Human evolution

Homo sapiens is a distinct species of the

hominid family of primates, which
includes all the great apes.[2] Over their
evolutionary history, humans gradually
developed traits such as bipedalism,
dexterity, and complex language.[3]
Modern humans interbred with archaic
humans,[4] indicating that their evolution
was not linear but weblike.[5][6][7][8] The
study of the origins of humans involves
several scientific disciplines, including
physical and evolutionary anthropology,
paleontology, and genetics; the field is
also known by the terms anthropogeny,
anthropogenesis, and A phylogenic model depicts (from lower right) Homo erectus
[9][10] spreading from Africa to Eurasia around 1.8 millions of years
anthropogony —with the latter two
ago. Its descendant H. heidelbergensis originated Denisovans,
sometimes used to refer to the related Neanderthals, and H. sapiens (humans) around 350,000 years
subject of hominization. ago.[1]

Primates diverged from other mammals

about 85 million years ago, in the Late Cretaceous period, with their earliest fossils appearing over
55 mya, during the Paleocene.[11] Primates produced successive clades leading to the ape superfamily,
which gave rise to the hominid and the gibbon families; these diverged some 15–20 mya. African and
Asian hominids (including orangutans) diverged about 14 mya. Hominins (including the
Australopithecine and Panina subtribes) parted from the Gorillini tribe between 8 and 9 mya;
Australopithecine (including the extinct biped ancestors of humans) separated from the Pan genus
(containing chimpanzees and bonobos) 4–7 mya.[12] The Homo genus is evidenced by the appearance of
H. habilis over 2 mya,[a] while anatomically modern humans emerged in Africa approximately
300,000 years ago.

Before Homo

Early evolution of primates

The evolutionary history of primates can be traced back 65 million years.[13][14][15][16][17] The details of
the origins and early evolution of primates are however still unknown to a large extent due to scarcity of
fossil evidence. One of the oldest known primate-like mammal species, the Plesiadapis, came from North
America;[18][19][20][21][22][23] another, Archicebus, came from China.[24] Other such early primates
include Altiatlasius and Algeripithecus, which were found in Northern Africa.[25][26] Other similar basal
primates were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and
Eocene.

David R. Begun theorized that early primates flourished first

in Eurasia and that a lineage leading to the African apes and
humans, including Dryopithecus, migrated south from Europe
or Western Asia into Africa.[27] However, earlier potential
ancestors of primates such as Altiatlasius are known to have
lived on the African continent much earlier than
Dryopithecus.[25]

The surviving tropical population of primates—which is seen

most completely in the Upper Eocene and lowermost Notharctus tenebrosus, American
Oligocene fossil beds of the Faiyum depression southwest of Museum of Natural History, New York
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 the uppermost Oligocene at Eragaleit in the
northern Great Rift Valley in Kenya, dated to 24 million years ago.[28] Its ancestry is thought to be
species related to Aegyptopithecus, Propliopithecus, and Parapithecus from the Faiyum, at around
35 mya.[29] In 2010, Saadanius was described as a close relative of the last common ancestor of the
crown catarrhines, and tentatively dated to 29–28 mya, helping to fill an 11-million-year gap in the fossil
record.[30]

In the Early Miocene, about 22 million years ago, the many

kinds of arboreally-adapted (tree-dwelling) 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.
Reconstructed tailless Proconsul
skeleton
The presence of other generalized non-cercopithecids of
Middle Miocene from sites far distant, such as 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 diverged from the line of great apes some 18–12
mya, and that of orangutans (subfamily Ponginae)[b] 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 Southeast
Asian hominoid population, but fossil
proto-orangutans may be represented by
Sivapithecus from India and
Griphopithecus from Turkey, dated to
around 10 mya.[31]

