Human Evolution, or anthropogenesis, is the part of biological evolution concerning the emergence of Homo sapiens as a distinct species from other hominans, great apes and placental mammals.
It is the subject of a broad scientific inquiry that seeks to understand and describe how this change occurred.
The study of human evolution encompasses many scientific disciplines, most notably physical anthropology, linguistics and genetics.
The term "human", in the context of human evolution, refers to the genus Homo, but studies of human evolution usually include other hominins, such as the australopithecines.
The Homo genus diverged from the australopithecines about 2 million years ago in Africa. Several typological species of Homo, now extinct, evolved. 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. Many scientists favor the view that H. sapiens evolved in Africa and spread across the globe, replacing populations of H. erectus and H. neanderthalensis.
Others view modern humans as having evolved as a single, widespread population from existing Homo species, particularly H. erectus. The fossil evidence so far is insufficient to resolve this vigorous debate.
Starting with H. habilis, humans began using stone tools of increasing sophistication. About 50,000 years ago, human technology and culture began to change more rapidly.
Paleoanthropology is the study of ancient humans based on fossil evidence, tools, and other signs of human habitation. The modern field of paleoanthropology began with the discovery of a Neanderthal skeleton in 1856, although there had been finds elsewhere since 1830. By 1859, the morphological similarity of humans to certain great apes had been discussed and argued for some time, but the idea of the biological evolution of species in general was not legitimized until Charles Darwin published On the Origin of Species in November of that year. Darwin's first book on evolution did not address the specific question of human evolution: "Light will be thrown on the origin of man and his history", was all Darwin wrote on the subject. Nevertheless, the implications of evolutionary theory were clear to contemporary readers. Debates between Thomas Huxley and Richard Owen focused on human evolution. Huxley convincingly illustrated many of the similarities and differences between humans and apes in his 1863 book Evidence as to Man's Place in Nature. By the time Darwin published his own book on the subject, The Descent of Man, it was already a well-known interpretation of his theory, and the interpretation which made the theory highly controversial. Even many of Darwin's original supporters (such as Alfred Russel Wallace and Charles Lyell) did not like the idea that human beings could have evolved their impressive mental capacities and moral sensibilities through natural selection. Since the time of Carolus Linnaeus, scientists have considered the great apes to be the closest relatives of human beings because they look very similar. In the 19th century, they speculated that the closest living relatives of humans are chimpanzees. Based on the natural range of these creatures, they surmised that humans share a common ancestor with other African great apes and that fossils of these ancestors would be found in Africa. It is now accepted by virtually all biologists that humans are not only similar to the great apes but, in fact, are great apes.Classification of Hominidae 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. This cave, in Taung, South Africa, was being mined for raw materials used to make concrete. 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. Another 20 years would pass before Dart's claims were taken seriously, following the discovery of more fossils that resembled his find. The prevailing view of the time was that a large brain evolved before bipedality. It was thought that intelligence on par with modern humans was a prerequisite to bipedalism. The australopithecines are now thought to be immediate ancestors of the genus Homo, the group to which modern humans belong. Both australopithecines and Homo sapiens are part of the tribe Hominini, but recent data has brought into doubt the position of A. africanus as a direct ancestor of modern humans; it may well have been a dead-end cousin. The australopithecines were originally classified as either gracile or robust. The robust variety of Australopithecus has since been reclassified as Paranthropus, although it is still regarded as a subgenus of Australopithecus by some authors. In the 1930s, when the robust specimens were first described, the Paranthropus genus was used. During the 1960s, the robust variety was moved into Australopithecus. The recent trend has been back to the original classification as a separate genus.
