|Part of a series on|
Human evolution is the evolutionary process leading to the appearance of anatomically modern humans. The topic typically focuses on the evolutionary history of the primates—in particular the genus Homo, and the emergence of Homo sapiens as a distinct species of the hominids (or "great apes")—rather than studying the earlier history that led to the primates. The study of human evolution involves many scientific disciplines, including physical anthropology, primatology, archaeology, paleontology, ethology, linguistics, evolutionary psychology, embryology and genetics.
Genetic studies show that primates diverged from other mammals about , in the Late Cretaceous period, and the earliest fossils appear in the Paleocene, around . Within the Hominoidea (apes) superfamily, the Hominidae family diverged from the Hylobatidae (gibbon) family some 15–20 million years ago; African great apes (subfamily Homininae) diverged from orangutans (Ponginae) about ; the Hominini tribe (humans, Australopithecines and other extinct biped genera, and chimpanzees) parted from the Gorillini tribe (gorillas) about ; and, in turn, the subtribes Hominina (humans and biped ancestors) and Panina (chimps) separated about .
The basic adaptation of the hominin line is bipedalism. The earliest bipedal hominin is considered to be either Sahelanthropus or Orrorin; alternatively, either Sahelanthropus or Orrorin may instead be the last shared ancestor between chimps and humans. Ardipithecus, a full biped, arose somewhat later, and the early bipeds eventually evolved into the australopithecines, and later into the genus Homo.
The earliest documented representative of the genus Homo is Homo habilis, which evolved around , and is arguably the earliest species for which there is positive evidence of the use of stone tools. The brains of these early hominins were about the same size as that of a chimpanzee, although it has been suggested that this was the time in which the human SRGAP2 gene doubled, producing a more rapid wiring of the frontal cortex. During the next million years a process of rapid encephalization occurred, and with the arrival of Homo erectus and Homo ergaster in the fossil record, cranial capacity had doubled to 850 cm3. (Such an increase in human brain size is equivalent to each generation having 125,000 more neurons than their parents.) It is believed that Homo erectus and Homo ergaster were the first to use fire and complex tools, and were the first of the hominin line to leave Africa, spreading throughout Africa, Asia, and Europe between .
According to the recent African origin of modern humans theory, modern humans evolved in Africa possibly from Homo heidelbergensis, Homo rhodesiensis or Homo antecessor and migrated out of the continent some 50,000 to 100,000 years ago, gradually replacing local populations of Homo erectus, Denisova hominins, Homo floresiensis and Homo neanderthalensis. Archaic Homo sapiens, the forerunner of anatomically modern humans, evolved in the Middle Paleolithic between 400,000 and 250,000 years ago. Recent DNA evidence suggests that several haplotypes of Neanderthal origin are present among all non-African populations, and Neanderthals and other hominins, such as Denisovans, may have contributed up to 6% of their genome to present-day humans, suggestive of a limited inter-breeding between these species. The transition to behavioral modernity with the development of symbolic culture, language, and specialized lithic technology happened around 50,000 years ago according to many anthropologists although some suggest a gradual change in behavior over a longer time span.
- 1 History of study
- 2 Anatomical changes
- 3 Evidence
- 4 Before Homo
- 5 Genus Homo
- 6 Use of tools
- 7 Transition to behavioral modernity
- 8 Recent and current human evolution
- 9 Species list
- 10 See also
- 11 References
- 12 Bibliography
- 13 Further reading
- 14 External links
History of study
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". Linnaeus and other scientists of his time also considered the great apes to be the closest relatives of humans based on morphological and anatomical similarities.
The possibility of linking humans with earlier apes by descent became clear only after 1859 with the publication of Charles Darwin's On the Origin of Species, in which he 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 Henry 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 initially agree that the origin of the mental capacities and the moral sensibilities of humans could be explained by natural selection, though this later changed. Darwin applied the theory of evolution and sexual selection to humans when he published The Descent of Man in 1871.
A major problem at that time was the lack of fossil intermediaries. Neanderthal remains were discovered in a limestone quarry in 1856, three years before the publication of On the Origin of Species, and Neanderthal fossils had been discovered in Gibraltar even earlier, but it was originally claimed that these were human remains of a creature suffering some kind of illness. Despite the 1891 discovery by Eugène Dubois of what is now called Homo erectus at Trinil, Java, it was only in the 1920s when such fossils were discovered in Africa, that intermediate species began to accumulate. In 1925, Raymond Dart described Australopithecus africanus. The type specimen was the Taung Child, an australopithecine infant which was discovered in a cave. The child's remains were a remarkably well-preserved tiny skull and an endocranial cast of the 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 (the hole in the skull where the spine enters) was evidence of bipedal locomotion. All of these traits convinced Dart that the Taung Child was a bipedal human ancestor, a transitional form between apes and humans.
The East African fossils—and Homo naledi in South Africa
During the 1960s and 1970s, hundreds of fossils were found in East Africa in the regions of the Olduvai Gorge and Lake Turkana. The driving force of these searches was the Leakey family, with Louis Leakey and his wife Mary Leakey, and later their son Richard and daughter-in-law Meave—all successful and world-renowned fossil hunters and palaeoanthropologists. From the fossil beds of Olduvai and Lake Turkana they amassed specimens of the early hominins: the australopithecines and Homo species, and even Homo erectus.
These finds cemented Africa as the cradle of humankind. In the late 1970s and the 1980s, Ethiopia emerged as the new hot spot of palaeoanthropology after "Lucy", the most complete fossil member of the species Australopithecus afarensis, was found in 1974 by Donald Johanson near Hadar in the desertic Afar Triangle region of northern Ethiopia. Although the specimen had a small brain, the pelvis and leg bones were almost identical in function to those of modern humans, showing with certainty that these hominins had walked erect. Lucy was classified as a new species, Australopithecus afarensis, which is thought to be more closely related to the genus Homo as a direct ancestor, or as a close relative of an unknown ancestor, than any other known hominid or hominin from this early time range; see terms "hominid" and "hominin". (The specimen was nicknamed "Lucy" after the Beatles' song "Lucy in the Sky with Diamonds", which was played loudly and repeatedly in the camp during the excavations.). The Afar Triangle area would later yield discovery of many more hominin fossils, particularly those uncovered or described by teams headed by Tim D. White in the 1990s, including Ardipithecus ramidus and Ardipithecus kadabba.
In 2013, fossil skeletons of Homo naledi, an extinct species of hominin assigned (provisionally) to the genus Homo, were found in the Rising Star Cave system, a site in South Africa's Cradle of Humankind region in Gauteng province near Johannesburg. As of September 2015[update], fossils of at least fifteen individuals, amounting to 1550 specimens, have been excavated from the cave. The species is characterized by a body mass and stature similar to small-bodied human populations, a smaller endocranial volume similar to Australopithecus, and a cranial morphology (skull shape) similar to early Homo species. The skeletal anatomy combines primitive features known from australopithecines with features known from early hominins. The individuals show signs of having been deliberately disposed of within the cave near the time of death. The fossils have not yet been dated.
The genetic revolution
The genetic revolution in studies of human evolution started when Vincent Sarich and Allan Wilson measured the strength of immunological cross-reactions of blood serum albumin between pairs of creatures, including humans and African apes (chimpanzees and gorillas). The strength of the reaction could be expressed numerically as an immunological distance, which was in turn proportional to the number of amino acid differences between homologous proteins in different species. By constructing a calibration curve of the ID of species' pairs with known divergence times in the fossil record, the data could be used as a molecular clock to estimate the times of divergence of pairs with poorer or unknown fossil records.
In their seminal 1967 paper in Science, Sarich and Wilson estimated the divergence time of humans and apes as four to five million years ago, at a time when standard interpretations of the fossil record gave this divergence as at least 10 to as much as 30 million years. Subsequent fossil discoveries, notably "Lucy", and reinterpretation of older fossil materials, notably Ramapithecus, showed the younger estimates to be correct and validated the albumin method.
Progress in DNA sequencing, specifically mitochondrial DNA (mtDNA) and then Y-chromosome DNA (Y-DNA) advanced the understanding of human origins. Application of the molecular clock principle revolutionized the study of molecular evolution.
On the basis of a separation from the orangutan between 10 and 20 million years ago, earlier studies of the molecular clock suggested that there were about 76 mutations per generation that were not inherited by human children from their parents; this evidence supported the divergence time between hominins and chimps noted above. However, a 2012 study in Iceland of 78 children and their parents suggests a mutation rate of only 36 mutations per generation; this datum extends the separation between humans and chimps to an earlier period greater than 7 million years ago (Ma). Additional research with 226 offspring of wild chimp populations in 8 locations suggests that chimps reproduce at age 26.5 years, on average; which suggests the human divergence from chimps occurred between 7 to 13 million years ago. And these data suggest that Ardipithecus (4.5 Ma), Orrorin (6 Ma) and Sahelanthropus (7 Ma) all may be on the hominin lineage, and even that the separation may have occurred outside the East African Rift region.
Furthermore, analysis of the two species' genes in 2006 provides evidence that after human ancestors had started to diverge from chimpanzees, interspecies mating between "proto-human" and "proto-chimps" nonetheless occurred regularly enough to change certain genes in the new gene pool:
- A new comparison of the human and chimp genomes suggests that after the two lineages separated, they may have begun interbreeding... A principal finding is that the X chromosomes of humans and chimps appear to have diverged about 1.2 million years more recently than the other chromosomes.
The research suggests:
- There were in fact two splits between the human and chimp lineages, with the first being followed by interbreeding between the two populations and then a second split. The suggestion of a hybridization has startled paleoanthropologists, who nonetheless are treating the new genetic data seriously.
The quest for the earliest hominin
In the 1990s, several teams of paleoanthropologists were working throughout Africa looking for evidence of the earliest divergence of the hominin lineage from the great apes. In 1994, Meave Leakey discovered Australopithecus anamensis. The find was overshadowed by Tim D. White's 1995 discovery of Ardipithecus ramidus, which pushed back the fossil record to .
In 2000, Martin Pickford and Brigitte Senut discovered, in the Tugen Hills of Kenya, a 6-million-year-old bipedal hominin which they named Orrorin tugenensis. And in 2001, a team led by Michel Brunet discovered the skull of Sahelanthropus tchadensis which was dated as , and which Brunet argued was a bipedal, and therefore a hominid—that is, a hominin (cf Hominidae; terms "hominids" and hominins).
Anthropologists in the 1980s were divided regarding some details of reproductive barriers and migratory dispersals of the Homo genus. Subsequently, genetics has been used to investigate and resolve these issues. According to the Sahara pump theory evidence suggests that genus Homo have migrated out of Africa at least three times (e.g. Homo erectus, Homo heidelbergensis and Homo sapiens).
The "out of Africa" model proposed that modern H. sapiens speciated in Africa recently (that is, approximately 200,000 years ago) and the subsequent migration through Eurasia resulted in nearly complete replacement of other Homo species. This model has been developed by Chris B. Stringer and Peter Andrews. In contrast, the multiregional hypothesis proposed that Homo genus contained only a single interconnected population as it does today (not separate species), and that its evolution took place worldwide continuously over the last couple million years. This model was proposed in 1988 by Milford H. Wolpoff.
Sequencing mtDNA and Y-DNA sampled from a wide range of indigenous populations revealed ancestral information relating to both male and female genetic heritage. Aligned in genetic tree differences were interpreted as supportive of a recent single origin. Analyses have shown a greater diversity of DNA patterns throughout Africa, consistent with the idea that Africa is the ancestral home of mitochondrial Eve and Y-chromosomal Adam.
"Out of Africa" has gained support from research using female mitochondrial DNA and the male Y chromosome. After analysing genealogy trees constructed using 133 types of mtDNA, researchers concluded that all were descended from a female African progenitor, dubbed Mitochondrial Eve. "Out of Africa" is also supported by the fact that mitochondrial genetic diversity is highest among African populations.
A broad study of African genetic diversity, headed by Sarah Tishkoff, found the San people had 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. The fossil evidence was insufficient for Richard Leakey to resolve this debate. Studies of haplogroups in Y-chromosomal DNA and mitochondrial DNA have largely supported a recent African origin. Evidence from autosomal DNA also predominantly supports a Recent African origin. However, evidence for archaic admixture in modern humans had been suggested by some studies.
Recent sequencing of Neanderthal and Denisovan genomes shows that some admixture occurred. Modern humans outside Africa have 2–4% Neanderthal alleles in their genome, and some Melanesians have an additional 4–6% of Denisovan alleles. These new results do not contradict the "out of Africa" model, except in its strictest interpretation. After recovery from a genetic bottleneck that might be due to the Toba supervolcano catastrophe, a fairly small group left Africa and briefly interbred with Neanderthals, probably in the middle-east or even North Africa before their departure. Their still predominantly African descendants spread to populate the world. A fraction in turn interbred with Denisovans, probably in south-east Asia, before populating Melanesia. HLA haplotypes of Neanderthal and Denisova origin have been identified in modern Eurasian and Oceanian populations.
There are still differing theories on whether there was a single exodus from Africa or several. A multiple dispersal model involves the Southern Dispersal theory, 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 may have dispersed across the Sinai Peninsula into Asia, resulting in the bulk of human population for Eurasia. This second group possibly 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 each glacial maximum. 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.
Human evolution is characterized by a number of morphological, developmental, physiological, and behavioral changes that have taken place since the split between the last common ancestor of humans and chimpanzees. The most significant of these adaptations are bipedalism, increased brain size, lengthened ontogeny (gestation and infancy), and decreased sexual dimorphism. The relationship between these changes is the subject of ongoing debate.[page needed] Other significant morphological changes included the evolution of a power and precision grip, a change first occurring in H. erectus.
Bipedalism is the basic adaptation of the hominin and is considered the main cause behind a suite of skeletal changes shared by all bipedal hominins. The earliest hominin, of presumably primitive bipedalism, is considered to be either Sahelanthropus or Orrorin, both of which arose some 6 to 7 million years ago. The non-bipedal knuckle-walkers, the gorilla and chimpanzee, diverged from the hominin line over a period covering the same time, so either of Sahelanthropus or Orrorin may be our last shared ancestor. Ardipithecus, a full biped, arose somewhat later.