Hominidae subfamily Homininae

(African hominids) diverged from
Ponginae (orangutans) about 14 mya.
Hominins (including humans and the
The ape superfamily (hominoids) comprises: the lesser apes
Australopithecine and Panina subtribes) (including gibbons) and great apes (chimpanzees, gorillas, and
parted from the Gorillini tribe (gorillas) humans)
between 8 and 9 mya; Australopithecine
(including the extinct biped ancestors of
humans) separated from the Pan genus (containing chimpanzees and bonobos) 4–7 mya.[12] The Homo
genus is evidenced by the appearance of H. habilis over 2 mya,[a] while anatomically modern humans
emerged in Africa approximately 300,000 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.
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, Genetic data suggest that
chimpanzees and humans may
pigs, elephants, horses, and others. The equatorial belt contracted after
have diverged from a common
about 8 million years ago, and there is very little fossil evidence for the ancestor between 9 and 7
split—thought to have occurred around that time—of the hominin million years ago.[32]
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.[33] 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 the
genus Pan, may have evolved via the process of self-
domestication. Consequently, arguing against the so-called
"chimpanzee referential model"[34] the authors suggest it is
no longer tenable to use 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
Chimpanzee nest. Later hominins may
has for the evolution of hominin social psychology, they have developed niche creating shelter-
wrote: building traditions from such earlier nest-
building practices.
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.[33]: 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.

Genus Australopithecus
The genus Australopithecus 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, A. afarensis, A. sediba, and
A. africanus. There is still some debate among academics whether certain African hominid species of this
time, such as P. robustus and P. boisei, constitute members of the same genus; if so, they would be
considered to be "robust australopiths" while the others would be considered "gracile australopiths".
However, if these species do indeed constitute their own genus, then they may be given their own name,
Paranthropus.

Australopithecus (4–1.8 Ma), with species A. anamensis, A. afarensis, A. africanus, A.

 bahrelghazali, A. garhi, and A. 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 A. afarensis.
There is debate whether A. deyiremeda is a new species or is A.
afarensis.[35] Australopithecus prometheus, otherwise known as Little
Foot, has recently been dated at 3.67 million years old through a new
dating technique, making the genus Australopithecus as old as
afarensis.[36] Given the opposable big toe found on Little Foot, it
seems that the specimen was a good climber. It is thought given the
night predators of the region that he built a nesting platform at night in
the trees in a similar fashion to chimpanzees and gorillas.
Reconstruction of "Lucy"

Evolution of genus Homo

The earliest documented
representative of the genus Hominin timeline
Homo is the Ledi jaw, 0— Modern humans
←
←
which is dated 2.75[37][38] – Homo sapiens ← Earliest clothes
Denisovans Earliest rock art
2.8 million years ago P
– l Neanderthals
H. heidelbergensis
(mya),[39][40] and is e
arguably the earliest species −1 — i H. erectus
s
for which there is positive t (H. antecessor) P
evidence of the use of stone –o (H. ergaster) a
c (Au. sediba) ← Earliest fire / cooking
tools. The brains of these e r

early hominins were about −2 — n a

e H. habilis ← Dispersal beyond Africa
the same size as that of a n
(H. rudolfensis)
– t
chimpanzee, although it has (Au. garhi)
been suggested that this
h ← Earliest sign of
−3 — r
Homo
was the time in which the Australopithecus
P (Au. africanus)
o
human SRGAP2 gene ← Earliest stone tools
– l (Au. afarensis) p
doubled, producing a more i
(Au. anamensis) u
o
rapid wiring of the frontal −4 — c s
cortex. During the next e
n Ardipithecus
← Earliest
million years a process of sign of
–e Australopithecus
(Ar. ramidus)
rapid encephalization
occurred, and with the −5 — Hominini
arrival of Homo erectus and
Homo ergaster in the fossil –
← Earliest
sign of
record, cranial capacity had (Ar. kadabba)
Ardipithecus
doubled to 850 cm3.[41] −6 — Orrorin
(Such an increase in human (O. praegens)
– (O. tugenensis)
brain size is equivalent to
M
7 Earliest bipedal
 −7 — i ← Earliest bipedal
each generation having Sahelanthropus H
o
125,000 more neurons than
–ce
Graecopithecus o
their parents.) It is believed Oreopithecus
n
that H. erectus and −8 — e Chororapithecus m
H. ergaster were the first to i
Sivapithecus
use fire and complex tools, – ← Chimpanzee split
and they were the first of Ouranopithecus n
the hominin line to leave −9 — (Ou. turkae) ← Gorilla split
i
(Ou. macedoniensis)
Africa, spreading
throughout Africa, Asia, – Samburupithecusd
and Europe between Nakalipithecus s
1.3 to 1.8 mya. −10 — ← Earlier apes
(million years ago)
According to the recent
African origin theory, modern humans evolved in Africa possibly from H. heidelbergensis,
H. rhodesiensis or H. antecessor and migrated out of the continent some 50,000 to 100,000 years ago,
gradually replacing local populations of H. erectus, Denisova hominins, H. floresiensis, H. luzonensis and
H. neanderthalensis, whose ancestors had left Africa in earlier migrations.[42][43][44][45][46] Archaic Homo
sapiens, the forerunner of anatomically modern humans, evolved in the Middle Paleolithic between
400,000 and 250,000 years ago.[47][48][49] 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 interbreeding between these species.[50][51][52] According to some anthropologists, the
transition to behavioral modernity with the development of symbolic culture, language, and specialized
lithic technology happened around 50,000 years ago (beginning of the Upper Paleolithic),[53] although
others point to evidence of a gradual change over a longer time span during the Middle Paleolithic.[54]