History Of Evolution
The evolutionary history of the primates can be traced back for some 85 million years, as one of the oldest of all surviving placental mammal groups. With the beginning of modern climates, marked by the formation of the first Antarctic ice in the early Oligocene around 40 million years ago, primates went extinct everywhere but Africa and southern Asia. One such primate from this time was Notharctus. Fossil evidence found in Germany 20 years ago was determined to be about 16.5 million years old, some 1.5 million years older than similar species from East Africa. It suggests that the primate lineage of the great apes first appeared in Eurasia and not Africa . The discoveries suggest that the early ancestors of the hominids (the family of great apes and humans) migrated to Eurasia from Africa about 17 million years ago, just before these two continents were cut off from each other by an expansion of the Mediterranean Sea. Begun says that these primates flourished in Eurasia and that their lineage leading to the African apes and humans Dryopithecusmigrated 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 mya (millions of years before present). Its ancestry is generally thought to be close to such genera as Aegyptopithecus, Propliopithecus, and Parapithecus from the Fayum, at around 35 mya. There are no fossils from the intervening 11 million years. No near ancestor to South American platyrrhines, whose fossil record begins at around 30 mya, can be identified among the North African fossil species, and possibly lies in other forms that lived in West Africa that were caught up in the still-mysterious transatlantic sweepstakes that sent primates, rodents, boa constrictors, and cichlid fishes from Africa to South America sometime in the Oligocene. In the early Miocene, after 22 mya, many kinds of arboreally adapted primitive catarrhines from East Africa suggest a long history of prior diversification. Because the fossils at 20 mya include fragments attributed to Victoriapithecus, the earliest cercopithecoid; the other forms are (by default) grouped as hominoids, without clear evidence as to which are closest to living apes and humans. Among the presently recognized genera in this group, which ranges up to 13 mya, we find 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 distantOtavipithecus 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 mya coal beds in Italy. Molecular evidence indicates that the lineage of gibbons (family Hylobatidae) became distinct between 18 and 12 Ma, and that of orangutans (subfamily Ponginae) at about 12 Ma; we have no fossils that clearly document the ancestry of gibbons, which may have originated in a so far unknown South East Asian hominid population, but fossil proto-orangutans may be represented by Ramapithecus from India and Griphopithecus from Turkey, dated to around 10 Ma. It has been suggested that species close to last common ancestors 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 mya, first the gorillas, and then the chimpanzee (genus Pan) split off from the line leading to the humans; human DNA is 98.4 percent identical to the DNA of chimpanzees. 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, however, likely adapted (along with antelopes, hyenas, dogs, pigs, elephants, and horses) to the somewhat drier environments outside the equatorial belt (which contracted after about 8 million years ago; reference needed) and their fossils are relatively well known. The earliest are Sahelanthropus tchadensis (7 mya) and Orrorin tugenensis (6 mya), followed by:
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Ardipithecus (5.5-4.4 mya), with species Ar. kadabba and Ar. ramidus;
Australopithecus (4-2 mya), with species Au. anamensis, Au. afarensis, Au. africanus, Au. bahrelghazali, and Au. garhi;
Kenyanthropus (3-2.7 mya), with species Kenyanthropus platyops
Paranthropus (3-1.2 mya), with species P. aethiopicus, P. boisei, and P. robustus;
Homo (2 mya-present), with species Homo habilis, Homo rudolfensis, Homo ergaster, Homo georgicus, Homo antecessor, Homo cepranensis, Homo erectus, Homo heidelbergensis, Homo rhodesiensis, Homo sapiens neanderthalensis, Homo sapiens idaltu, Archaic Homo sapiens, Homo floresiensis.
The word homo is Latin for "human", chosen originally by Carolus Linnaeus in his classification system. It is often translated as "man"; however, this can lead to confusion, given that the English word "man" can refer specifically to males, and such gender-biased usage is increasingly rejected by modern audiences. The word "human" is from the Latin humanus, the adjectival form of homo. (Latin for "man" in the gender-specific sense is vir, as in "virile".) In modern taxonomy, Homo sapiens is the only extant species of its genus, Homo. Likewise, the ongoing study of the origins of Homo sapiens often demonstrates that 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. There is not yet a consensus as to which of these groups should count as separate species and which as subspecies of another species. 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 provides an explanation of the early variation in the genus Homo. There are now two main schools of thought about the factors that drove human evolution. One theory, the Savannah Theory, first propounded by Raymond Dart, says that the arboreal existence was replaced by a move to the savannah for hunting animals. Another theory, which is still strongly disputed by many researchers, is the aquatic ape hypothesis (AAH). This asserts that wading, swimming and diving for food exerted a strong evolutionary effect on the ancestors of the genus Homo and is in part responsible for the split between the common ancestors of humans and other great apes. The AAH attempts to explain the large number of physical differences between humans and other apes such as lack of body hair, larger brains and upright posture in terms of the methods of feeding and the types of food utilized by early hominids living in coastal and river regions. Though no fossil evidence of an aquatic ape has been found, certain physical differences between humans and other apes seem to support the theory, such as the human's subcutaneous layer of fat, webbing between the fingers and toes, vernix caseosa, and hair growth that follows the direction of water flowing over the body.
H. habilis lived from about 2.4 to 1.4 million years ago (mya). 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 mya, 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 (mode 1). Some scientists have proposed moving this species out of Homo and into Australopithecus due to its postcranial morphology being more adapted to an arboreal existence rather than bipedalism than that of H. sapiens.