The early bipeds eventually evolved into the australopithecines and later the genus Homo. There are several theories of the adaptation value of bipedalism. It is possible that bipedalism was favored because it freed the hands for reaching and carrying food, saved energy during locomotion, enabled long distance running and hunting, provided an enhanced field of vision, and helped avoid hyperthermia by reducing the surface area exposed to direct sun; features all advantageous for thriving in the new savanna environment versus the previous forest habitat. A new study provides support for the hypothesis that walking on two legs, or bipedalism, evolved because it used less energy than quadrupedal knuckle-walking.
Anatomically, the evolution of bipedalism has been accompanied by a large number of skeletal changes, not just to the legs and pelvis, but also to the vertebral column, feet and ankles, and skull. The femur evolved into a slightly more angular position to move the center of gravity toward the geometric center of the body. The knee and ankle joints became increasingly robust to better support increased weight. To support the increased weight on each vertebra in the upright position, the human vertebral column became S-shaped and the lumbar vertebrae became shorter and wider. In the feet the big toe moved into alignment with the other toes to help in forward locomotion. The arms and forearms shortened relative to the legs making it easier to run. The foramen magnum migrated under the skull and more anterior.
The most significant changes occurred in the pelvic region, where the long downward facing iliac blade was shortened and widened as a requirement for keeping the center of gravity stable while walking; bipedal hominids have a shorter but broader, bowl-like pelvis due to this. A drawback is that the birth canal of bipedal apes is smaller than in knuckle-walking apes, though there has been a widening of it in comparison to that of australopithecine and modern humans, permitting the passage of newborns due to the increase in cranial size but this is limited to the upper portion, since further increase can hinder normal bipedal movement.
The shortening of the pelvis and smaller birth canal evolved as a requirement for bipedalism and had significant effects on the process of human birth which is much more difficult in modern humans than in other primates. During human birth, because of the variation in size of the pelvic region, the fetal head must be in a transverse position (compared to the mother) during entry into the birth canal and rotate about 90 degrees upon exit. The smaller birth canal became an limiting factor to brain size increases in early humans and prompted a shorter gestation period leading to the relative immaturity of human offspring, who are unable to walk much before 12 months and have greater neoteny, compared to other primates, who are mobile at a much earlier age. The increased brain growth after birth and the increased dependency of children on mothers had a big effect upon the female reproductive cycle, and the more frequent appearance of monogamous relationships in humans when compared with other hominids. Delayed human sexual maturity also led to the evolution of menopause with one explanation providing that elderly women could better pass on their genes by taking care of their daughter's offspring, as compared to having more of their own.
The human species developed a much larger brain than that of other primates—typically 1,330 cm3 in modern humans, over twice the size of that of a chimpanzee or gorilla. The pattern of encephalization started with Homo habilis, which at approximately 600 cm3 had a brain slightly larger than that of chimpanzees, and continued with Homo erectus (800–1,100 cm3), reaching a maximum in Neanderthals with an average size of (1,200–1,900 cm3), larger even than Homo sapiens. The pattern of human postnatal brain growth differs from that of other apes (heterochrony) and allows for extended periods of social learning and language acquisition in juvenile humans. However, the differences between the structure of human brains and those of other apes may be even more significant than differences in size.
The increase in volume over time has affected areas within the brain unequally—the temporal lobes, which contain centers for language processing, have increased disproportionately,and seems to favor a belief that there was evolution after leaving Africa, as has the prefrontal cortex which has been related to complex decision-making and moderating social behavior. Encephalization has been tied to an increasing emphasis on meat in the diet, or with the development of cooking, and it has been proposed that intelligence increased as a response to an increased necessity for solving social problems as human society became more complex. The human brain was able to expand because of the changes in the morphology of smaller mandibles and mandible muscle attachments to the skull into allowing more room for the brain to grow.
The increase in volume of the neocortex also included a rapid increase in size of the cerebellum. Traditionally the cerebellum has been associated with a paleocerebellum and archicerebellum as well as a neocerebellum. Its function has also traditionally been associated with balance, fine motor control but more recently speech and cognition. The great apes including humans and its antecessors had a more pronounced development of the cerebellum relative to the neocortex than other primates. It has been suggested that because of its function of sensory-motor control and assisting in learning complex muscular action sequences, the cerebellum may have underpinned the evolution of human's technological adaptations including the preadaptation of speech.
The reason for this encephalisation is difficult to discern, as the major changes from Homo erectus to Homo heidelbergensis were not associated with major changes in technology. It has been suggested that the changes have been associated with social changes, increased empathic abilities  and increases in size of social groupings
The reduced degree of sexual dimorphism is visible primarily in the reduction of the male canine tooth relative to other ape species (except gibbons) and reduced brow ridges and general robustness of males. Another important physiological change related to sexuality in humans was the evolution of hidden estrus. Humans and bonobos are the only apes in which the female is fertile year round and in which no special signals of fertility are produced by the body (such as genital swelling during estrus).
Nonetheless, humans retain a degree of sexual dimorphism in the distribution of body hair and subcutaneous fat, and in the overall size, males being around 15% larger than females. These changes taken together have been interpreted as a result of an increased emphasis on pair bonding as a possible solution to the requirement for increased parental investment due to the prolonged infancy of offspring.
A number of other changes have also characterized the evolution of humans, among them an increased importance on vision rather than smell; a smaller gut; loss of body hair; evolution of sweat glands; a change in the shape of the dental arcade from being u-shaped to being parabolic; development of a chin (found in Homo sapiens alone); development of styloid processes; and the development of a descended larynx.
The evidence on which scientific accounts of human evolution are based comes from many fields of natural science. The main source of knowledge about the evolutionary process has traditionally been the fossil record, but since the development of genetics beginning in the 1970s, DNA analysis has come to occupy a place of comparable importance. The studies of ontogeny, phylogeny and especially evolutionary developmental biology of both vertebrates and invertebrates offer considerable insight into the evolution of all life, including how humans evolved. The specific study of the origin and life of humans is anthropology, particularly paleoanthropology which focuses on the study of human prehistory.
Evidence from molecular biology
The closest living relatives of humans are bonobos and chimpanzees (both genus Pan) and gorillas (genus Gorilla). With the sequencing of both the human and chimpanzee genome, current estimates of the similarity between their DNA sequences range between 95% and 99%. By using the technique called the molecular clock which estimates the time required for the number of divergent mutations to accumulate between two lineages, the approximate date for the split between lineages can be calculated.
The gibbons (family Hylobatidae) and then orangutans (genus Pongo) were the first groups to split from the line leading to the hominins, including humans—followed by gorillas, and, ultimately, by the chimpanzees (genus Pan). The splitting date between hominin and chimpanzee lineages is placed by some between Late Miocene.Speciation, however, appears to have been unusually drawn-out. Initial divergence occurred sometime between , but ongoing hybridization blurred the separation and delayed complete separation during several millions of years. Patterson (2006) dated the final divergence at ., that is, during the
Genetic evidence has also been employed to resolve the question of whether there was any gene flow between early modern humans and Neanderthals, and to enhance our understanding of the early human migration patterns and splitting dates. By comparing the parts of the genome that are not under natural selection and which therefore accumulate mutations at a fairly steady rate, it is possible to reconstruct a genetic tree incorporating the entire human species since the last shared ancestor.
Each time a certain mutation (Single-nucleotide polymorphism) appears in an individual and is passed on to his or her descendants a haplogroup is formed including all of the descendants of the individual who will also carry that mutation. By comparing mitochondrial DNA which is inherited only from the mother, geneticists have concluded that the last female common ancestor whose genetic marker is found in all modern humans, the so-called mitochondrial Eve, must have lived around 200,000 years ago.
Human evolutionary genetics studies how one human genome differs from the other, the evolutionary past that gave rise to it, and its current effects. Differences between genomes have anthropological, medical and forensic implications and applications. Genetic data can provide important insight into human evolution.
Evidence from the fossil record
There is little fossil evidence for the divergence of the gorilla, chimpanzee and hominin lineages. The earliest fossils that have been proposed as members of the hominin lineage are Sahelanthropus tchadensis dating from , Orrorin tugenensis dating from , and Ardipithecus kadabba dating to . Each of these have been argued to be a bipedal ancestor of later hominins but, in each case, the claims have been contested. It is also possible that one or more of these species are ancestors of another branch of African apes, or that they represent a shared ancestor between hominins and other apes.
The question then of the relationship between these early fossil species and the hominin lineage is still to be resolved. From these early species, the australopithecines arose around robust (also called Paranthropus) and gracile branches, one of which (possibly A. garhi) probably went on to become ancestors of the genus Homo. The australopithecine species that is best represented in the fossil record is Australopithecus afarensis with more than one hundred fossil individuals represented, found from Northern Ethiopia (such as the famous "Lucy"), to Kenya, and South Africa. Fossils of robust australopithecines such as Au. robustus (or alternatively Paranthropus robustus) and Au./P. boisei are particularly abundant in South Africa at sites such as Kromdraai and Swartkrans, and around Lake Turkana in Kenya.and diverged into
The earliest member of the genus Homo is Homo habilis which evolved around  Homo habilis is the first species for which we have positive evidence of the use of stone tools. They developed the Oldowan lithic technology, named after the Olduvai Gorge in which the first specimens were found. Some scientists consider Homo rudolfensis, a larger bodied group of fossils with similar morphology to the original H. habilis fossils, to be a separate species while others consider them to be part of H. habilis—simply representing intraspecies variation, or perhaps even sexual dimorphism. The brains of these early hominins were about the same size as that of a chimpanzee, and their main adaptation was bipedalism as an adaptation to terrestrial living..
During the next million years, a process of encephalization began and, by the arrival (about) of Homo erectus in the fossil record, cranial capacity had doubled. Homo erectus were the first of the hominins to emigrate from Africa, and, from , this species spread through Africa, Asia, and Europe. One population of H. erectus, also sometimes classified as a separate species Homo ergaster, remained in Africa and evolved into Homo sapiens. It is believed that these species, H. erectus and H. ergaster, were the first to use fire and complex tools.
The earliest transitional fossils between H. ergaster/erectus and archaic H. sapiens are from Africa, such as Homo rhodesiensis, but seemingly transitional forms were also found at Dmanisi, Georgia. These descendants of African H. erectus spread through Eurasia from ca. 500,000 years ago evolving into H. antecessor, H. heidelbergensis and H. neanderthalensis. The earliest fossils of anatomically modern humans are from the Middle Paleolithic, about 200,000 years ago such as the Omo remains of Ethiopia; later fossils from Es Skhul cave in Israel and Southern Europe begin around 90,000 years ago ( ).
As modern humans spread out from Africa, they encountered other hominins such as Homo neanderthalensis and the so-called Denisovans, who may have evolved from populations of Homo erectus that had left Africa around . The nature of interaction between early humans and these sister species has been a long-standing source of controversy, the question being whether humans replaced these earlier species or whether they were in fact similar enough to interbreed, in which case these earlier populations may have contributed genetic material to modern humans.
This migration out of Africa is estimated to have begun about 70,000 years BP (Before Present) and modern humans subsequently spread globally, replacing earlier hominins either through competition or hybridization. They inhabited Eurasia and Oceania by 40,000 years BP, and the Americas by at least 14,500 years BP.
Early evolution of primates
Evolutionary history of the primates can be traced back 65 million years. The oldest known primate-like mammal species, the Plesiadapis, came from North America, but they were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and Eocene.
David R. Begun  concluded that early primates flourished in Eurasia and that a lineage leading to the African apes and humans, including to Dryopithecus, migrated south from Europe or Western Asia into Africa. The surviving tropical population of primates—which is seen most completely in the Upper Eocene and lowermost Oligocene fossil beds of the Faiyum depression southwest of Cairo—gave rise to all extant primate species, including the lemurs of Madagascar, lorises of Southeast Asia, galagos or "bush babies" of Africa, and to the anthropoids, which are the Platyrrhines or New World monkeys, the Catarrhines or Old World monkeys, and the great apes, including humans and other hominids.
The earliest known catarrhine is Kamoyapithecus from uppermost Oligocene at Eragaleit in the northern Great Rift Valley in Kenya, dated to 24 million years ago. Its ancestry is thought to be species related to Aegyptopithecus, Propliopithecus, and Parapithecus from the Faiyum, at around 35 million years ago. In 2010, Saadanius was described as a close relative of the last common ancestor of the crown catarrhines, and tentatively dated to 29–28 million years ago, helping to fill an 11-million-year gap in the fossil record.
In the Early Miocene, about 22 million years ago, the many kinds of arboreally adapted primitive catarrhines from East Africa suggest a long history of prior diversification. Fossils at 20 million years ago include fragments attributed to Victoriapithecus, the earliest Old World Monkey. Among the genera thought to be in the ape lineage leading up to 13 million years ago are Proconsul, Rangwapithecus, Dendropithecus, Limnopithecus, Nacholapithecus, Equatorius, Nyanzapithecus, Afropithecus, Heliopithecus, and Kenyapithecus, all from East Africa.
The presence of other generalized non-cercopithecids of Middle Miocene from sites far distant—Otavipithecus from cave deposits in Namibia, and Pierolapithecus and Dryopithecus from France, Spain and Austria—is evidence of a wide diversity of forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the Early and Middle Miocene. The youngest of the Miocene hominoids, Oreopithecus, is from coal beds in Italy that have been dated to 9 million years ago.
Molecular evidence indicates that the lineage of gibbons (family Hylobatidae) diverged from the line of great apes some 18–12 million years ago, and that of orangutans (subfamily Ponginae) diverged from the other great apes at about 12 million years; there are no fossils that clearly document the ancestry of gibbons, which may have originated in a so-far-unknown South East Asian hominoid population, but fossil proto-orangutans may be represented by Sivapithecus from India and Griphopithecus from Turkey, dated to around 10 million years ago.
Divergence of the human clade from other great apes
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, however, of gorillas and chimpanzees is limited; both poor preservation—rain forest soils tend to be acidic and dissolve bone—and sampling bias probably contribute to this problem.
Other hominins probably adapted to the drier environments outside the equatorial belt; and there they encountered antelope, hyenas, dogs, pigs, elephants, horses, and others. The equatorial belt contracted after about 8 million years ago, and there is very little fossil evidence for the split—thought to have occurred around that time—of the hominin lineage from the lineages of gorillas and chimpanzees. The earliest fossils argued by some to belong to the human lineage are Sahelanthropus tchadensis (7 Ma) and Orrorin tugenensis (6 Ma), followed by Ardipithecus (5.5–4.4 Ma), with species Ar. kadabba and Ar. ramidus.