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.[55][56] There is yet no consensus as to which of these groups should be
considered a separate species and which should be subspecies; this may be due to the dearth of fossils or
to the slight differences used to classify species in the genus Homo.[56] The Sahara pump theory
(describing an occasionally passable "wet" Sahara desert) provides one possible explanation of the
intermittent migration and speciation in the genus Homo.

Based on archaeological and paleontological evidence, it has been possible to infer, to some extent, the
ancient dietary practices[57] of various Homo species and to study the role of diet in physical and
behavioral evolution within Homo.[58][59][60][61][62]

Some anthropologists and archaeologists subscribe to the Toba catastrophe theory, which posits that the
supereruption of Lake Toba on Sumatra in Indonesia some 70,000 years ago caused global starvation,[63]
killing the majority of humans and creating a population bottleneck that affected the genetic inheritance
of all humans today.[64] The genetic and archaeological evidence for this remains in question however.[65]
A 2023 genetic study suggests that a similar human population bottleneck of between 1,000 and 100,000
survivors occurred "around 930,000 and 813,000 years ago ... lasted for about 117,000 years and brought
human ancestors close to extinction."[66][67]
H. habilis and H. gautengensis
Homo habilis lived from about 2.8[40] 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 with the
development of smaller molars and larger brains. One of the first known hominins, it made tools from
stone and perhaps animal bones, leading to its name homo habilis (Latin 'handy man') bestowed by
discoverer Louis Leakey. Some scientists have proposed moving this species from Homo into
Australopithecus due to the morphology of its skeleton being more adapted to living in trees rather than
walking on two legs like later hominins.[68]

In May 2010, a new species, Homo gautengensis, was proposed based on specimens discovered in South
Africa.[69]

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 a specimen of Homo habilis, but this has not been confirmed.[70] It is currently
thought that H. rudolfensis is an early hominid species, alongside H. erectus and H.
habilis.[71]
Homo georgicus, from Georgia, may be an intermediate form between Homo habilis and
Homo erectus,[72] or a subspecies of Homo erectus.[73]

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 Anthropopithecus
erectus (1892–1893, considered at this point as a chimpanzee-
like fossil primate) and Pithecanthropus erectus (1893–1894,
changing his mind as of based on its morphology, which he
considered to be intermediate between that of humans and
apes).[74] Years later, in the 20th century, the German
physician and paleoanthropologist Franz Weidenreich (1873–
1948) compared in detail the characters of Dubois' Java Man, Reconstruction of Turkana Boy who
then named Pithecanthropus erectus, with the characters of lived 1.5 to 1.6 million years ago
the Peking Man, then named Sinanthropus pekinensis.
Weidenreich concluded in 1940 that because of their
anatomical similarity with modern humans it was necessary to gather all these specimens of Java and
China in a single species of the genus Homo, the species H. erectus.[75][76]

Homo erectus lived from about 1.8 Ma to about 108,000 years ago.[77] This population appears to have
died out when the savannah corridors closed, and tropical jungle took over.;[78] however, nearby
H. floresiensis survived it. The early phase of H. erectus, from 1.8 to 1.25 Ma, is considered by some to
be a separate species, H. ergaster, or as H. erectus ergaster, a subspecies of H. erectus. Many
paleoanthropologists now use the term Homo ergaster for the non-Asian forms of this group, and reserve
H. erectus only for those fossils that are found in Asia and meet certain skeletal and dental requirements
which differ slightly from H. ergaster.