These are proposed species names for fossils from about 1.9-1.6 mya, the relation of which with H. habilis is not yet clear. H. rudolfensis refers to a single, incomplete skull from Kenya. Scientists have suggested that this was just another habilis, but this has not been confirmed. H. georgicus, from Georgia, may be an intermediate form between H. habilis and H. erectus, or a sub-species of H. erectus.
Homo ergaster and Homo erectus:
A famous example of Homo erectus is Peking Man; others were found in Asia (notably in Indonesia), Africa, and Europe. Many paleoanthropologists are now using 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. However, this article does not follow that usage.
Homo cepranensis and Homo antecessor:
These are proposed as species that may be intermediate between H. erectus and H. heidelbergensis.
H. antecessor is known from fossils from Spain and England that are 1.2 mya-500,000 years old. H. cepranensis refers to a single skull cap from Italy, estimated to be about 800,000 years old.
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.
Homo rhodesiensis, and the Gawis cranium:
H. rhodesiensis, estimated to be 300,000-125,000 years old. Most current experts believe Rhodesian Man to be within the group of Homo heidelbergensis though other designations such as Archaic Homo sapiens and Homo sapiens rhodesiensis have also been proposed. In February 2006 a fossil, the Gawis cranium, was found which might possibly be a species intermediate between H. erectus and H. sapiens or one of many evolutionary dead ends. The skull from Gawis, Ethiopia, is believed to be 500,000-250,000 years old. Only summary details are known, and no peer reviewed studies have been released by the finding team. Gawis man's facial features suggest its being either an intermediate species or an example of a "Bodo man" female.
H. neanderthalensis lived from about 250,000 to as recent as 30,000 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. 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.
H. sapiens ("sapiens" means 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 cranial 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 (there is little evidence that this speciation occurred elsewhere). Then 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 single-origin theory. However, the current evidence does not preclude multiregional speciation, either. This is a hotly debated area in 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 Toba catastrophe. 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 and include such outward "racial" characteristics 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, lived from about 160,000 years ago (proposed subspecies). It is the oldest known anatomically modern human.
H. floresiensis, which lived about 100,000-12,000 years ago has been nicknamed hobbit for its small size, possibly a result of insular dwarfism. 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. Some scientists presently believe that H. floresiensis was a modern H. sapiens suffering from pathological dwarfism. This hypothesis is supported in part, because the 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.
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 for hunting and consuming meat, which is more energy-rich than plants. Researchers have suggested that early hominids 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. There is some evidence that the australopithecines (4 mya) may have used broken bones as tools, but this is debated.
Stone tools are first attested around 2.6 million years ago, when H. habilis in Eastern Africa used so-called pebble tools, choppers made out of round pebbles that had been split by simple strikes. 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. 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.
Modern humans and the "Great Leap Forward" debate Until about 50,000-40,000 years ago the use of stone tools seems to have progressed stepwise. Each phase (habilis, ergaster, neanderthal) started at a higher level than the previous one; but once that phase started, further development was slow. In other words, these particular Homo species were culturally conservative. After 50,000 BP, however, human culture apparently started to change at a much greater speed. Modern humans started burying their dead carefully, making clothing out of hides, developing sophisticated hunting techniques (such as using trapping pits or driving animals off cliffs), and engaging in cave painting. This speed-up of cultural change seems connected with the arrival of behaviorally modern humans, Homo sapiens. 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, neanderthalensis populations are found with technology similar to other contemporary neanderthalensis populations.
Theoretically, modern human behavior is taken to include four ingredient capabilities: 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). Among concrete examples of modern human behaviour, anthropologists include specialization of tools, use of jewelry 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. Nevertheless, debate continues as to whether a "revolution" led to modern humans ("the big bang of human consciousness"), or whether the evolution was more gradual.
Today, all humans are classified as belonging to the species Homo sapiens sapiens. However, this is not the first species of hominids: the first species of genus Homo, Homo habilis, evolved in East Africa at least 2 million years ago, and members of this species populated different parts of Africa in a relatively short time. Homo erectus evolved more than 1.8 million years ago, and by 1.5 million years ago had spread throughout the Old World. Virtually all physical anthropologists agree that Homo sapiens evolved out of Homo erectus. Anthropologists have been divided as to whether Homo sapiens evolved as one interconnected species from H. erectus (called the Multiregional Model, or the Regional Continuity Model), or evolved only in East Africa, and then migrated out of Africa and replaced H. erectus populations throughout the Old World (called the Out of Africa Model or the Complete Replacement Model). Anthropologists continue to debate both possibilities, and the evidence is technically ambiguous as to which model is correct, although most anthropologists currently favor the Out of Africa model.
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