It has been argued in a study of the life history of Ar. ramidus that the species provides evidence for a suite of anatomical and behavioral adaptations in very early hominins unlike any species of extant great ape. This study demonstrated affinities between the skull morphology of Ar. ramidus and that of infant and juvenile chimpanzees, suggesting the species evolved a juvenalised or paedomorphic craniofacial morphology via heterochronic dissociation of growth trajectories. It was also argued that the species provides support for the notion that very early hominins, akin to bonobos (Pan paniscus) the less aggressive species of chimpanzee, may have evolved via the process of self-domestication. Consequently, arguing against the so-called "chimpanzee referential model" the authors suggest it is no longer tenable to use common chimpanzee (Pan troglodytes) social and mating behaviors in models of early hominin social evolution. When commenting on the absence of aggressive canine morphology in Ar. ramidus and the implications this has for the evolution of hominin social psychology, they wrote:
Of course Ar. ramidus differs significantly from bonobos, bonobos having retained a functional canine honing complex. However, the fact that Ar. ramidus shares with bonobos reduced sexual dimorphism, and a more paedomorphic form relative to chimpanzees, suggests that the developmental and social adaptations evident in bonobos may be of assistance in future reconstructions of early hominin social and sexual psychology. In fact the trend towards increased maternal care, female mate selection and self-domestication may have been stronger and more refined in Ar. ramidus than what we see in bonobos.:128
The authors argue that many of the basic human adaptations evolved in the ancient forest and woodland ecosystems of late Miocene and early Pliocene Africa. Consequently, they argue that humans may not represent evolution from a chimpanzee-like ancestor as has traditionally been supposed. This suggests many modern human adaptations represent phylogenetically deep traits and that the behavior and morphology of chimpanzees may have evolved subsequent to the split with the common ancestor they share with humans.
The Australopithecus genus evolved in eastern Africa around 4 million years ago before spreading throughout the continent and eventually becoming extinct 2 million years ago. During this time period various forms of australopiths existed, including Australopithecus anamensis, Au. afarensis, Au. sediba, and Au. africanus. There is still some debate among academics whether certain African hominid species of this time, such as Au. robustus and Au. boisei, constitute members of the same genus; if so, they would be considered to be Au. robust australopiths whilst the others would be considered Au. gracile australopiths. However, if these species do indeed constitute their own genus, then they may be given their own name, the Paranthropus.
- Australopithecus (4–1.8 Ma), with species Au. anamensis, Au. afarensis, Au. africanus, Au. bahrelghazali, Au. garhi, and Au. sediba;
- Kenyanthropus (3–2.7 Ma), with species K. platyops;
- Paranthropus (3–1.2 Ma), with species P. aethiopicus, P. boisei, and P. robustus
A new proposed species Australopithecus deyiremeda is claimed to have been discovered living at the same time period of Au. afarensis. There is debate if Au. deyiremeda is a new species or is Au. afarensis.
Homo sapiens is the only extant species of its genus, Homo. While some (extinct) Homo species might have been ancestors of Homo sapiens, many, perhaps most, were likely "cousins," having speciated away from the ancestral hominin line. There is yet no consensus as to which of these groups should be considered a separate species and which should be a subspecies; this may be due to the dearth of fossils or to the slight differences used to classify species in the Homo genus. The Sahara pump theory (describing an occasionally passable "wet" Sahara desert) provides one possible explanation of the early variation in the genus Homo.
Based on archaeological and paleontological evidence, it has been possible to infer, to some extent, the ancient dietary practices of various Homo species and to study the role of diet in physical and behavioral evolution within Homo.
Some anthropologists and archaeologists subscribe to the Toba catastrophe theory, which posits that the supereruption of Lake Toba on Sumatran island in Indonesia some 70,000 years ago caused global consequences, killing the majority of humans and creating a population bottleneck that affected the genetic inheritance of all humans today.
H. habilis and H. gautengensis
Homo habilis lived from about 2.8 to 1.4 Ma. The species evolved in South and East Africa in the Late Pliocene or Early Pleistocene, 2.5–2 Ma, when it diverged from the australopithecines. Homo habilis had smaller molars and larger brains than the australopithecines, and made tools from stone and perhaps animal bones. One of the first known hominins, it was nicknamed 'handy man' by 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 Homo sapiens.
H. rudolfensis and H. georgicus
These are proposed species names for fossils from about 1.9–1.6 Ma, whose relation to Homo habilis is not yet clear.
- Homo rudolfensis refers to a single, incomplete skull from Kenya. Scientists have suggested that this was another Homo habilis, but this has not been confirmed.
- Homo georgicus, from Georgia, may be an intermediate form between Homo habilis and Homo erectus, or a sub-species of Homo erectus.
H. ergaster and H. erectus
The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in 1891 on the Indonesian island of Java. He originally named the material Pithecanthropus erectus based on its morphology, which he considered to be intermediate between that of humans and apes. Homo 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; however, nearby Homo floresiensis survived it. The early phase of Homo erectus, from 1.8 to 1.25 Ma, is considered by some to be a separate species, Homo ergaster, or as Homo erectus ergaster, a subspecies of Homo erectus.
In Africa in the Early Pleistocene, 1.5–1 Ma, some populations of Homo habilis are thought to have evolved larger brains and to have made more elaborate stone tools; these differences and others are sufficient for anthropologists to classify them as a new species, Homo erectus—in Africa. The evolution of locking knees and the movement of the foramen magnum are thought to be likely drivers of the larger population changes. This species also may have used fire to cook 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 reserve Homo erectus only for those fossils that are found in Asia and meet certain skeletal and dental requirements which differ slightly from H. ergaster.
H. cepranensis and H. 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 dated 1.2 Ma–500 ka.
- 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.
H. rhodesiensis, and the Gawis cranium
- H. rhodesiensis, estimated to be 300,000–125,000 years old. Most current researchers place Rhodesian Man within the group of Homo heidelbergensis, though other designations such as archaic Homo sapiens and Homo sapiens rhodesiensis have 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 the finders have not yet released a peer-reviewed study. Gawis man's facial features suggest its being either an intermediate species or an example of a "Bodo man" female.
Neanderthal and Denisovan
Homo neanderthalensis, alternatively designated as Homo sapiens neanderthalensis, lived in Europe and Asia from 400,000 to about 30,000 years ago. Evidence from sequencing mitochondrial DNA indicated that no significant gene flow occurred between H. neanderthalensis and H. sapiens, and that the two were separate species that shared a common ancestor about 660,000 years ago. However, a sequencing of the Neanderthal genome in 2010 indicated that Neanderthals did indeed interbreed with anatomically modern humans circa 45,000 to 80,000 years ago (at the approximate time that modern humans migrated out from Africa, but before they dispersed into Europe, Asia and elsewhere).
Nearly all modern non-African humans have 1% to 4% of their DNA derived from Neanderthal DNA, and this finding is consistent with recent studies indicating that the divergence of some human alleles dates to one Ma, although the interpretation of these studies has been questioned. Neanderthals and Homo sapiens could have co-existed in Europe for as long as 10,000 years, during which human populations exploded vastly outnumbering Neanderthals, possibly outcompeting them by sheer numerical strength.
In 2008, archaeologists working at the site of Denisova Cave in the Altai Mountains of Siberia uncovered a small bone fragment from the fifth finger of a juvenile member of Denisovans. Artifacts, including a bracelet, excavated in the cave at the same level were carbon dated to around 40,000 BP. As DNA had survived in the fossil fragment due to the cool climate of the Denisova Cave, both mtDNA and nuclear DNA were sequenced.
While the divergence point of the mtDNA was unexpectedly deep in time, the full genomic sequence suggested the Denisovans belonged to the same lineage as Neanderthals, with the two diverging shortly after their line split from the lineage that gave rise to modern humans. Modern humans are known to have overlapped with Neanderthals in Europe for more than 10,000 years, and the discovery raises the possibility that Neanderthals, Denisovans, and modern humans may have co-existed. The existence of this distant branch creates a much more complex picture of humankind during the Late Pleistocene than previously thought. Evidence has also been found that as much as 6% of the DNA of some modern Melanesians derive from Denisovans, indicating limited interbreeding in Southeast Asia.
Alleles thought to have originated in Neanderthals and Denisovans have been identified at several genetic loci in the genomes of modern humans outside of Africa. HLA haplotypes from Denisovans and Neanderthal represent more than half the HLA alleles of modern Eurasians, indicating strong positive selection for these introgressed alleles.
H. floresiensis, which lived from approximately 100,000 to 12,000 years before present, 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 an example of a recent species of the genus Homo that exhibits derived traits not shared with modern humans. In other words, H. floresiensis shares 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 hold that H. floresiensis was a modern H. sapiens with pathological dwarfism. This hypothesis is supported in part, because some modern humans who live on Flores, the Indonesian island where the skeleton was found, are pygmies. This, coupled with pathological dwarfism, could have resulted in a significantly diminutive human. The other major attack on H. floresiensis as a separate species is that it was found with tools only associated with H. sapiens.
The hypothesis of pathological dwarfism, however, fails to explain additional anatomical features that are unlike those of modern humans (diseased or not) but much like those of ancient members of our genus. Aside from cranial features, these features include the form of bones in the wrist, forearm, shoulder, knees, and feet. Additionally, this hypothesis fails to explain the find of multiple examples of individuals with these same characteristics, indicating they were common to a large population, and not limited to one individual.
H. sapiens (the adjective sapiens is Latin for "wise" or "intelligent") have 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 intra-cranial volume 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 (both 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 hypothesis" 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 humans 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. 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 at high altitudes.
H. sapiens idaltu, from Ethiopia, is an extinct sub-species from about 160,000 years ago who is argued to be the direct ancestor of all modern humans.
|Species||Temporal range Mya||Habitat||Adult height||Adult mass||Cranial capacity (cm³)||Fossil record||Discovery / publication of name|
|H. habilis||2.1 – 1.5||Africa||150 cm (4 ft 11 in)||33–55 kg (73–121 lb)||510–660||Many||1960/1964|
|H. erectus||1.9 – 0.07||Africa, Eurasia (Java, China, India, Caucasus)||180 cm (5 ft 11 in)||60 kg (130 lb)||850 (early) – 1,100 (late)||Many||1891/1892|
membership in Homo uncertain
also classified as H. habilis
|1.9 – 0.6||South Africa||100 cm (3 ft 3 in)||3 individuals||2010/2010|
also classified as H. erectus
|1.8 – 1.3||Eastern and Southern Africa||700–850||Many||1975|
also classified as H. heidelbergensis
|1.2 – 0.8||Spain||175 cm (5 ft 9 in)||90 kg (200 lb)||1,000||2 sites||1997|
a single fossil, possibly H. erectus
|0.9 – 0.35||Italy||1,000||1 skull cap||1994/2003|
|H. heidelbergensis||0.6 – 0.35||Europe, Africa, China||180 cm (5 ft 11 in)||90 kg (200 lb)||1,100–1,400||Many||1908|
possibly a subspecies of H. sapiens
|0.35 – 0.04||Europe, Western Asia||170 cm (5 ft 7 in)||55–70 kg (121–154 lb) (heavily built)||1,200–1,900||Many||(1829)/1864|
||2.5||South Africa||150 centimetres (4 ft 11 in) tall||45 kilograms (99 lb)||450||15 individuals||2013/2015|
possibly H. erectus
|0.19 – 0.01||Taiwan||1 individual||pre-2008/2015|
also classified as H. heidelbergensis
|0.3 – 0.12||Zambia||1,300||Very few||1921|
|Worldwide||150 - 190 cm (4 ft 7 in - 6 ft 3 in)||50–100 kg (110–220 lb)||950–1,800||(extant)||—/1758|
|0.10 – 0.012||Indonesia||100 cm (3 ft 3 in)||25 kg (55 lb)||400||7 individuals||2003/2004|
possible H. sapiens subspecies or hybrid
|Red Deer Cave people
possible H. sapiens subspecies or hybrid
Use of tools
The use of tools has been interpreted as a sign of intelligence, and it has been theorized that tool use may have stimulated certain aspects of human evolution, especially the continued expansion of the human brain. 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 13 watts (260 kilocalories per day), a fifth of the body's total energy consumption. 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 hominins were thus under evolutionary pressure to increase their capacity to create and use tools.
Precisely when early humans started to use tools is difficult to determine, because the more primitive these tools are (for example, sharp-edged stones) the more difficult it is to decide whether they are natural objects or human artifacts. 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 genus that dominates the areas of making and using more complex tools. The oldest known tools are the Oldowan stone tools from Ethiopia, 2.5–2.6 million years old. 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 a possibility but does not yet represent solid evidence. The Third metacarpal styloid process enables the hand bone to lock into the wrist bones, allowing for greater amounts of pressure to be applied to the wrist and hand from a grasping thumb and fingers. It allows humans the dexterity and strength to make and use complex tools. This unique anatomical feature separates humans from apes and other nonhuman primates, and is not seen in human fossils older than 1.8 million years.
Bernard Wood noted that Paranthropus co-existed with the early Homo species in the area of the "Oldowan Industrial Complex" over roughly the same span of time. Although there is no direct evidence which identifies Paranthropus as the tool makers, their anatomy lends to indirect evidence of their capabilities in this area. Most paleoanthropologists agree that the early Homo species were indeed responsible for most of the Oldowan tools found. They argue that when most of the Oldowan tools were found in association with human fossils, Homo was always present, but Paranthropus was not.
In 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 which are lacking in chimpanzees. Humans also have thicker metacarpals with broader heads, allowing more precise grasping than the chimpanzee hand can perform. Susman posited that modern anatomy of the human opposable thumb is an evolutionary response to the requirements associated with making and handling tools and that both species were indeed toolmakers.
Stone tools are first attested around 2.6 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, (Late Stone Age), 50,000-10,000 years ago.
Archaeologists working in the Great Rift Valley in Kenya claim to have discovered the oldest known stone tools in the world. Dated to around 3.3 million years ago, the implements are some 700,000 years older than stone tools from Ethiopia that previously held this distinction.
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 the more refined so-called Levallois technique was developed, a series of consecutive strikes, by which scrapers, slicers ("racloirs"), needles, and flattened needles were made. 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.