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.[79] This species
also may have used fire to cook meat. Richard Wrangham notes that Homo seems to have been ground
dwelling, with reduced intestinal length, smaller dentition, and "brains [swollen] to their current,
horrendously fuel-inefficient size",[80] and hypothesizes that control of fire and cooking, which released
increased nutritional value, was the key adaptation that separated Homo from tree-sleeping
Australopithecines.[81]

H. cepranensis and H. antecessor

These are proposed as species 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.[82][83] Two H. antecessor scapulae fossils were discovered at the Gran Dolina cave site
in Spain. Researchers compared the found scapulae with other fossils of chimpanzees,
modern humans, and troglodytes. They found that the H. antecessor fossils were more
developmentally similar to modern humans. This corroborates with previous findings on H.
antecessor skulls, where their cranial bones display features that both Chinese Middle
Pleistocene hominins and modern humans share. These findings suggest that H.
cepranensis may be the last common ancestor between H. sapiens, H. neanderthalensis
and Denisovans, though more research would need to be done to verify this hypothesis.[84]
A 2020 study argues that H. antecessor, with a facial structure similar to that of H. sapiens,
belongs to the genus Homo, whereas the cranial structure of Neanderthals is derivative.[85]
H. cepranensis refers to a single skull cap from Italy, estimated to be about 800,000 years
old.[86] In 1994, researchers in Italy discovered skull fragments. After assembly, the distinct
shape separated them from other hominid species, leading researchers to believe it was
part of a previously unknown hominid species. They named it H. cepranensis sp. nov. after
the site it was found in (Campo Grande of Ceprano). Comparing the skull to other known
hominid species, they found that H. cepranensis sp. nov. bones differed in bone thickness,
length, and angles. This led to the conclusion that H. cepranensis sp. nov. is a distinct
hominid species.[87]

H. heidelbergensis
H. heidelbergensis ("Heidelberg Man") lived from about 800,000 to about 300,000 years ago. It is also
proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.[88]

In 1907, miners in Germany discovered the Mauer mandible, a very primitive fossilized jaw. An analysis
of the jaw by Otto Schoetensack prompted the attribution of this jaw to a new species: H. heidelbergensis.
The validity of this species as its own species and an ancestor to the Neandertals has been debated. In
2009, a study was conducted to reevaluate the Mauer mandible against other fossilized remains to
determine where on the human evolutionary scale H. heidelbergensis may lie or if it is even valid as a
separate species. With this reevaluation, researchers concluded that H. heidelbergensis was most likely an
Afro-European species that was a common ancestor to both
Neandertals and H. sapiens, though it is also possible that H.
heidelbergensis was a European species that was an ancestor
to only Neandertals, while another species from Africa was
the ancestor to H. sapiens.[89]

H. rhodesiensis, and the Gawis cranium

In 1921, a cranium was found in cave deposits
during a mining excursion in what is now known as
Zambia. H. rhodesiensis, estimated to be 300,000–
125,000 years old.[90] Most current researchers
place Rhodesian Man within the group of Homo Reconstruction of H. heidelbergensis
heidelbergensis, though other designations such as
archaic Homo sapiens and Homo sapiens
rhodesiensis have been proposed. This fossil has expanded the idea of the evolution of H.
sapiens; current estimates place the emergence of H. sapiens at around 500,000 years
ago.[91]
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 that it is either an intermediate species or an example of a
"Bodo man" female.[92]

Neanderthal and Denisovan

Homo neanderthalensis, alternatively designated as Homo sapiens
neanderthalensis,[93] lived in Europe and Asia from 400,000[94] to
about 28,000 years ago.[95] There are a number of clear anatomical
differences between anatomically modern humans (AMH) and
Neanderthal specimens, many relating to the superior Neanderthal
adaptation to cold environments. Neanderthal surface to volume ratio
was even lower than that among modern Inuit populations, indicating
superior retention of body heat.

