens sapiens. However, according to the "Out of Africa" model this is not the first species of hominids: the first species of genus Homo, Homo habilis, evolved in East Africa at least 2 Ma, and members of this species populated different parts of Africa in a relatively short time. Homo erectus evolved more than 1.8 Ma, and by 1.5 Ma had spread throughout the Old World.
Anthropologists have been divided as to whether current human population evolved as one interconnected population (as postulated by the Multiregional Evolution hypothesis), or evolved only in East Africa, speciated, and then migrating out of Africa and replaced human populations in Eurasia (called the "Out of Africa" Model or the "Complete Replacement" Model).
Multiregional evolution, a model to account for the pattern of human evolution, was proposed by Milford H. Wolpoff[49] in 1988[50]. Multiregional evolution holds that human evolution from the beginning of the Pleistocene 2.5 million years BP to the present day has been within a single, continuous human species, evolving worldwide to modern Homo sapiens.
According to the multiregional hypothesis, fossil and genomic data are evidence for worldwide human evolution and contradict the recent speciation postulated by the Recent African origin hypothesis. The fossil evidence was insufficient for Richard Leakey to resolve this debate.[51]. Studies of haplogroups in Y-chromosomal DNA and mitochondrial DNA have largely supported a recent African origin.[52] Evidence from autosomal DNA also supports the Recent African origin. However the presence of archaic admixture in modern humans remains a possibility and has been suggested by some studies.[53]
According to the Out of Africa model, developed by Chris Stringer and Peter Andrews, modern H. sapiens evolved in Africa 200,000 years ago. Homo sapiens began migrating from Africa between 70,000 – 50,000 years ago and eventually replaced existing hominid species in Europe and Asia.[54][55] Out of Africa has gained support from research using mitochondrial DNA (mtDNA). After analysing genealogy trees constructed using 133 types of mtDNA, researchers concluded that all were descended from a woman from Africa, dubbed Mitochondrial Eve. Out of Africa is also supported by the fact that mitochondrial genetic diversity is highest among African populations.[56]
There are differing theories on whether there was a single exodus or several. A multiple dispersal model involves the Southern Dispersal theory,[57] 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 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. A second wave of humans dispersed across the Sinai peninsula into Asia, resulting in the bulk of human population for Eurasia. This second group possessed a more sophisticated 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 the Holocene era.[57] 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 lineages, which support a single migration out of Africa that gave rise to all non-African populations.[58]
The broad study of African genetic diversity headed by Dr.Sarah Tishkoff found the San people to express the greatest genetic diversity among the 113 distinct populations sampled, making them one of 14 "ancestral population clusters".The research also located the origin of modern human migration in south-western Africa, near the coastal border of Namibia and Angola.[59]
Natural selection has been observed in contemporary human populations. Evolutionary trends reported include a lengthening of the reproductive period, reduction in cholesterol levels, blood glucose and blood pressure.[60]
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.
This list is in chronological order across the page by genus.
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