Transition to behavioral modernity
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 after each phase started, further development was slow. Currently paleoanthropologists are debating whether these Homo species possessed some or many of the cultural and behavioral traits associated with modern humans such as language, complex symbolic thinking, technological creativity etc. It seems that they were culturally conservative maintaining simple technologies and foraging patterns over very long periods.
Around 50,000 BP, modern human culture started to evolve more rapidly. The transition to behavioral modernity has been characterized by some as a Eurasian "Great Leap Forward," or as the "Upper Palaeolithic Revolution," due to the sudden appearance of distinctive signs of modern behavior and big game hunting in the archaeological record. Other scholars consider the transition to have been more gradual, noting that some features had already appeared among archaic African Homo sapiens since 200,000 years ago.
Modern humans started burying their dead, using animal hides to make clothing, hunting with more sophisticated techniques (such as using trapping pits or driving animals off cliffs), and engaging in cave painting. 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, although the Chatelperronian assemblages have been found to be Neanderthal innovations produced as a result of exposure to the Homo sapiens Aurignacian technologies.
Among concrete examples of modern human behavior, anthropologists include specialization of tools, use of jewellery and images (such as cave drawings), organization of living space, rituals (for example, burials with grave gifts), specialized hunting techniques, exploration of less hospitable geographical areas, and barter trade networks. Debate continues as to whether a "revolution" led to modern humans ("the big bang of human consciousness"), or whether the evolution was more gradual.
Recent and current human evolution
Natural selection still affects modern human populations. For example, the population at risk of the severe debilitating disease kuru has significant over-representation of an immune variant of the prion protein gene G127V versus non-immune alleles. The frequency of this genetic variant is due to the survival of immune persons. Other reported trends appear to include lengthening of the human reproductive period and reduction in cholesterol levels, blood glucose and blood pressure in some populations .
It has been argued that human evolution has accelerated since the development of agriculture and civilization some 10,000 years ago, resulting, it is claimed, in substantial genetic differences between different current human populations. The retention of lactase persistence into adulthood is an example of such recent evolution. Recent human evolution seems to have been largely confined to genetic resistance to infectious disease that have appeared in human populations by crossing the species barrier from domesticated animals.
It is a common misconception that humans have stopped evolving and current genetic changes are purely genetic drift. Although selection pressure on some traits has decreased in modern human life (for instance, we are no longer evolving to survive smallpox), humans are still undergoing natural selection for many other traits (for instance, menopause is evolving to occur later).
This list is in chronological order across the page by genus.
- Adaptive evolution in the human genome
- The Ancestor's Tale
- Chimpanzee–human last common ancestor
- Dawn of Humanity (2015 PBS film)
- Dual inheritance theory
- Evolution of hair
- Evolution of human intelligence
- Evolution of morality
- Evolutionary anthropology
- Evolutionary medicine
- Evolutionary neuroscience
- Evolutionary origin of religions
- Evolutionary psychology
- Human behavioral ecology
- Human evolution (origins of society and culture)
- Human skeletal changes due to bipedalism
- Human vestigiality
- List of human evolution fossils
- March of Progress
- Molecular paleontology
- Origin of language
- Origin of speech
- Obstetrical dilemma
- Recent African origin of modern humans
- Sahara pump theory
- Sexual selection in human evolution
- Timeline of human evolution
- Heng, Henry H. Q. (May 2009). "The genome-centric concept: resynthesis of evolutionary theory". BioEssays (Hoboken, NJ: John Wiley & Sons) 31 (5): 512–525. doi:10.1002/bies.200800182. ISSN 0265-9247. PMID 19334004.
- Tyson, Peter (July 1, 2008). "Meet Your Ancestors". NOVA scienceNOW. PBS; WGBH Educational Foundation. Retrieved 2015-04-18.
- Dawkins 2004
- "Find Time of Divergence: Hominidae versus Hylobatidae". TimeTree. Retrieved 2015-04-18.
- Ghosh, Pallab (March 4, 2015). "'First human' discovered in Ethiopia". BBC News (London: BBC). Retrieved 2015-04-19.
- Swisher, Curtis & Lewin 2001
- Stringer 1994, p. 242
- McHenry 2009, p. 265
- "Out of Africa Revisited". Science (This Week in Science) (Washington, D.C.: American Association for the Advancement of Science) 308 (5724): 921. May 13, 2005. doi:10.1126/science.308.5724.921g. ISSN 0036-8075.
- Stringer, Chris (June 12, 2003). "Human evolution: Out of Ethiopia". Nature (London: Nature Publishing Group) 423 (6941): 692–695. doi:10.1038/423692a. ISSN 0028-0836. PMID 12802315.
- Johanson, Donald (May 2001). "Origins of Modern Humans: Multiregional or Out of Africa?". actionbioscience. Washington, D.C.: American Institute of Biological Sciences. Retrieved 2009-11-23.
- Mixon, Bobbie; Ehardt, Carolyn; Hammer, Michael (September 6, 2011). "Evolution's Past Is Modern Human's Present" (Press release). National Science Foundation. Press Release 11-181. Retrieved 2015-04-20.
- O'Neil, Dennis. "Early Modern Homo sapiens". Evolution of Modern Humans: A Survey of the Biological and Cultural Evolution of Archaic and Modern Homo sapiens (Tutorial). San Marcos, CA: Palomar College. Retrieved 2015-04-20.
- "Fossil Reanalysis Pushes Back Origin of Homo sapiens". Scientific American (Stuttgart: Georg von Holtzbrinck Publishing Group). February 17, 2005. ISSN 0036-8733. Retrieved 2015-04-20.
- Reich, David; Green, Richard E.; Kircher, Martin; et al. (December 23, 2010). "Genetic history of an archaic hominin group from Denisova Cave in Siberia". Nature (London: Nature Publishing Group) 468 (7327): 1053–1060. Bibcode:2010Natur.468.1053R. doi:10.1038/nature09710. ISSN 0028-0836. PMID 21179161.
- Noonan, James P. (May 2010). "Neanderthal genomics and the evolution of modern humans". Genome Research (Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press) 20 (5): 547–553. doi:10.1101/gr.076000.108. ISSN 1088-9051. PMC 2860157. PMID 20439435.
- Abi-Rached, Laurent; Jobin, Matthew J.; Kulkarni, Subhash; et al. (October 7, 2011). "The Shaping of Modern Human Immune Systems by Multiregional Admixture with Archaic Humans". Science (Washington, D.C.: American Association for the Advancement of Science) 334 (6052): 89–94. Bibcode:2011Sci...334...89A. doi:10.1126/science.1209202. ISSN 0036-8075. PMC 3677943. PMID 21868630.
- Mellars, Paul (June 20, 2006). "Why did modern human populations disperse from Africa ca. 60,000 years ago? A new model". Proc. Natl. Acad. Sci. U.S.A. (Washington, D.C.: National Academy of Sciences) 103 (25): 9381–9386. Bibcode:2006PNAS..103.9381M.. doi:10.1073/pnas.0510792103. ISSN 0027-8424. PMC 1480416. PMID 16772383.
- Mcbrearty, Sally; Brooks, Alison S. (November 2000). "The revolution that wasn't: a new interpretation of the origin of modern human behavior". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 39 (5): 453–563. doi:10.1006/jhev.2000.0435. ISSN 0047-2484. PMID 11102266.
- American Heritage Dictionaries (editors) 2006
- Darwin 1981
- Montgomery 1988, pp. 95–96
- Dart, Raymond (February 7, 1925). "Australopithecus africanus: The Man-Ape of South Africa" (PDF). Nature (London: Nature Publishing Group) 115 (2884): 195–199. doi:10.1038/115195a0. ISSN 0028-0836. Retrieved 2015-05-11.
- Johanson 1981, pp. 20–22, 184–185
- Cartmill, Matt; Fred H. Smith; Kaye B. Brown (2009). The Human Lineage. Wiley-Blackwell. p. 151. ISBN 978-0-471-21491-5.
- Johanson 1981, p. 22
- Shreeve, Jamie (July 2010). "The Evolutionary Road". National Geographic (Washington, D.C.: National Geographic Society). ISSN 0027-9358. Retrieved 2015-05-28.
- Berger, Lee R. et al. (10 September 2015). "Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa". eLife 4. doi:10.7554/eLife.09560. Retrieved 19 September 2015. Lay summary.
- Shreeve, Jamie (10 September 2015). "This Face Changes the Human Story. But How?". National Geographic News. Retrieved 10 September 2015.
- Berger et al. (2015): "If the fossils prove to be substantially older than 2 million years, H. naledi would be the earliest example of our genus that is more than a single isolated fragment. [...] A date younger than 1 million years ago would demonstrate the coexistence of multiple Homo morphs in Africa, including this small-brained form, into the later periods of human evolution."
- Sarich, V. M.; Wilson, A. C. (1967). "Immunological time scale for hominid evolution". Science 158 (3805): 1200–1203. doi:10.1126/science.158.3805.1200. PMID 4964406.
- M'charek 2005, p. 96
- DeSalle & Tattersall 2008, p. 146
- Trent 2005, p. 6
- Wade, Nicholas (May 18, 2006). "Two Splits Between Human and Chimp Lines Suggested". The New York Times. Retrieved 2015-04-20.
- Behar, Doron M.; Villems, Richard; Soodyall, Himla; et al. (May 9, 2008). "The Dawn of Human Matrilineal Diversity" (PDF). American Journal of Human Genetics (Cambridge, MA: Cell Press on behalf of the American Society of Human Genetics) 82 (5): 1130–1140. doi:10.1016/j.ajhg.2008.04.002. ISSN 0002-9297. PMC 2427203. PMID 18439549. Retrieved 2015-04-20.
- Gonder, Mary Katherine; Mortensen, Holly M.; Reed, Floyd A.; et al. (March 2007). "Whole-mtDNA Genome Sequence Analysis of Ancient African Lineages" (PDF). Molecular Biology and Evolution (Oxford, UK: Oxford University Press on behalf of the Society for Molecular Biology and Evolution) 24 (3): 757–768. doi:10.1093/molbev/msl209. ISSN 0737-4038. PMID 17194802. Retrieved 2015-04-20.
- Tishkoff and Reed (2009)
- Templeton, Alan R. (2005). "Haplotype Trees and Modern Human Origins". American Journal of Physical Anthropology (Hoboken, NJ: John Wiley & Sons for the American Association of Physical Anthropologists) 128 (Supplement 41 (Yearbook of Physical Anthropology)): 33–59. doi:10.1002/ajpa.20351. ISSN 0002-9483. PMID 16369961.
- Owen, James (July 18, 2007). "Modern Humans Came Out of Africa, 'Definitive' Study Says". National Geographic News (Washington, D.C.: National Geographic Society). Retrieved 2011-05-14.
- Stringer, Chris B.; Andrews, Peter (March 11, 1988). "Genetic and fossil evidence for the origin of modern humans". Science (Washington, D.C.: American Association for the Advancement of Science) 239 (4845): 1263–1268. Bibcode:1988Sci...239.1263S. doi:10.1126/science.3125610. ISSN 0036-8075. PMID 3125610.
- Wolpoff, Milford H.; Hawks, John; Caspari, Rachel (May 2000). "Multiregional, Not Multiple Origins" (PDF). American Journal of Physical Anthropology (Hoboken, NJ: John Wiley & Sons for the American Association of Physical Anthropologists) 112 (1): 129–136. doi:10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-K. ISSN 0002-9483. PMID 10766948.
- Wolpoff, Milford H.; Spuhler, James N.; Smith, Fred H.; et al. (August 12, 1988). "Modern Human Origins". Science (Washington, D.C.: American Association for the Advancement of Science) 241 (4867): 772–774. Bibcode:1988Sci...241..772W. doi:10.1126/science.3136545. ISSN 0036-8075. PMID 3136545.
- Webster 2010, p. 53
- Ramachandran et al. 2010, p. 606
- Kutty 2009, p. 40
- Cann, Rebecca L.; Stoneking, Mark; Wilson, Allan C. (January 1, 1987). "Mitochondrial DNA and human evolution". Nature (London: Nature Publishing Group) 325 (6099): 31–36. Bibcode:1987Natur.325...31C. doi:10.1038/325031a0. ISSN 0028-0836. PMID 3025745. Archived from the original on 2010-08-13. Retrieved 2015-04-21.
- Gill, Victoria (May 1, 2009). "Africa's genetic secrets unlocked". BBC News (London: BBC). Retrieved 2011-06-08. The results were published in the online edition of the journal Science.
- Leakey 1994, pp. 87–89
- Jorde, Lynn B.; Bamshad, Michael; Rogers, Alan R. (February 1998). "Using mitochondrial and nuclear DNA markers to reconstruct human evolution". BioEssays (Hoboken, NJ: John Wiley & Sons) 20 (2): 126–136. doi:10.1002/(SICI)1521-1878(199802)20:2<126::AID-BIES5>3.0.CO;2-R. ISSN 0265-9247. PMID 9631658.
- Wall, Jeffrey D.; Lohmueller, Kirk E.; Plagnol, Vincent (August 2009). "Detecting Ancient Admixture and Estimating Demographic Parameters in Multiple Human Populations". Molecular Biology and Evolution (Oxford, UK: Oxford University Press on behalf of the Society for Molecular Biology and Evolution) 26 (8): 1823–1827. doi:10.1093/molbev/msp096. ISSN 0737-4038. PMC 2734152. PMID 19420049.
- Green, Richard E.; Krause, Johannes; Briggs, Adrian W.; et al. (May 7, 2010). "A Draft Sequence of the Neandertal Genome". Science (Washington, D.C.: American Association for the Advancement of Science) 328 (5979): 710–722. Bibcode:2010Sci...328..710G. doi:10.1126/science.1188021. ISSN 0036-8075. PMID 20448178.
- Reich, David; Patterson, Nick; Kircher, Martin; et al. (October 7, 2011). "Denisova Admixture and the First Modern Human Dispersals into Southeast Asia and Oceania". American Journal of Human Genetics (Cambridge, MA: Cell Press on behalf of the American Society of Human Genetics) 89 (4): 516–528. doi:10.1016/j.ajhg.2011.09.005. ISSN 0002-9297. PMC 3188841. PMID 21944045. Retrieved 2015-04-21.