Neanderthals also had significantly larger brains, as shown from brain

endocasts, casting doubt on their intellectual inferiority to modern
humans. However, the higher body mass of Neanderthals may have
required larger brain mass for body control.[96] Also, recent research by
Pearce, Stringer, and Dunbar has shown important differences in brain
architecture. The larger size of the Neanderthal orbital chamber and Reconstruction of an elderly
occipital lobe suggests that they had a better visual acuity than modern Neanderthal man
humans, useful in the dimmer light of glacial Europe.

Neanderthals may have had less brain capacity available for social functions. Inferring social group size
from endocranial volume (minus occipital lobe size) suggests that Neanderthal groups may have been
limited to 120 individuals, compared to 144 possible relationships for modern humans.[97] Larger social
groups could imply that modern humans had less risk of inbreeding within their clan, trade over larger
areas (confirmed in the distribution of stone tools), and faster spread of social and technological
innovations. All these may have contributed to modern Homo sapiens replacing Neanderthal populations
by 28,000 BP.[96]

Earlier evidence from sequencing mitochondrial DNA suggested 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.[98][99][100] However, a sequencing of the Neanderthal genome
in 2010 indicated that Neanderthals did indeed interbreed with anatomically modern humans c. 45,000-
80,000 years ago, around the time modern humans migrated out from Africa, but before they dispersed
throughout Europe, Asia and elsewhere.[101] The genetic sequencing of a 40,000-year-old human skeleton
from Romania showed that 11% of its genome was Neanderthal, implying the individual had a
Neanderthal ancestor 4–6 generations previously,[102] in addition to a contribution from earlier
interbreeding in the Middle East. Though this interbred Romanian population seems not to have been
ancestral to modern humans, the finding indicates that interbreeding happened repeatedly.[103]

All modern non-African humans have about 1% to 4% (or 1.5% to 2.6% by more recent data) of their
DNA derived from Neanderthals.[104][101][105] This finding is consistent with recent studies indicating
that the divergence of some human alleles dates to one Ma, although this interpretation has been
questioned.[106][107] Neanderthals and AMH Homo sapiens could have co-existed in Europe for as long
as 10,000 years, during which AMH populations exploded, vastly outnumbering Neanderthals, possibly
outcompeting them by sheer numbers.[108]

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 another human species, the
Denisovans.[109] 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.[50][110]

While the divergence point of the mtDNA was unexpectedly deep in time,[111] 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.[50] Modern humans are known to
have overlapped with Neanderthals in Europe and the Near East for possibly more than 40,000 years,[112]
and the discovery raises the possibility that Neanderthals, Denisovans, and modern humans may have co-
existed and interbred. The existence of this distant branch creates a much more complex picture of
humankind during the Late Pleistocene than previously thought.[110][113] 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.[114][115]

Alleles thought to have originated in Neanderthals and Denisovans have been identified at several genetic
loci in the genomes of modern humans outside Africa. Human leukocyte antigen (HLA) haplotypes from
Denisovans and Neanderthal represent more than half the HLA alleles of modern Eurasians,[52] indicating
strong positive selection for these introgressed alleles. Corinne Simoneti at Vanderbilt University, in
Nashville and her team have found from medical records of 28,000 people of European descent that the
presence of Neanderthal DNA segments may be associated with a higher rate of depression.[116]

The flow of genes from Neanderthal populations to modern humans was not all one way. Sergi Castellano
of the Max Planck Institute for Evolutionary Anthropology reported in 2016 that while Denisovan and
Neanderthal genomes are more related to each other than they are to us, Siberian Neanderthal genomes
show more similarity to modern human genes than do European Neanderthal populations. This suggests
Neanderthal populations interbred with modern humans around 100,000 years ago, probably somewhere
in the Near East.[117]

Studies of a Neanderthal child at Gibraltar show from brain development and tooth eruption that
Neanderthal children may have matured more rapidly than Homo sapiens.[118]

