Human evolution 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 species of Homo evolved, including Homo erectus, which spread to Asia, and Homo neanderthalensis, which spread to Europe. Homo sapiens evolved between 400,000 and 250,000 years ago. Most scientists favor the view that modern humans evolved in Africa and spread across the globe, replacing populations of H. erectus and Neanderthals. Others view modern humans as having evolved as a single, widespread population.
Starting with habilis, humans have used stone tools of increasing sophistication. Starting about 50,000 years ago, human technology and culture began to change more rapidly.
History of paleoanthropologyPaleoanthropology is the study of ancient humans based on fossil evidence, tools, and other signs of human habitation. The modern field of paleoanthropology began in the 19th century with the discovery of "Neanderthal man". The eponymous skeleton was found in 1856, but 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.
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 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 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 (7–6 mya) and Orrorin tugenensis (6 mya), followed by:
- 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
Genus HomoThe word homo is Latin for "human", chosen originally by Carolus Linnaeus in his classification system. It is often translated as "man", although this can lead to confusion, given that the English word "man" can be generic like homo, but can also specifically refer to males. Latin for "man" in the gender-specific sense is vir (pronounced weer), cognate with "virile" and "werewolf". The word "human" is from humanus, the adjectival form of homo.
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.
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. Some scientists have proposed moving this species out of Homo and into Australopithecus.
Homo rudolfensis and Homo georgicus
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.
Homo ergaster and Homo erectus
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.
seealso Control of fire by early humans
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 ergaster.
Homo cepranensis and Homo antecessor
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.
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 mitochondrial DNA and Y-chromosomal DNA sequencing indicates that little or no gene flow occurred between H. neanderthalensis and H. sapiens, and, therefore, the two were separate species. 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.
Homo sapiensH. 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.
Homo floresiensisH. 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.
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. 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 began to create novelty in existing technologies. Artifacts such as fish hooks, buttons and bone needles begin to 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, or 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. Debate continues whether there was indeed a "revolution" leading to modern humans ("the big bang of human consciousness"), or a more gradual evolution.
Models of human evolutionToday, 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.
Multiregional modelAdvocates of the Multiregional model, primarily Milford Wolpoff and his associates, have argued that the simultaneous evolution of H. sapiens in different parts of Europe and Asia would have been possible if there was a degree of gene flow between archaic populations. Similarities of morphological features between archaic European and Chinese populations and modern H. sapiens from the same regions, Wolpoff argues, support a regional continuity only possible within the Multiregional model. Wolpoff and others further argue that this model is consistent with clinal patterns of phenotypic variation (Wolpoff 1993).
Out of Africa modelseealso Recent single origin hypothesis
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.
Lieberman and Jackson have related the multiregional theory to race with the following statement:
Lieberman and Jackson have related the Out of Africa theory to race with the following comment:
Lieberman and Jackson have 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.
Notable human evolution researchers
- James Burnett, Lord Monboddo, a British judge most famous today as a founder of modern comparative historical linguistics
- Charles Darwin, a British naturalist who documented considerable evidence that species originate through evolutionary change
- Richard Dawkins, a British ethologist, evolutionary biologist who has promoted a gene-centered view of evolution
- J. B. S. Haldane, a British geneticist and evolutionary biologist
- William D. Hamilton, a British Evolutionary Biologist who expounded a rigorous genetic basis for kin selection, and on the evolution of HIV and other human diseases.
- Sir Alister Hardy, a British zoologist, who first hypothesised the aquatic ape theory of human evolution
- Henry McHenry, an American anthropologist who specializes in studies of human evolution, the origins of bipedality, and paleoanthropology
- Louis Leakey, an African archaeologist and naturalist whose work was important in establishing human evolutionary development in Africa
- Richard Leakey, an African paleontologist and archaeologist, son of Louis Leakey
- Svante Pääbo, a Swedish biologist specializing in evolutionary genetics
- Jeffrey H. Schwartz, an American physical anthropologist and professor of biological anthropology
- Chris Stringer, anthropologist, leading proponent of the recent single origin hypothesis
- Alan Templeton, geneticist and statistician, proponent of the multiregional hypothesis
- Erik Trinkaus, a prominent American paleoanthropologist and expert on Neanderthal biology and human evolution
- Milford H. Wolpoff, an American paleoanthropologist who is the leading proponent of the multiregional evolution hypothesis
This list is in chronological order by genus.