- Lahr, Marta Mirazón; Petraglia, Mike; Stokes, Stephen; et al. "Searching for traces of the Southern Dispersal". Leverhulme Centre for Human Evolutionary Studies. Cambridge, UK: University of Cambridge. Archived from the original on 2012-05-10. Retrieved 2015-04-21.
- Macaulay, Vincent; Hill, Catherine; Achilli, Alessandro; et al. (May 13, 2005). "Single, Rapid Coastal Settlement of Asia Revealed by Analysis of Complete Mitochondrial Genomes". Science (Washington, D.C.: American Association for the Advancement of Science) 308 (5724): 1034–1036. Bibcode:2005Sci...308.1034M. doi:10.1126/science.1109792. ISSN 0036-8075. PMID 15890885.
- Boyd & Silk 2003
- Brues & Snow 1965, pp. 1–39
- Brunet, Michel; Guy, Franck; Pilbeam, David; et al. (July 11, 2002). "A new hominid from the Upper Miocene of Chad, Central Africa". Nature (London: Nature Publishing Group) 418 (6894): 145–151. doi:10.1038/nature00879. ISSN 0028-0836. PMID 12110880.
- Srivastava 2009, p. 87
- Curry 2008, pp. 106–109
- "Study Identifies Energy Efficiency As Reason For Evolution Of Upright Walking". ScienceDaily (Rockville, MD: ScienceDaily, LLC). July 17, 2007. Retrieved 2015-04-09.
- "Study identifies energy efficiency as reason for evolution of upright walking". UANews. Tucson, AZ: The University of Arizona Office of University Communications. July 16, 2007. Retrieved 2015-04-23.
- Sockol, Michael D.; Raichlen, David A.; Pontzer, Herman (July 24, 2007). "Chimpanzee locomotor energetics and the origin of human bipedalism". Proc. Natl. Acad. Sci. U.S.A. (Washington, D.C.: National Academy of Sciences) 104 (30): 12265–12269. doi:10.1073/pnas.0703267104. ISSN 0027-8424. PMC 1941460. PMID 17636134.
- Aiello & Dean 1990
- Kondo 1985
- Strickberger 2000, pp. 475–476
- Trevathan 2011, p. 20
- Wayman, Erin (August 19, 2013). "Killer whales, grandmas and what men want: Evolutionary biologists consider menopause". Science News (Washington, D.C.: Society for Science & the Public). ISSN 0036-8423. Retrieved 2015-04-24.
- Schoenemann, P. Thomas (October 2006). "Evolution of the Size and Functional Areas of the Human Brain". Annual Review of Anthropology (Palo Alto, CA: Annual Reviews) 35: 379–406. doi:10.1146/annurev.anthro.35.081705.123210. ISSN 0084-6570.
- Brown, Graham; Fairfax, Stephanie; Sarao, Nidhi. Tree of Life Web Project: Human Evolution. Link: http://tolweb.org/treehouses/?treehouse_id=3710
- Park, Min S.; Nguyen, Andrew D.; Aryan, Henry E.; et al. (March 2007). "Evolution of the human brain: changing brain size and the fossil record". Neurosurgery (Philadelphia, PA: Lippincott Williams & Wilkins) 60 (3): 555–562. doi:10.1227/01.NEU.0000249284.54137.32. ISSN 0148-396X. PMID 17327801.
- Bruner, Emiliano (December 2007). "Cranial shape and size variation in human evolution: structural and functional perspectives". Child's Nervous System (Heidelberg: Springer-Verlag) 23 (12): 1357–1365. doi:10.1007/s00381-007-0434-2. ISSN 0256-7040. PMID 17680251. CiteSeerX: 10
.1 .1 .391 .288.
- Potts, Richard (October 2012). "Evolution and Environmental Change in Early Human Prehistory". Annual Review of Anthropology (Palo Alto, CA: Annual Reviews) 41: 151–167. doi:10.1146/annurev-anthro-092611-145754. ISSN 0084-6570.
- Leonard, William R.; Snodgrass, J. Josh; Robertson, Marcia L. (August 2007). "Effects of brain evolution on human nutrition and metabolism". Annual Review of Nutrition (Palo Alto, CA: Annual Reviews) 27: 311–327. doi:10.1146/annurev.nutr.27.061406.093659. ISSN 0199-9885. PMID 17439362.
- McBroom, Patricia (June 14, 1999). "Meat-eating was essential for human evolution, says UC Berkeley anthropologist specializing in diet" (Press release). Berkeley, CA: University of California, Berkeley. Retrieved 2015-04-25.
- Mann, Neil (September 2007). "Meat in the human diet: An anthropological perspective". Nutrition & Dietetics (Hoboken, NJ: Wiley-Blackwell) 64 (Supplement s4): S102–S107. doi:10.1111/j.1747-0080.2007.00194.x. ISSN 1747-0080. Retrieved 2012-01-31.
- Zimmer, Carl (August 13, 2015). "For Evolving Brains, a ‘Paleo’ Diet Full of Carbs". New York Times. Retrieved August 14, 2015.
- Organ, Chris; Nunn, Charles L.; Machanda, Zarin; Wrangham, Richard W. (August 30, 2011). "Phylogenetic rate shifts in feeding time during the evolution of Homo". Proc. Natl. Acad. Sci. U.S.A. (Washington, D.C.: National Academy of Sciences) 108 (35): 14555–14559. doi:10.1073/pnas.1107806108. ISSN 0027-8424. PMC 3167533. PMID 21873223.
- Bown & Rose 1987
- Barton, Robert A.; Venditti, Chris (October 20, 2014). "Rapid Evolution of the Cerebellum in Humans and Other Great Apes". Current Biology (Cambridge, MA: Cell Press) 24 (20): 2440–2444. doi:10.1016/j.cub.2014.08.056. ISSN 0960-9822.
- Starowicz-Filip, Anna; Milczarek, Olga; Kwiatkowski, Stanisław; et al. (2013). "Cerebellar cognitive affective syndrome CCAS – a case report" (PDF). Archives of Psychiatry and Psychotherapy (Kraków, Poland: Polish Psychiatric Association) 3: 57–64. OCLC 220954272.
- Feng Yu; Qing-jun Jiang; Xi-yan Sun; et al. (August 22, 2014). "A new case of complete primary cerebellar agenesis: clinical and imaging findings in a living patient". Brain (Oxford, UK: Published by Oxford University Press on behalf of the Guarantors of Brain). doi:10.1093/brain/awu239. ISSN 1460-2156. PMID 25149410.
- Weaver, Anne H. (March 8, 2005). "Reciprocal evolution of the cerebellum and neocortex in fossil humans". Proc. Natl. Acad. Sci. U.S.A. (Washington, D.C.: National Academy of Sciences) 102 (10): 3576–3580. doi:10.1073/pnas.0500692102. ISSN 0027-8424. PMC 553338. PMID 15731345.
- Klein, Stefan (2012) "Evolution of the Nicest"
- Sallet, J.; Mars, R. B.; Noonan, M. P.; Andersson, J. L.; O’Reilly, J. X.; Jbabdi, S.; Croxson, P. L.; Jenkinson, M.; Miller, K. L. (2011-11-04). "Social Network Size Affects Neural Circuits in Macaques". Science 334 (6056): 697–700. doi:10.1126/science.1210027. ISSN 0036-8075. PMID 22053054.
- Dunbar, R. I. M. (1992). "Neocortex size as a constraint on group size in primates". Journal of Human Evolution 22 (6): 469–493.
- Dávid-Barrett, T.; Dunbar, R. I. M. (2013-08-22). "Processing power limits social group size: computational evidence for the cognitive costs of sociality". Proceedings of the Royal Society of London B: Biological Sciences 280 (1765): 20131151. doi:10.1098/rspb.2013.1151. ISSN 0962-8452. PMC 3712454. PMID 23804623.
- Shultz, Susanne; Dunbar, Robin (2010-12-14). "Encephalization is not a universal macroevolutionary phenomenon in mammals but is associated with sociality". Proceedings of the National Academy of Sciences 107 (50): 21582–21586. doi:10.1073/pnas.1005246107. ISSN 0027-8424. PMC 3003036. PMID 21098277.
- Stanford, Allen & Antón 2009
- Wood, Bernard A.; Richmond, Brian G. (July 2000). "Human evolution: taxonomy and paleobiology". Journal of Anatomy (Hoboken, NJ: Wiley-Blackwell on behalf of the Anatomical Society of Great Britain and Ireland) 197 (1): 19–60. doi:10.1046/j.1469-7580.2000.19710019.x. ISSN 1469-7580. PMC 1468107. PMID 10999270.
- Ajit, Varki; Nelson, David L. (October 2007). "Genomic Comparisons of Humans and Chimpanzees" (PDF). Annual Review of Anthropology (Palo Alto, CA: Annual Reviews) 36: 191–209. doi:10.1146/annurev.anthro.36.081406.094339. ISSN 0084-6570. Retrieved 2015-04-26.
Sequence differences from the human genome were confirmed to be ∼1% in areas that can be precisely aligned, representing ∼35 million single base-pair differences. Some 45 million nucleotides of insertions and deletions unique to each lineage were also discovered, making the actual difference between the two genomes ∼4%.
- Sayers, Ken; Raghanti, Mary Ann; Lovejoy, C. Owen (October 2012). "Human Evolution and the Chimpanzee Referential Doctrine". Annual Review of Anthropology (Palo Alto, CA: Annual Reviews) 41: 119–138. doi:10.1146/annurev-anthro-092611-145815. ISSN 0084-6570.
- Ruvolo, Maryellen (October 1997). "Genetic Diversity in Hominoid Primates". Annual Review of Anthropology (Palo Alto, CA: Annual Reviews) 26: 515–540. doi:10.1146/annurev.anthro.26.1.515. ISSN 0084-6570.
- Ruvolo, Maryellen (March 1997). "Molecular Phylogeny of the Hominoids: Inferences from Multiple Independent DNA Sequence Data Sets" (PDF). Molecular Biology and Evolution (Oxford, UK: Oxford University Press on behalf of the Society for Molecular Biology and Evolution) 14 (3): 248–265. doi:10.1093/oxfordjournals.molbev.a025761. ISSN 0737-4038. PMID 9066793.
- Patterson N, Richter DJ, Gnerre S, Lander ES, Reich D (June 2006). "Genetic evidence for complex speciation of humans and chimpanzees". Nature 441 (7097): 1103–8. doi:10.1038/nature04789. PMID 16710306.
- Begun, David R. (October 2010). "Miocene Hominids and the Origins of the African Apes and Humans". Annual Review of Anthropology (Palo Alto, CA: Annual Reviews) 39: 67–84. doi:10.1146/annurev.anthro.012809.105047. ISSN 0084-6570.
- Wood 2009, pp. 17–27
- Mitchell, Alanna (January 30, 2012). "DNA Turning Human Story Into a Tell-All". The New York Times. Retrieved 2012-02-13.
- Wood, Bernard A. (December 1996). "Human evolution". BioEssays (Hoboken, NJ: John Wiley & Sons) 18 (12): 945–954. doi:10.1002/bies.950181204. ISSN 0265-9247. PMID 8976151.
- Maxwell 1984, p. 296
- Rui Zhang; Yin-Qiu Wang; Bing Su (July 2008). "Molecular Evolution of a Primate-Specific microRNA Family" (PDF). Molecular Biology and Evolution (Oxford, UK: Oxford University Press on behalf of the Society for Molecular Biology and Evolution) 25 (7): 1493–1502. doi:10.1093/molbev/msn094. ISSN 0737-4038. PMID 18417486. Retrieved 2015-04-27.
- Willoughby, Pamela R. (2005). "Palaeoanthropology and the Evolutionary Place of Humans in Nature". International Journal of Comparative Psychology (International Society for Comparative Psychology) 18 (1): 60–91. ISSN 0889-3667. Retrieved 2015-04-27.
- Martin 2001, pp. 12032–12038
- Tavaré, Simon; Marshall, Charles R.; Will, Oliver; et al. (April 18, 2002). "Using the fossil record to estimate the age of the last common ancestor of extant primates". Nature (London: Nature Publishing Group) 416 (6882): 726–729. doi:10.1038/416726a. ISSN 0028-0836. PMID 11961552.
- Rose, Kenneth D. (1994). "The earliest primates". Evolutionary Anthropology: Issues, News, and Reviews (Hoboken, NJ: John Wiley & Sons) 3 (5): 159–173. doi:10.1002/evan.1360030505. ISSN 1060-1538.
- Fleagle, John; Gilbert, Chris (2011). Rowe, Noel; Myers, Marc, eds. "Primate Evolution". All The World's Primates. Charlestown, RI: Primate Conservation, Inc. Retrieved 2015-04-27.
- Roach, John (March 3, 2008). "Oldest Primate Fossil in North America Discovered". National Geographic News (Washington, D.C.: National Geographic Society). Retrieved 2015-04-27.
- McMains, Vanessa (December 5, 2011). "Found in Wyoming: New fossils of oldest American primate". The Gazette (Baltimore, MD: Johns Hopkins University). Retrieved 2015-04-27.
- Caldwell, Sara B. (May 19, 2009). "Missing link found, early primate fossil 47 million years old". Digital Journal (Toronto, Canada: digitaljournal.com). Retrieved 2015-04-27.
- Watts, Alex (May 20, 2009). "Scientists Unveil Missing Link In Evolution". Sky News Online (London: BSkyB). Archived from the original on 2011-07-28. Retrieved 2015-04-27.
- Kordos, László; Begun, David R. (January 2001). "Primates from Rudabánya: allocation of specimens to individuals, sex and age categories". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 40 (1): 17–39. doi:10.1006/jhev.2000.0437. ISSN 0047-2484. PMID 11139358.
- Cameron 2004, p. 76
- Wallace 2004, p. 240
- Zalmout, Iyad S.; Sanders, William J.; MacLatchy, Laura M.; et al. (July 15, 2010). "New Oligocene primate from Saudi Arabia and the divergence of apes and Old World monkeys". Nature (London: Nature Publishing Group) 466 (7304): 360–364. Bibcode:2010Natur.466..360Z. doi:10.1038/nature09094. ISSN 0028-0836. PMID 20631798.
- Clark, G.; Henneberg, M. (June 2015). "The life history of Ardipithecus ramidus: a heterochronic model of sexual and social maturation". Anthropological Review 78 (2). Retrieved 2015-07-22.
- Sayers, K.; et al. "Human Evolution and the Chimpanzee refernatial Model". Annual Review of Anthropology 41: 119–138.