H. floresiensis
H. floresiensis, which lived from approximately 190,000 to
50,000 years before present (BP), has been nicknamed the
hobbit for its small size, possibly a result of insular
dwarfism.[119] 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
A facial reconstruction of Homo estimated to be one meter in height, with a brain volume of
floresiensis just 380 cm3 (considered small for a chimpanzee and less
than a third of the H. sapiens average of 1400 cm3).[119]

However, there is an ongoing debate over whether H. floresiensis is indeed a separate species.[120] Some
scientists hold that H. floresiensis was a modern H. sapiens with pathological dwarfism.[121] 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.[121]

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.[120]

In 2016, fossil teeth and a partial jaw from hominins assumed to be ancestral to H. floresiensis were
discovered[122] at Mata Menge, about 74 km (46 mi) from Liang Bua. They date to about 700,000 years
ago[123] and are noted by Australian archaeologist Gerrit van den Bergh for being even smaller than the
later fossils.[124]

H. luzonensis
A small number of specimens from the island of Luzon, dated 50,000 to 67,000 years ago, have recently
been assigned by their discoverers, based on dental characteristics, to a novel human species, H.
luzonensis.[125]

H. sapiens
H. sapiens (the adjective sapiens is Latin for "wise" or
"intelligent") emerged in Africa around 300,000 years ago,
likely derived from H. heidelbergensis or a related
lineage.[126][127] In September 2019, scientists reported the
computerized determination, based on 260 CT scans, of a
virtual skull shape of the last common human ancestor to
modern humans (H. sapiens), representative of the earliest
modern humans, and suggested that modern humans arose
between 260,000 and 350,000 years ago through a merging of
populations in East and South Africa.[128][129]

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 Reconstruction of early Homo sapiens
Africa) eventually replaced the earlier dispersed H. erectus. from Jebel Irhoud, Morocco
This migration and origin theory is usually referred to as the c. 315,000 years BP
"recent single-origin hypothesis" or "out of Africa" theory.
H. sapiens interbred with archaic humans both in Africa and
in Eurasia, in Eurasia notably with Neanderthals and Denisovans.[50][114]

The Toba catastrophe theory, which postulates a population bottleneck for H. sapiens about 70,000 years
ago,[130] was controversial from its first proposal in the 1990s and by the 2010s had very little
support.[131] Distinctive human genetic variability has arisen as the result of the founder effect, by archaic
admixture and by recent evolutionary pressures.

Anatomical changes
Since Homo sapiens separated from its last common ancestor shared with chimpanzees, human evolution
is characterized by a number of morphological, developmental, physiological, behavioral, and
environmental changes.[10] Environmental (cultural) evolution discovered much later during the
Pleistocene played a significant role in human evolution observed via human transitions between
subsistence systems.[132][10] 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.[133] Other significant morphological changes
included the evolution of a power and precision grip, a change first occurring in H. erectus.[134]
Bipedalism
Bipedalism (walking on two legs) is the basic adaptation of the
hominid and is considered the main cause behind a suite of skeletal
changes shared by all bipedal hominids. Fossil records regarding
bipedalism since the 1990s have placed the emergence of the earliest
hominins to the late Miocene era.[135] The earliest hominin, of
presumably primitive bipedalism, is considered to be either
Sahelanthropus[136] or Orrorin, both of which arose some 6 to
7 million years ago. The non-bipedal knuckle-walkers, the gorillas and
chimpanzees, diverged from the hominin line over a period covering
the same time, so either Sahelanthropus or Orrorin may be our last
shared ancestor. Ardipithecus, a full biped, arose approximately
5.6 million years ago.[137]

The early bipeds eventually evolved into the australopithecines and

still later into the genus Homo. There are several theories of the
adaptation value of bipedalism. It is possible that bipedalism was Bipedalism shown by a man
and a woman
favored because it freed the hands for reaching and carrying food,
saved energy during locomotion,[138] 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 and woodland
environment created as a result of the East African Rift Valley uplift versus the previous closed forest
habitat.[138][139][140] A 2007 study provides support for the hypothesis that bipedalism evolved because it
used less energy than quadrupedal knuckle-walking.[141][142] However, recent studies suggest that
bipedality without the ability to use fire would not have allowed global dispersal.[143] This change in gait
saw a lengthening of the legs proportionately when compared to the length of the arms, which were
shortened through the removal of the need for brachiation. Another change is the shape of the big toe.
Recent studies suggest that australopithecines still lived part of the time in trees as a result of maintaining
a grasping big toe. This was progressively lost in habilines.