- The validity of evolution and the origins of humanity have often been a subject of great political and religious controversy within the non-scientific community (see Creation-evolution controversy and Hybrid-origin).
- The classification of humans and their relatives has changed considerably over time (see History of hominoid taxonomy).
- Speculation about the future evolution of humans is often explored in science fiction as continued speciation of humans as they fill various ecological niches (see Adaptive radiation and Co-evolution), as well as deliberate self-modification (see Participant evolution).
- Currently, scientists have estimated that humans branched off from their common ancestor with chimpanzees about 5–7 mya.
- Flinn, M. V., Geary, D. C., & Ward, C. V. (2005). Ecological dominance, social competition, and coalitionary arms races: Why humans evolved extraordinary intelligence. Evolution and Human Behavior, 26, 10-46.
- Rozsa L 2008. The rise of non-adaptive intelligence in humans under pathogen pressure. Medical Hypotheses, 70, 685-690.
- DNA Shows Neandertals Were Not Our Ancestors
- —two ancestral ape chromosomes fused to give rise to human chromosome 2.
- BBC: Finds test human origins theory. 2007-08-08 Homo habilis and Homo erectus are sister species that overlapped in time.
- Aquatic ape hypothesis
- Sociocultural evolution
- Dual inheritance theory
- Evolutionary anthropology
- Evolutionary medicine
- Evolutionary neuroscience
- Evolutionary psychology
- History of Earth
- Hominid intelligence
- Human behavioral ecology
- Human vestigiality
- Hunting hypothesis
- Mitochondrial Eve ("African Eve" theory)
- Most recent common ancestor
- Multi-regional origin
- Physical anthropology
- Single origin hypothesis
- Timeline of human evolution
- Y-chromosomal Adam
protohuman in Arabic: تطور الإنسان
protohuman in Bosnian: Ljudska evolucija
protohuman in Bulgarian: Еволюция на човека
protohuman in Catalan: Procés d'hominització
protohuman in Czech: Vývoj člověka
protohuman in Danish: Menneskets udvikling
protohuman in German: Hominisation
protohuman in Estonian: Inimese evolutsioon
protohuman in Modern Greek (1453-): Ανθρώπινη εξέλιξη
protohuman in Spanish: Evolución humana
protohuman in Basque: Giza eboluzioa
protohuman in Persian: تکامل بشر
protohuman in French: Évolution humaine
protohuman in Korean: 인류의 진화
protohuman in Hindi: मानव का विकास
protohuman in Croatian: Evolucija čovjeka
protohuman in Ido: Evoluciono di homo
protohuman in Icelandic: Þróun mannsins
protohuman in Italian: Evoluzione umana
protohuman in Kannada: ಮಾನವನ ವಿಕಾಸ
protohuman in Latin: Evolutio Hominis
protohuman in Lithuanian: Žmogaus evoliucija
protohuman in Hungarian: Az ember evolúciója
protohuman in Dutch: Evolutie van de mens
protohuman in Norwegian: Menneskets evolusjon
protohuman in Polish: Antropogeneza
protohuman in Portuguese: Evolução humana
protohuman in Romanian: Antropogeneză
protohuman in Russian: Антропогенез
protohuman in Sicilian: Evuluzzioni umana
protohuman in Simple English: Human evolution
protohuman in Slovenian: Nastanek in razvoj človeka
protohuman in Serbian: Развој човека
protohuman in Serbo-Croatian: Ljudska evolucija
protohuman in Finnish: Ihmisen evoluutio
protohuman in Swedish: Människans utveckling
protohuman in Thai: วิวัฒนาการของมนุษย์
protohuman in Turkish: İnsan evrimi
protohuman in Ukrainian: Антропогенез
protohuman in Chinese: 人類演化
Bronze Age man, Hominidae, Iron Age man, Stone Age man, aboriginal, aborigine, ancestral, ancient, antediluvian, antepatriarchal, anthropoid, ape-man, atavistic, autochthon, autochthonous, bushman, cave dweller, caveman, fossil man, hominid, humanoid, man of old, missing link, patriarchal, preadamite, preglacial, prehistoric, prehistoric man, prehuman, primate, prime, primeval, primitive, primogenial, primoprimitive, primordial, pristine, protohistoric, troglodyte