- Zimmer, Carl (May 27, 2015). "The Human Family Tree Bristles With New Branches". The New York Times. Retrieved 2015-05-30.
- "Figure 5. Temporal and Geographical Distribution of Hominid Populations Redrawn from Stringer (2003)" (edited from source), in Reed, David L.; Smith, Vincent S.; Hammond, Shaless L.; et al. (November 2004). "Genetic Analysis of Lice Supports Direct Contact between Modern and Archaic Humans". PLOS Biology (San Francisco, CA: PLOS) 2 (11): e340. doi:10.1371/journal.pbio.0020340. ISSN 1545-7885. PMC 521174. PMID 15502871.
- Strait, David S.; Grine, Frederick E.; Moniz, Marc A. (January 1997). "A reappraisal of early hominid phylogeny". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 32 (1): 17–82. doi:10.1006/jhev.1996.0097. ISSN 0047-2484. PMID 9034954.
- Bryson 2004, pp. 522–543
- Walker 2007, pp. 3–10
- Ungar & Teaford 2002
- Bogin 1997, pp. 96–142
- Barnicot, Nigel A. (April–June 2005). "Human nutrition: Evolutionary perspectives". Integrative Physiological & Behavioral Science (New York: Springer US) 40 (2): 114–117. doi:10.1007/BF02734246. ISSN 1932-4502. PMID 17393680.
- "The new batch - 150,000 years ago". BBC - Science & Nature - The evolution of man (London: BBC). Archived from the original on 2006-01-18. Retrieved 2015-04-28.
- Whitehouse, David (June 9, 2003). "When humans faced extinction". BBC News (London: BBC). Retrieved 2007-01-05.
- Wood, Bernard; Collard, Mark (1999). "The changing face of Genus Homo". Evolutionary Anthropology: Issues, News, and Reviews (Hoboken, NJ: John Wiley & Sons) 8 (6): 195–207. doi:10.1002/(SICI)1520-6505(1999)8:6<195::AID-EVAN1>3.0.CO;2-2. ISSN 1060-1538.
- Viegas, Jennifer (May 21, 2010). "Toothy Tree-Swinger May Be Earliest Human". Discovery News (Silver Spring, MD: Discovery Communications, LLC.). Retrieved 2015-04-28.
- Wood, Bernard A. (January 1999). "‘Homo rudolfensis’ Alexeev, 1986—fact or phantom?". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 36 (1): 115–118. doi:10.1006/jhev.1998.0246. ISSN 0047-2484. PMID 9924136.
- Gabounia, Léo; de Lumley, Marie-Antoinette; Vekua, Abesalom; et al. (September 2002). "Découverte d’un nouvel hominidé à Dmanissi (Transcaucasie, Géorgie)" [Discovery of a new hominid at Dmanisi (Transcaucasia, Georgia)]. Comptes Rendus Palevol (in French) (Elsevier on behalf of the French Academy of Sciences) 1 (4): 243–253. doi:10.1016/S1631-0683(02)00032-5. ISSN 1631-0683.
- Lordkipanidze, David; Vekua, Abesalom; Ferring, Reid; et al. (November 2006). "A fourth hominin skull from Dmanisi, Georgia". The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology (New York: Wiley-Liss) 288A (11): 1146–1157. doi:10.1002/ar.a.20379. ISSN 1552-4884. PMID 17031841.
- Turner, William (April 1895). "On M. Dubois' Description of Remains recently found in Java, named by him Pithecanthropus erectus. With Remarks on so-called Transitional Forms between Apes and Man". Journal of Anatomy and Physiology 29 (Pt 3): 424–445. PMC 1328414. PMID 17232143.
- Spoor, Fred; Wood, Bernard A.; Zonneveld, Frans (June 23, 1994). "Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion". Nature (London: Nature Publishing Group) 369 (6482): 645–648. Bibcode:1994Natur.369..645S. doi:10.1038/369645a0. ISSN 0028-0836. PMID 8208290.
- Bermúdez de Castro, José María; Arsuaga, Juan Luis; Carbonell, Eudald; et al. (May 30, 1997). "A Hominid from the Lower Pleistocene of Atapuerca, Spain: Possible Ancestor to Neandertals and Modern Humans". Science (Washington, D.C.: American Association for the Advancement of Science) 276 (5317): 1392–1395. doi:10.1126/science.276.5317.1392. ISSN 0036-8075. PMID 9162001.
- Carbonell, Eudald; Bermúdez de Castro, José María; Parés, Josep M.; et al. (March 27, 2008). "The first hominin of Europe". Nature (London: Nature Publishing Group) 452 (7186): 465–469. Bibcode:2008Natur.452..465C. doi:10.1038/nature06815. ISSN 0028-0836. PMID 18368116.
- Manzi, Giorgio; Mallegni, Francesco; Ascenzi, Antonio (August 14, 2001). "A cranium for the earliest Europeans: Phylogenetic position of the hominid from Ceprano, Italy". Proc. Natl. Acad. Sci. U.S.A. (Washington, D.C.: National Academy of Sciences) 98 (17): 10011–10016. Bibcode:2001PNAS...9810011M. doi:10.1073/pnas.151259998. ISSN 0027-8424. PMC 55569. PMID 11504953.
- Czarnetzki, Alfred; Jakob, Tina; Pusch, Carsten M. (April 2003). "Palaeopathological and variant conditions of the Homo heidelbergensis type specimen (Mauer, Germany)". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 44 (4): 479–495. doi:10.1016/S0047-2484(03)00029-0. ISSN 0047-2484. PMID 12727464.
- Semaw, Sileshi; Toth, Nicholas; Schick, Kathy; et al. (March 27, 2006). "Scientists discover hominid cranium in Ethiopia" (Press release). Bloomington, IN: Indiana University. Retrieved 2006-11-26.
- Harvati, Katerina (January 2003). "The Neanderthal taxonomic position: models of intra- and inter-specific craniofacial variation". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 44 (1): 107–132. doi:10.1016/S0047-2484(02)00208-7. ISSN 0047-2484. PMID 12604307.
- Herrera, K. J.; Somarelli, J. A.; Lowery, R. K.; Herrera, R. J. (2009). "To what extent did Neanderthals and modern humans interact?". Biological Reviews 84 (2): 245–257. doi:10.1111/j.1469-185X.2008.00071.x. PMID 19391204.
- Krings, Matthias; Stone, Anne; Schmitz, Ralf W.; et al. (July 11, 1997). "Neandertal DNA sequences and the origin of modern humans". Cell (Cambridge, MA: Cell Press) 90 (1): 19–30. doi:10.1016/S0092-8674(00)80310-4. ISSN 0092-8674. PMID 9230299.
- Green, Richard E.; Malaspinas, Anna-Sapfo; Krause, Johannes; et al. (August 8, 2008). "A Complete Neandertal Mitochondrial Genome Sequence Determined by High-Throughput Sequencing". Cell (Cambridge, MA: Cell Press) 134 (3): 416–426. doi:10.1016/j.cell.2008.06.021. ISSN 0092-8674. PMC 2602844. PMID 18692465.
- Serre, David; Langaney, André; Chech, Mario; et al. (March 2004). "No Evidence of Neandertal mtDNA Contribution to Early Modern Humans". PLOS Biology (San Francisco, CA: PLOS) 2 (3): e57. doi:10.1371/journal.pbio.0020057. ISSN 1545-7885. PMC 368159. PMID 15024415.
- Viegas, Jennifer (May 6, 2010). "Neanderthals, Humans Interbred, DNA Proves". Discovery News (Silver Spring, MD: Discovery Communications, LLC.). Retrieved 2015-04-30.
- Gutiérrez, Gabriel; Sánchez, Diego; Marín, Antonio (August 2002). "A Reanalysis of the Ancient Mitochondrial DNA Sequences Recovered from Neandertal Bones". Molecular Biology and Evolution (Oxford, UK: Oxford University Press on behalf of the Society for Molecular Biology and Evolution) 19 (8): 1359–1366. doi:10.1093/oxfordjournals.molbev.a004197. ISSN 0737-4038. PMID 12140248.
- Hebsgaard, Martin B.; Wiuf, Carsten; Gilbert, M. Thomas; et al. (January 2007). "Evaluating Neanderthal Genetics and Phylogeny". Journal of Molecular Evolution (New York: Springer US) 64 (1): 50–60. doi:10.1007/s00239-006-0017-y. ISSN 0022-2844. PMID 17146600.
- Mellars, Paul; French, Jennifer C. (July 29, 2011). "Tenfold Population Increase in Western Europe at the Neandertal–to–Modern Human Transition Paul". Science (Washington, D.C.: American Association for the Advancement of Science) 333 (6042): 623–627. Bibcode:2011Sci...333..623M. doi:10.1126/science.1206930. ISSN 0036-8075. PMID 21798948.
- Brown, Terence A. (April 8, 2010). "Human evolution: Stranger from Siberia". Nature (London: Nature Publishing Group) 464 (7290): 838–839. Bibcode:2010Natur.464..838B. doi:10.1038/464838a. ISSN 0028-0836. PMID 20376137.
- Krause, Johannes; Qiaomei Fu; Good, Jeffrey M.; et al. (April 8, 2010). "The complete mitochondrial DNA genome of an unknown hominin from southern Siberia". Nature (London: Nature Publishing Group) 464 (7290): 894–897. Bibcode:2010Natur.464..894K. doi:10.1038/nature08976. ISSN 0028-0836. PMID 20336068.
- Katsnelson, Alla (March 24, 2010). "New hominin found via mtDNA". The Nutshell (Blog) (Philadelphia, PA: The Scientist). ISSN 0890-3670. Retrieved 2015-05-01.
- Bokma, Folmer; van den Brink, Valentijn; Stadler, Tanja (September 2012). "Unexpectedly many extinct hominins". Evolution (Hoboken, NJ: John Wiley & Sons for the Society for the Study of Evolution) 66 (9): 2969–2974. doi:10.1111/j.1558-5646.2012.01660.x. ISSN 0014-3820. PMID 22946817.
- Martinón-Torres, María; Dennell, Robin; Bermúdez de Castro, José María (February 2011). "The Denisova hominin need not be an out of Africa story". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 60 (2): 251–255. doi:10.1016/j.jhevol.2010.10.005. ISSN 0047-2484. PMID 21129766.
- Brown, Peter; Sutikna, Thomas; Morwood, Michael J.; Soejono, Raden Panji; et al. (October 28, 2004). "A new small-bodied hominin from the Late Pleistocene of Flores, Indonesia". Nature (London: Nature Publishing Group) 431 (7012): 1055–1061. Bibcode:2004Natur.431.1055B. doi:10.1038/nature02999. ISSN 0028-0836. PMID 15514638.
- Argue, Debbie; Donlon, Denise; Groves, Colin; et al. (October 2006). "Homo floresiensis: Microcephalic, pygmoid, Australopithecus, or Homo?". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 51 (4): 360–374. doi:10.1016/j.jhevol.2006.04.013. ISSN 0047-2484. PMID 16919706.
- Martin, Robert D.; Maclarnon, Ann M.; Phillips, James L.; et al. (November 2006). "Flores hominid: New species or microcephalic dwarf?". The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology (New York: Wiley-Liss) 288A (11): 1123–1145. doi:10.1002/ar.a.20389. ISSN 1552-4884. PMID 17031806.
- Cusack, Sinéad (narrator) (February 3, 2000). "Supervolcanoes". Horizon. Series 36. Episode 08. Transcript. BBC2.
- Ambrose, Stanley H. (June 1998). "Late Pleistocene human population bottlenecks, volcanic winter, and differentiation of modern humans". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 34 (6): 623–651. doi:10.1006/jhev.1998.0219. ISSN 0047-2484. PMID 9650103.
- Ambrose, Stanley H. "Volcanic Winter, and Differentiation of Modern Humans". bradshawfoundation.com (Extract from the Journal of Human Evolution). Geneva, Switzerland: Bradshaw Foundation. Retrieved 2015-05-02.
- Schrenk, Friedemann; Kullmer, Ottmar; Bromage, Timothy (2007). "The Earliest Putative Homo Fossils". In Henke, Winfried; Tattersall, Ian. Handbook of Paleoanthropology 1. In collaboration with Thorolf Hardt. Berlin, Heidelberg: Springer. pp. 1611–1631. doi:10.1007/978-3-540-33761-4_52. ISBN 978-3-540-32474-4. Confirmed H. habilis fossils are dated to between 2.1 and 1.5 million years ago. This date range overlaps with the emergence of Homo erectus. Wilford, John Noble (August 9, 2007). "Fossils in Kenya Challenge Linear Evolution". The New York Times. Retrieved 2015-05-04.
- DiMaggio, Erin N.; Campisano, Christopher J.; Rowan, John; et al. (March 20, 2015). "Late Pliocene fossiliferous sedimentary record and the environmental context of early Homo from Afar, Ethiopia". Science (Washington, D.C.: American Association for the Advancement of Science) 347 (6228): 1355–1359. doi:10.1126/science.aaa1415. ISSN 0036-8075. PMID 25739409. Hominins with "proto-Homo" traits may have lived as early as 2.8 million years ago, as suggested by a fossil jawbone classified as transitional between Australopithecus and Homo discovered in 2015.
- Haviland, William A.; Walrath, Dana; Prins, Harald E. L.; McBride, Bunny (2007). Evolution and Prehistory: The Human Challenge (8th ed.). Belmont, CA: Thomson Wadsworth. p. 162. ISBN 978-0-495-38190-7.H. erectus may have appeared some 2 million years ago. Fossils dated to as much as 1.8 million years ago have been found both in Africa and in Southeast Asia, and the oldest fossils by a narrow margin (1.85 to 1.77 million years ago) were found in the Caucasus, so that it is unclear whether H. erectus emerged in Africa and migrated to Eurasia, or if, conversely, it evolved in Eurasia and migrated back to Africa.
- Ferring, R.; Oms, O.; Agusti, J.; Berna, F.; Nioradze, M.; Shelia, T.; Tappen, M.; Vekua, A.; Zhvania, D.; Lordkipanidze, D. (2011). "Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85-1.78 Ma". Proceedings of the National Academy of Sciences 108 (26): 10432. doi:10.1073/pnas.1106638108.