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.[144] 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.[145]

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;[31]
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, thus permitting the passage of newborns due
to the increase in cranial size. This is limited to the upper portion, since further increase can hinder
normal bipedal movement.[146]
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.[147] The smaller birth canal became a 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.[140] The increased brain growth after birth and the
increased dependency of children on mothers had a major effect upon the female reproductive cycle,[148]
and the more frequent appearance of alloparenting in humans when compared with other hominids.[149]
Delayed human sexual maturity also led to the evolution of menopause with one explanation, the
grandmother hypothesis, providing that elderly women could better pass on their genes by taking care of
their daughter's offspring, as compared to having more children of their own.[150][151]

Encephalization

Skulls of successive (or near-successive, depending on the source) human evolutionary ancestors,[c] up until
'modern' Homo sapiens
* Mya – million years ago, kya – thousand years ago

The human species eventually developed

a much larger brain than that of other
primates—typically 1,330 cm3 (81 cu in)
in modern humans, nearly three times the
size of a chimpanzee or gorilla
brain.[154] After a period of stasis with
Australopithecus anamensis and
Ardipithecus, species which had smaller
brains as a result of their bipedal
locomotion,[155] the pattern of
encephalization started with Homo
habilis, whose 600 cm3 (37 cu in) brain
was slightly larger than that of
chimpanzees. This evolution continued in
Homo erectus with 800–1,100 cm3 (49– Brain size and tooth size in hominins

67 cu in), and reached a maximum in

Neanderthals with 1,200–1,900 cm3 (73–
116 cu in), larger even than modern Homo sapiens. This brain increase manifested during postnatal brain
growth, far exceeding that of other apes (heterochrony). It also allowed for extended periods of social
learning and language acquisition in juvenile humans, beginning as much as 2 million years ago.
Encephalization may be due to a dependency on calorie-dense, difficult-to-acquire food.[156]
Furthermore, the changes in the structure of human brains may be even more significant than the increase
in size.[157][158][159][58] Fossilized skulls shows the brain size in early humans fell within the range of
modern humans 300,000 years ago, but only got its present-day brain shape between 100,000 and 35,000
years ago.[160]

The temporal lobes, which contain centers for language

processing, have increased disproportionately, as has the
prefrontal cortex, which has been related to complex
decision-making and moderating social behavior.[154]
Encephalization has been tied to increased starches[57] and
meat[161][162] in the diet, however a 2022 meta study called
into question the role of meat.[163] Other factors are the
development of cooking,[164] 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.[165] Changes in skull morphology, such as smaller
The size and shape of the skull mandibles and mandible muscle attachments, allowed more
changed over time. The leftmost, and
room for the brain to grow.[166]
largest, is a replica of a modern human
skull.
The increase in volume of the neocortex also included a rapid
increase in size of the cerebellum. Its function has
traditionally been associated with balance and fine motor control, but more recently with speech and
cognition. The great apes, including hominids, had a more pronounced cerebellum relative to the
neocortex than other primates. It has been suggested that because of its function of sensory-motor control
and learning complex muscular actions, the cerebellum may have underpinned human technological
adaptations, including the preconditions of speech.[167][168][169][170]

The immediate survival advantage of encephalization is difficult to discern, as the major brain changes
from Homo erectus to Homo heidelbergensis were not accompanied by major changes in technology. It
has been suggested that the changes were mainly social and behavioural, including increased empathic
abilities,[171][172] increases in size of social groups,[165][173][174] and increased behavioral plasticity.[175]
Humans are unique in the ability to acquire information through social transmission and adapt that
information.[176] The emerging field of cultural evolution studies human sociocultural change from an
evolutionary perspective.[177]
 Evolution of the shape, size, and contours of the human (Homo)
skull[178][179][a][182][d][4][184][185][186][187][120][188][189]

