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. The dominant view among scientists is the recent African origin of modern humans (RAO) that H. sapiens evolved in Africa and spread across the globe, replacing populations of H. erectus and H. neanderthalensis. Scientists supporting the alternative hypothesis on the multiregional origin of modern humans (MTO) view modern humans as having evolved as a single, widespread population from existing Homo species, particularly H. erectus. The fossil evidence 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.
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.
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.
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. Most paleontologists consider that primates share a common ancestor with the bats, another extremely ancient lineage, and that this ancestor probably lived during the late Cretaceous, together with the last dinosaurs. The oldest known primates come from North America, but they were widespread in Eurasia and Africa as well, 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. 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—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 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 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 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:
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.
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 human (Homo) physical and behavioral evolution.
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 to the bipedalism 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.
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. H. erectus lived from about 1.8 mya to 70,000 years ago. Often the early phase, from 1.8 to 1.25 mya, 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 mya, in Africa, Asia, and Europe, presumably, some populations of Homo habilis 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. 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 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.
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.
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. In 1997, Dr. Mark Stoneking, then an associate professor of anthropology at Pennsylvania State University, 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. However, supporters of the multiregional hypothesis point to recent studies indicating non-African nuclear DNA heritage dating to one mya, although the reliability of these studies has been questioned.
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 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, lived from about 160,000 years ago (proposed subspecies). It is the oldest known anatomically modern human.
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.
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 .
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 would eventually replace existing hominid species in Europe and Asia. The Out of Africa Model has gained support by recent research using mitochondrial DNA (mtDNA). After analysing genealogy trees constructed using 133 types of mtDNA, they concluded that all were descended from a woman from Africa, dubbed Mitochondrial Eve.
There are differing theories on whether there was a single exodus, or several (a Multiple Dispersal Model). A Multiple Dispersal Model involves the Southern Dispersal theory, 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.. The multiple dispersals models 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. The study further indicates that there was most likely only one single migration out of Africa that gave rise to all Non-African populations.
The multiregional evolution model describes an evolutionary trend throughout the Old World toward modern Homo sapiens, with little population migration and no population replacement, whereas the 'Out of Africa' model calls for the evolution of Homo sapiens in one location only, followed by extensive population migration across the Old World, resulting in the replacement of existing premodern populations.
The Multiregional model suggests that the fossil record should show the same regional differences in anatomical characteristics that are currently visible today. Racial differences would be deep-rooted, going back as much as two million years. In the Out of Africa model, however, the fossil record would not be expected to show continuity of anatomical characteristics over time; on the contrary, the regional characteristics of older fossils would likely be replaced globally by more modern African ones. Racial differences would be shallow-rooted, having evolved over a relatively short period of time.
Implications for the concept of race
In a 1995 article, Leonard Lieberman and Fatimah Jackson point out that the concepts of cline, population, and ethnicity, as well as humanitarian and political concerns, have led many scientists away from the notion of race; nevertheless, a recent survey showed that physical anthropologists were evenly divided as to whether race is a valid biological concept. They note that among physical anthropologists the vast majority of opposition to the race concept comes from population geneticists, so any new support for a biological concept of race would likely come from another source, namely, the study of human evolution. They therefore ask what, if any, implications the current models of human evolution may have for any biological conception of race.
The major implication for race in the multiregional evolution continuity model involves the time depth of a million or more years in which race differentiation might evolve in diverse ecological regions [...]. This must be balanced against the degree of gene flow and the transregional operation of natural selection on encephalization due to development of tools and, more broadly, culture.
Lieberman and Jackson identified these implications of the Out of Africa model on the race concept:
The shallow time dimension minimizes the degree to which racial differences could have evolved [...]. [T]he mitochondrial DNA model presents a view ... minimizing race differences and avoiding racist implications. However, the model, as interpreted by Wainscoat et al. (1989:34), does describe "a major division of human populations into an African and a Eurasian group." This conclusion could best be used to emphasize the degree of biological differences, and thereby provide support for the race concept.
Lieberman and Jackson argued that while advocates of both the Multiregional model and the Out of Africa model use the word race and make racial assumptions, none define the term. They conclude, "Each model has implications that both magnify and minimize the differences between races. Yet each model seems to take race and races as a conceptual reality. The net result is that those anthropologists who prefer to view races as a reality are encouraged to do so." However, Lieberman and Jackson conclude that students of human evolution would be better off avoiding the word race, instead describing genetic differences in terms of populations and clinal gradations.
This list is in chronological order by genus.