- "New discovery suggests Homo erectus originated from Asia". Daily News and Analysis (Mumbai, India: Diligent Media Corporation Ltd.). Asian News International. June 8, 2011. Retrieved 2015-05-04.
- Frazier, Kendrick (November–December 2006). "Leakey Fights Church Campaign to Downgrade Kenya Museum's Human Fossils". Skeptical Inquirer (Amherst, NY: Committee for Skeptical Inquiry) 30 (6). ISSN 0194-6730. Retrieved 2015-05-04.
- Now also included in H. erectus are Peking Man (formerly Sinanthropus pekinensis) and Java Man (formerly Pithecanthropus erectus). H. erectus is now grouped into various subspecies, including Homo erectus erectus, Homo erectus yuanmouensis, Homo erectus lantianensis, Homo erectus nankinensis, Homo erectus pekinensis, Homo erectus palaeojavanicus, Homo erectus soloensis, Homo erectus tautavelensis, Homo erectus georgicus. The distinction from descendant species such as Homo ergaster, Homo floresiensis, Homo antecessor, Homo heidelbergensis and indeed Homo sapiens is not entirely clear.
- Curnoe, Darren (June 2010). "A review of early Homo in southern Africa focusing on cranial, mandibular and dental remains, with the description of a new species (Homo gautengensis sp. nov.)". HOMO - Journal of Comparative Human Biology (Amsterdam, the Netherlands: Elsevier) 61 (3): 151–177. doi:10.1016/j.jchb.2010.04.002. ISSN 0018-442X. PMID 20466364. A species proposed in 2010 based on the fossil remains of three individuals dated between 1.9 and 0.6 million years ago. The same fossils were also classified as H. habilis, H. ergaster or Australopithecus by other anthropologists.
- Hazarika, Manjil (2007). "Homo erectus/ergaster and Out of Africa: Recent Developments in Paleoanthropology and Prehistoric Archaeology" (PDF). EAA Summer School eBook 1. European Anthropological Association. pp. 35–41. Retrieved 2015-05-04. "Intensive Course in Biological Anthrpology, 1st Summer School of the European Anthropological Association, 16–30 June, 2007, Prague, Czech Republic"
- The type fossil is Mauer 1, dated to ca. 0.6 million years ago. The transition from H. heidelbergensis to H. neanderthalensis at about 0.35 to 0.25 million years ago is largely conventional. Relevant examples are fossils found at Bilzingsleben (also classified as Homo erectus bilzingslebensis).
- Bischoff, James L.; Shamp, Donald D.; Aramburu, Arantza; et al. (March 2003). "The Sima de los Huesos Hominids Date to Beyond U/Th Equilibrium (>350 kyr) and Perhaps to 400–500 kyr: New Radiometric Dates". Journal of Archaeological Science (Amsterdam, the Netherlands: Elsevier) 30 (3): 275–280. doi:10.1006/jasc.2002.0834. ISSN 0305-4403. The first humans with "proto-Neanderthal traits" lived in Eurasia as early as 0.6 to 0.35 million years ago (classified as H. heidelbergensis, also called a chronospecies because it represents a chronological grouping rather than being based on clear morphological distinctions from either H. erectus or H. neanderthalensis), with the first "true Neanderthals" appearing between 0.25 and 0.2 million years ago.
- Chang, Chun-Hsiang; Kaifu, Yousuke; Takai, Masanaru; Kono, Reiko T.; Grün, Rainer; Matsu’ura, Shuji; Kinsley, Les; Lin, Liang-Kong (2015). "The first archaic Homo from Taiwan". Nature Communications 6: 6037. doi:10.1038/ncomms7037.
- "Fossil Reanalysis Pushes Back Origin of Homo sapiens". Scientific American (Stuttgart: Georg von Holtzbrinck Publishing Group). February 17, 2005. ISSN 0036-8733. Retrieved 2015-05-04. The oldest fossil remains of anatomically modern humans are the Omo remains, which date to 195,000 (±5,000) years ago and include two partial skulls as well as arm, leg, foot and pelvis bones.
- McDougall, Ian; Brown, Francis H.; Fleagle, John G. (February 17, 2005). "Stratigraphic placement and age of modern humans from Kibish, Ethiopia". Nature (London: Nature Publishing Group) 433 (7027): 733–736. Bibcode:2005Natur.433..733M. doi:10.1038/nature03258. ISSN 0028-0836. PMID 15716951. H. sapiens idaltu is a confirmed subspecies, based on 3 craniums dated 0.16 – 0.15 Mya found in Ethiopia (1997/2003).
- Jabr, Ferris (July 18, 2012). "Does Thinking Really Hard Burn More Calories?". Scientific American (Stuttgart: Georg von Holtzbrinck Publishing Group). ISSN 0036-8733. Retrieved 2015-05-03.
- Gibbons, Ann (May 29, 1998). "Solving the Brain's Energy Crisis". Science (Washington, D.C.: American Association for the Advancement of Science) 280 (5368): 1345–1347. doi:10.1126/science.280.5368.1345. ISSN 0036-8075. PMID 9634409.
- Robinson 2008, p. 398
- Freeman & Herron 2007, pp. 786–788
- Ward, Carol V.; Tocheri, Matthew W.; Plavcan, J. Michael; et al. (January 7, 2014). "Early Pleistocene third metacarpal from Kenya and the evolution of modern human-like hand morphology". Proc. Natl. Acad. Sci. U.S.A. (Washington, D.C.: National Academy of Sciences) 111 (1): 121–124. doi:10.1073/pnas.1316014110. ISSN 0027-8424. PMC 3890866. PMID 24344276.
- Plummer, Thomas (2004). "Flaked stones and old bones: Biological and cultural evolution at the dawn of technology". American Journal of Physical Anthropology (Hoboken, NJ: John Wiley & Sons for the American Association of Physical Anthropologists). Supplement 39 (Yearbook of Physical Anthropology): 118–164. doi:10.1002/ajpa.20157. ISSN 0002-9483. PMID 15605391.
- Wong, Kate (April 15, 2015). "Archaeologists Take Wrong Turn, Find World’s Oldest Stone Tools". Scientific American (Blog) (Stuttgart: Georg von Holtzbrinck Publishing Group). ISSN 0036-8733. Retrieved 2015-05-03.
- Balter, Michael (April 14, 2015). "World's oldest stone tools discovered in Kenya". Science (News) (Washington, D.C.: American Association for the Advancement of Science). doi:10.1126/science.aab2487. ISSN 0036-8075. Retrieved 2015-05-03.
- Drake, Nadia (April 16, 2015). "Oldest Stone Tools Discovered in Kenya". National Geographic News (Washington, D.C.: National Geographic Society). Retrieved 2015-05-03.
- Diamond 1999, p. 39
- Bar-Yosef, Ofer (October 2002). "The Upper Paleolithic Revolution". Annual Review of Anthropology (Palo Alto, CA: Annual Reviews) 31: 363–393. doi:10.1146/annurev.anthro.31.040402.085416. ISSN 0084-6570.
- Oppenheimer, Stephen (2012) "Out of Eden: the peopling of the world" (Robinson)
- Nowell, April (October 2010). "Defining Behavioral Modernity in the Context of Neandertal and Anatomically Modern Human Populations". Annual Review of Anthropology (Palo Alto, CA: Annual Reviews) 39: 437–452. doi:10.1146/annurev.anthro.012809.105113. ISSN 0084-6570.
- d'Errico, Francesco; Stringer, Chris B. (April 12, 2011). "Evolution, revolution or saltation scenario for the emergence of modern cultures?". Philosophical Transactions of the Royal Society B (London: Royal Society) 366 (1567): 1060–1069. doi:10.1098/rstb.2010.0340. ISSN 0962-8436. PMC 3049097. PMID 21357228.
- Ambrose, Stanley H. (March 2, 2001). "Paleolithic Technology and Human Evolution". Science (Washington, D.C.: American Association for the Advancement of Science) 291 (5509): 1748–1753. Bibcode:2001Sci...291.1748A. doi:10.1126/science.1059487. ISSN 0036-8075. PMID 11249821.
- Mellars P. (2010). Neanderthal symbolism and ornament manufacture: The bursting of a bubble? Proc Natl Acad Sci U S A. doi:10.1073/pnas.1014588107
- Medical Research Council (UK) (November 21, 2009). "Brain Disease 'Resistance Gene' evolves in Papua New Guinea community; could offer insights Into CJD". ScienceDaily (Rockville, MD: ScienceDaily, LLC). Retrieved 2009-11-22.
- Mead, S.; Whitfield, J.; Poulter, M.; Shah, P.; Uphill, J.; Campbell, T.; Al-Dujaily, H.; Hummerich, H.; Beck, J.; Mein, C. A.; Verzilli, C.; Whittaker, J.; Alpers, M. P.; Collinge, J. (2009). "A Novel Protective Prion Protein Variant that Colocalizes with Kuru Exposure". The New England Journal of Medicine 361 (21): 2056–2065. doi:10.1056/NEJMoa0809716. PMID 19923577.
- Byars, S. G.; Ewbank, D.; Govindaraju, D. R.; Stearns, S. C. (2009). "Natural selection in a contemporary human population". Proceedings of the National Academy of Sciences 107 (suppl_1): 1787–1792. Bibcode:2010PNAS..107.1787B. doi:10.1073/pnas.0906199106. PMC 2868295. PMID 19858476.
- Cochran & Harpending 2009
- Diamond 1999
- Aiello, Leslie; Dean, Christopher (1990). An Introduction to Human Evolutionary Anatomy. London; San Diego: Elsevier Academic Press. ISBN 0-12-045591-9. LCCN 95185095. OCLC 33408268.
- American Heritage Dictionaries (editors) (2006). More Word Histories and Mysteries: From Aardvark to Zombie. Boston, MA: Houghton Mifflin. ISBN 978-0-618-71681-4. LCCN 2006020835. OCLC 70199867.
- Bogin, Barry (1997). "The Evolution of Human Nutrition". In Romanucci-Ross, Lola; Moerman, Daniel E.; Tancredi, Laurence R. The Anthropology of Medicine: From Culture to Method (3 ed.). Westport, CT: Bergin & Garvey. ISBN 0-89789-516-9. LCCN 96053993. OCLC 36165190.
- Bown, Thomas M.; Rose, Kenneth D. (1987). Patterns of Dental Evolution in Early Eocene Anaptomorphine Primates (Omomyidae) From the Bighorn Basin, Wyoming (PDF). Memoir (Paleontological Society) 23. Tulsa, OK: Paleontological Society. OCLC 16997265.
- Boyd, Robert; Silk, Joan B. (2003). How Humans Evolved (3rd ed.). New York: Norton. ISBN 0-393-97854-0. LCCN 2002075336. OCLC 49959461.
- Brues, Alice M.; Snow, Clyde C. (1965). "Physical Anthropology". In Siegel, Bernard J. Biennial Review of Anthropology 1965. Biennial Review of Anthropology 4 (Stanford, CA: Stanford University Press). ISBN 0-8047-1746-X. ISSN 0067-8503. LCCN 59012726. OCLC 01532912.
- Bryson, Bill (2004) [Originally published 2003]. "The Mysterious Biped". A Short History of Nearly Everything. Toronto, Canada: Anchor Canada. ISBN 0-385-66004-9. LCCN 2003046006. OCLC 55016591.
- Cameron, David W. (2004). Hominid Adaptations and Extinctions. Sydney, NSW: UNSW Press. ISBN 0-86840-716-X. LCCN 2004353026. OCLC 57077633.
- Cochran, Gregory; Harpending, Henry (2009). The 10,000 Year Explosion: How Civilization Accelerated Human Evolution. New York: Basic Books. ISBN 978-0-465-00221-4. LCCN 2008036672. OCLC 191926088.
- Curry, James R. (2008). Children of God: Children of Earth. Bloomington, IN: AuthorHouse. ISBN 978-1-4389-1846-4. OCLC 421466369.
- Darwin, Charles (1981) [Originally published 1871; London: John Murray]. The Descent of Man, and Selection in Relation to Sex. Introduction by John Tyler Bonner and Robert M. May (Reprint ed.). Princeton, NJ: Princeton University Press. ISBN 0-691-02369-7. LCCN 80008679. OCLC 7197127.
- Dawkins, Richard (2004). The Ancestor's Tale: A Pilgrimage to the Dawn of Evolution. Boston, MA: Houghton Mifflin. ISBN 0-618-00583-8. LCCN 2004059864. OCLC 56617123.
- DeSalle, Rob; Tattersall, Ian (2008). Human Origins: What Bones and Genomes Tell Us About Ourselves. Texas A&M University Anthropology Series 13 (1st ed.). College Station, TX: Texas A&M University Press. ISBN 978-1-58544-567-7. OCLC 144520427.
- Diamond, Jared (1999). Guns, Germs, and Steel: The Fates of Human Societies. New York: W. W. Norton & Company. ISBN 0-393-31755-2. LCCN 2005284124. OCLC 35792200.
- Freeman, Scott; Herron, Jon C. (2007). Evolutionary Analysis (4th ed.). Upper Saddle River, NJ: Pearson Prentice Hall. ISBN 978-0-13-227584-2. LCCN 2006034384. OCLC 73502978.
- Montgomery, William M. (1988) [Originally published 1974]. "Germany". In Glick, Thomas F. The Comparative Reception of Darwinism. Chicago, IL: University of Chicago Press. ISBN 0-226-29977-5. LCCN 87035814. OCLC 17328115. "The Conference on the Comparative Reception of Darwinism was held in Austin, Texas, on April 22 and 23, 1972, under the joint sponsorship of the American Council of Learned Societies and the University of Texas at Austin"
- Kondo, Shiro, ed. (1985). Primate Morphophysiology, Locomotor Analyses, and Human Bipedalism. Tokyo: University of Tokyo Press. doi:10.1002/ajpa.1330700214. ISBN 4-13-066093-4. LCCN 85173489. OCLC 12352830.
- Kutty, Ahamed V. P. (2009). Adam's Gene and the Mitochondrial Eve: The Non-incestuous Descent of Man from Adam and Eve. Bloomington, IN: Xlibris Corporation. ISBN 978-1-4415-0729-7. OCLC 741127863.
- Leakey, Richard E. (1994). The Origin of Humankind. Science Masters Series. New York: Basic Books. ISBN 0-465-03135-8. LCCN 94003617. OCLC 30739453.
- M'charek, Amade (2005). The Human Genome Diversity Project: An Ethnography of Scientific Practice. Cambridge Studies in Society and the Life Sciences. Cambridge; New York: Cambridge University Press. ISBN 0-521-83222-5. LCCN 2004052648. OCLC 55600894.
- Martin, Robert D. (2001). "Primates, Evolution of". In Smelser, Neil J.; Baltes, Paul B. International Encyclopedia of the Social & Behavioral Sciences (1st ed.). Amsterdam; New York: Elsevier. doi:10.1016/B0-08-043076-7/03083-7. ISBN 978-0-08-043076-8. LCCN 2001044791. OCLC 47869490.
- Maxwell, Mary (1984). Human Evolution: A Philosophical Anthropology. New York: Columbia University Press. ISBN 0-231-05946-9. LCCN 83024005. OCLC 10163036.
- McHenry, Henry M. (2009). "Human Evolution". In Ruse, Michael; Travis, Joseph. Evolution: The First Four Billion Years. Foreword by Edward O. Wilson. Cambridge, MA: Belknap Press of Harvard University Press. ISBN 978-0-674-03175-3. LCCN 2008030270. OCLC 225874308.
- Ramachandran, Sohini; Hua Tang; Gutenkunst, Ryan N.; Bustamante, Carlos D. (2010). "Genetics and Genomics of Human Population Structure". In Speicher, Michael R.; Antonarakis, Stylianos E.; Motulsky, Arno G. Vogel and Motulsky's Human Genetics: Problems and Approaches (4th completely rev. ed.). Heidelberg; London; New York: Springer. doi:10.1007/978-3-540-37654-5. ISBN 978-3-540-37653-8. LCCN 2009931325. OCLC 549541244.
- Robinson, J. T. (2008) [Originally published 1963; Chicago, IL: Aldine Transaction]. "Adaptive Radiation in the Australopithecines and the Origin of Man". In Howell, F. Clark; Bourlière, François. African Ecology and Human Evolution. New Brunswick, NJ: Transaction Publishers. ISBN 978-0-202-36136-9. LCCN 2007024716. OCLC 144770218.
- Srivastava, R. P. (2009). Morphology of the Primates and Human Evolution. New Delhi: PHI Learning Private Limited. ISBN 978-81-203-3656-8. OCLC 423293609.
- Stanford, Craig; Allen, John S.; Antón, Susan C. (2009). Biological Anthropology: The Natural History of Humankind (2nd ed.). Upper Saddle River, NJ: Pearson Prentice Hall. ISBN 978-0-13-601160-6. LCCN 2007052429. OCLC 187548835.
- Strickberger, Monroe W. (2000). Evolution (3rd ed.). Sudbury, MA: Jones and Bartlett Publishers. ISBN 0-7637-1066-0. LCCN 99032072. OCLC 41431683.
- Stringer, Chris B. (1994) [First published 1992]. "Evolution of Early Humans". In Jones, Steve; Martin, Robert D.; Pilbeam, David. The Cambridge Encyclopedia of Human Evolution. Foreword by Richard Dawkins (1st paperback ed.). Cambridge, UK: Cambridge University Press. ISBN 0-521-32370-3. LCCN 92018037. OCLC 444512451.
- Swisher, Carl C., III; Curtis, Garniss H.; Lewin, Roger (2001) [Originally published 2000]. Java Man: How Two Geologists Changed Our Understanding of Human Evolution. Chicago, IL: University of Chicago Press. ISBN 0-226-78734-6. LCCN 2001037337. OCLC 48066180.
- Tishkoff, S.A.; Reed, F.A.; et al. (22 May 2009). "The Genetic Structure and History of Africans and African Americans". Science 324 (5930): 1035–1044. doi:10.1126/science.1172257. PMC 2947357.
- Trent, Ronald J. (2005). Molecular Medicine: An Introductory Text (3rd ed.). Burlington, MA: Elsevier Academic Press. ISBN 0-12-699057-3. LCCN 2004028087. OCLC 162577235.
- Trevathan, Wenda R. (2011) [Originally published 1987; New York: Aldine De Gruyter]. Human Birth: An Evolutionary Perspective. New Brunswick, NJ: Transaction Publishers. ISBN 978-1-4128-1502-4. LCCN 2010038249. OCLC 669122326.
- Ungar, Peter S.; Teaford, Mark F., eds. (2002). Human Diet: Its Origin and Evolution. Westport, CT: Bergin & Garvey. ISBN 0-89789-736-6. LCCN 2001043790. OCLC 537239907.
- Walker, Alan (2007). "Early Hominin Diets: Overview and Historical Perspectives". In Ungar, Peter. Evolution of the Human Diet: The Known, the Unknown, and the Unknowable. Human Evolution Series. Oxford; New York: Oxford University Press. ISBN 978-0-19-518346-7. LCCN 2005036120. OCLC 132816551.
- Wallace, David Rains (2004). Beasts of Eden: Walking Whales, Dawn Horses, and Other Enigmas of Mammal Evolution. Berkeley, CA: University of California Press. ISBN 0-520-24684-5. LCCN 2003022857. OCLC 53254011.
- Webster, Donovan (2010). Meeting the Family: One Man's Journey Through His Human Ancestry. Foreword by Spencer Wells. Washington, D.C.: National Geographic Society. ISBN 978-1-4262-0573-6. LCCN 2009050471. OCLC 429022321.
- Wood, Bernard A. (2009). "Where Does the Genus Homo Begin, and How Would We Know?". In Grine, Frederick E.; Fleagle, John G.; Leakey, Richard E. The First Humans: Origin and Early Evolution of the Genus Homo. Vertebrate Paleobiology and Paleoanthropology. Dordrecht, the Netherlands: Springer Netherlands. doi:10.1007/978-1-4020-9980-9_3. ISBN 978-1-4020-9979-3. ISSN 1877-9077. LCCN 2009927083. OCLC 310400980. "Contributions from the Third Stony Brook Human Evolution Symposium and Workshop October 3–7, 2006."
- Alexander, Richard D. (1990). How Did Humans Evolve? Reflections on the Uniquely Unique Species (PDF). Special Publication (1). Ann Arbor, MI: Museum of Zoology, University of Michigan. pp. 1–38. LCCN 90623893. OCLC 22860997.
- Barton, Nicholas H.; Briggs, Derek E. G.; Eisen, Jonathan A.; et al. (2007). Evolution. Cold Spring Harbor, NY: Cold Spring Harbor Laboratory Press. ISBN 978-0-87969-684-9. LCCN 2007010767. OCLC 86090399.
- Enard, Wolfgang; Przeworski, Molly; Fisher, Simon E.; et al. (August 22, 2002). "Molecular evolution of FOXP2, a gene involved in speech and language". Nature (London: Nature Publishing Group) 418 (6900): 869–872. doi:10.1038/nature01025. ISSN 0028-0836. PMID 12192408.
- Flinn, Mark V.; Geary, David C.; Ward, Carol V. (January 2005). "Ecological dominance, social competition, and coalitionary arms races: Why humans evolved extraordinary intelligence" (PDF). Evolution and Human Behavior (Amsterdam, the Netherlands: Elsevier) 26 (1): 10–46. doi:10.1016/j.evolhumbehav.2004.08.005. ISSN 1090-5138. Retrieved 2015-05-05.
- Gibbons, Ann (2006). The First Human: The Race to Discover our Earliest Ancestors (1st ed.). New York: Doubleday. ISBN 978-0-385-51226-8. LCCN 2005053780. OCLC 61652817.
- Hartwig, Walter C., ed. (2002). The Primate Fossil Record. Cambridge Studies in Biological and Evolutionary Anthropology 33. Cambridge; New York: Cambridge University Press. ISBN 0-521-66315-6. LCCN 2001037847. OCLC 47254191.
- Heizmann, Elmar P. J.; Begun, David R. (November 2001). "The oldest Eurasian hominoid". Journal of Human Evolution (Amsterdam, the Netherlands: Elsevier) 41 (5): 463–481. doi:10.1006/jhev.2001.0495. ISSN 0047-2484. PMID 11681862.
- Hill, Andrew; Ward, Steven (1988). "Origin of the hominidae: The record of African large hominoid evolution between 14 my and 4 my". American Journal of Physical Anthropology (Hoboken, NJ: John Wiley & Sons for the American Association of Physical Anthropologists) 31 (59): 49–83. doi:10.1002/ajpa.1330310505. ISSN 0002-9483.
- Ijdo, Jacob W.; Baldini, Antonio; Ward, David C.; et al. (October 15, 1991). "Origin of human chromosome 2: An ancestral telomere-telomere fusion" (PDF). Proc. Natl. Acad. Sci. U.S.A. (Washington, D.C.: National Academy of Sciences) 88 (20): 9051–9055. Bibcode:1991PNAS...88.9051I. doi:10.1073/pnas.88.20.9051. ISSN 0027-8424. PMC 52649. PMID 1924367. Retrieved 2015-05-05. — two ancestral ape chromosomes fused to give rise to human chromosome 2
- Johanson, Donald; Wong, Kate (2010). Lucy's Legacy: The Quest for Human Origins. New York: Three Rivers Press. ISBN 978-0-307-39640-2. LCCN 2010483830. OCLC 419801728.
- Jones, Steve; Martin, Robert D.; Pilbeam, David, eds. (1994) [First published 1992]. The Cambridge Encyclopedia of Human Evolution. Foreword by Richard Dawkins (1st paperback ed.). Cambridge, UK: Cambridge University Press. ISBN 0-521-32370-3. LCCN 92018037. OCLC 444512451. (Note: this book contains very useful, information dense chapters on primate evolution in general, and human evolution in particular, including fossil history).
- Leakey, Richard E.; Lewin, Roger (1992). Origins Reconsidered: In Search of What Makes us Human. New York: Doubleday. ISBN 0-385-41264-9. LCCN 92006661. OCLC 25373161.
- Lewin, Roger (1997). Bones of Contention: Controversies in the Search for Human Origins (2nd ed.). Chicago, IL: University of Chicago Press. ISBN 0-226-47651-0. LCCN 97000972. OCLC 36181117.
- Morwood, Mike; van Oosterzee, Penny (2007). A New Human: The Startling Discovery and Strange Story of the 'Hobbits' of Flores, Indonesia (1st Smithsonian Books ed.). New York: Smithsonian Books/HarperCollins. ISBN 978-0-06-089908-0. LCCN 2006052267. OCLC 76481584.
- Oppenheimer, Stephen (2003). Out of Eden: The Peopling of the World. london: Constable & Robinson. ISBN 1-84119-697-5. LCCN 2005482222. OCLC 52195607.
- Ovchinnikov, Igor V.; Götherström, Anders; Romanova, Galina P.; et al. (March 30, 2000). "Molecular analysis of Neanderthal DNA from the Northern Caucasus". Nature (London: Nature Publishing Group) 404 (6777): 490–493. doi:10.1038/35006625. ISSN 0028-0836. PMID 10761915.
- Roberts, Alice M. (2009). The Incredible Human Journey: The Story of How We Colonised the Planet. London: Bloomsbury Publishing. ISBN 978-0-7475-9839-8. OCLC 310156315.
- Shreeve, James (1995). The Neandertal Enigma: Solving the Mystery of Modern Human Origins. New York: Morrow. ISBN 0-688-09407-4. LCCN 95006337. OCLC 32088673.
- Stringer, Chris B. (2011). The Origin of Our Species. London: Allen Lane. ISBN 978-1-84614-140-9. LCCN 2011489742. OCLC 689522193.
- Stringer, Chris B.; Andrews, Peter (2005). The Complete World of Human Evolution. London; New York: Thames & Hudson. ISBN 0-500-05132-1. LCCN 2004110563. OCLC 224377190.
- Stringer, Christopher; McKie, Robin (1997). African Exodus: The Origins of Modern Humanity (1st American ed.). New York: Henry Holt and Company. ISBN 0-8050-2759-9. LCCN 96037718. OCLC 36001167.
- Tattersall, Ian (2008). The Fossil Trail: How We Know What We Think We Know About Human Evolution (2nd ed.). New York: Oxford University. ISBN 978-0195367669. LCCN 2008013654. OCLC 218188644.
- van Oosterzee, Penny (1999). Dragon Bones: The Story of Peking Man. St Leonards, New South Wales: Allen & Unwin. ISBN 1-86508-123-X. LCCN 00300421. OCLC 45853997.
- Wade, Nicholas (2006). Before the Dawn: Recovering the Lost History of Our Ancestors. New York: Penguin Press. ISBN 1-59420-079-3. LCCN 2005055293. OCLC 62282400.
- Walker, Alan; Shipman, Pat (1996). The Wisdom of the Bones: In Search of Human Origins. London: Weidenfeld & Nicolson. ISBN 0-297-81670-5. OCLC 35202130.
- Weiss, Mark L.; Mann, Alan E. (1985). Human Biology and Behavior: An Anthropological Perspective (4th ed.). Boston: Little Brown. ISBN 0-316-92894-1. LCCN 85000158. OCLC 11726796. (Note: this book contains very accessible descriptions of human and non-human primates, their evolution, and fossil history).
- Wells, Spencer (2003) [Originally published 2002; Princeton, NJ: Princeton University Press]. The Journey of Man: A Genetic Odyssey (Random House trade paperback ed.). New York: Random House Trade Paperbacks. ISBN 0-8129-7146-9. LCCN 2003066679. OCLC 53287806.
|Wikimedia Commons has media related to Human evolution.|
- "BBC - Science & Nature - The evolution of man". BBC. Retrieved 2015-05-06.
- "Becoming Human". Arizona State University's Institute of Human Origins. Retrieved 2015-05-06.
- "Bones, Stones and Genes: The Origin of Modern Humans" (Video lecture series). Howard Hughes Medical Institute. Retrieved 2015-05-06.
- "Evolution Figures: Chapter 25". Cold Spring Harbor Laboratory Press. Retrieved 2015-05-06. — Illustrations from the book Evolution (2007)
- "Human Evolution". Smithsonian Institution's Human Origins Program. Retrieved 2013-06-24.
- "Human Evolution Timeline". ArchaeologyInfo.com. Retrieved 2013-06-24.
- Lambert, Tim (Producer) (June 24, 2015). First Peoples. London: Wall to Wall Television. OCLC 910115743. Retrieved 2015-07-18.
- Shaping Humanity Video 2013 Yale University