Sexual dimorphism
The reduced degree of sexual dimorphism in humans 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 are the only hominoids 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 or overt
changes in proceptivity during estrus).[190]

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.[191] 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.[192]

Ulnar opposition
The ulnar opposition—the contact between the thumb and the tip of the little finger of the same hand—is
unique to the genus Homo,[193] including Neanderthals, the Sima de los Huesos hominins and
anatomically modern humans.[194][195] In other primates, the thumb is short and unable to touch the little
finger.[194] The ulnar opposition facilitates the precision grip and power grip of the human hand,
underlying all the skilled manipulations.
Other changes
A number of other changes have also characterized the
evolution of humans, among them an increased reliance on
vision rather than smell (highly reduced olfactory bulb); a
longer juvenile developmental period and higher infant
dependency;[196] a smaller gut and small, misaligned teeth;
faster basal metabolism;[197] loss of body hair;[198] an
increase in eccrine sweat gland density that is ten times
higher than any other catarrhinian primates,[199] yet humans
use 30% to 50% less water per day compared to chimps and
gorillas;[200] more REM sleep but less sleep in total;[201] a
change in the shape of the dental arcade from u-shaped to
parabolic; development of a chin (found in Homo sapiens
alone); styloid processes; and a descended larynx. As the
human hand and arms adapted to the making of tools and
were used less for climbing, the shoulder blades changed too.
As a side effect, it allowed human ancestors to throw objects
with greater force, speed and accuracy.[202]

Use of tools
Only the human is able to touch the
The use of tools has been interpreted as a sign of intelligence, little finger with the thumb.
and it has been theorized that tool use may have stimulated
certain aspects of human evolution, especially the continued
expansion of the human brain.[204] 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, on
average, about 13 watts (260 kilocalories per day), a fifth of
the body's resting power consumption.[205] 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
"A sharp rock", an Oldowan pebble
create and use tools.[206] tool, the most basic of human stone 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.[204] There is some evidence that the
australopithecines (4 Ma) may have used broken bones as tools, but this is debated.[207]

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 flakes from West Turkana, Kenya, which date to
3.3 million years ago.[208] The next oldest stone tools are from Gona, Ethiopia, and are considered the
beginning of the Oldowan technology. These tools date to about 2.6 million years ago.[209] 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.[210] 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 other apes and other nonhuman The harnessing of fire was a pivotal
primates, and is not seen in human fossils older than milestone in human history.
1.8 million years.[211]

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.[210]

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 Acheulean hand-axes from
hand can perform. Susman posited that modern anatomy of the human Kent. H. erectus flint work. The
opposable thumb is an evolutionary response to the requirements types shown are (clockwise
associated with making and handling tools and that both species were from top) cordate, ficron and
ovate.
indeed toolmakers.[210]

Transition to behavioral modernity

Anthropologists describe modern human behavior to include cultural and behavioral traits such as
specialization of tools, use of jewellery and images (such as cave drawings), organization of living space,
rituals (such as grave gifts), specialized hunting techniques, exploration of less hospitable geographical
areas, and barter trade networks, as well as more general traits such as language and complex symbolic
thinking. Debate continues as to whether a "revolution" led to modern humans ("big bang of human
consciousness"), or whether the evolution was more gradual.[54]

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) marked a new technology, followed by very slow
development until the next phase. Currently paleoanthropologists are debating whether these Homo
species possessed some or many modern human behaviors. They seem
to have been culturally conservative, maintaining the same
technologies and foraging patterns over very long periods.

Around 50,000 BP, human culture started to evolve more rapidly. The
transition to behavioral modernity has been characterized by some as a
"Great Leap Forward",[212] or as the "Upper Palaeolithic
Revolution",[213] due to the sudden appearance in the archaeological
record of distinctive signs of modern behavior and big game
hunting.[214] Evidence of behavioral modernity significantly earlier
also exists from Africa, with older evidence of abstract imagery,
widened subsistence strategies, more sophisticated tools and weapons,
and other "modern" behaviors, and many scholars have recently argued
that the transition to modernity occurred sooner than previously
believed.[54][215][216][217]
Venus of Willendorf, an
Other scholars consider the transition to have been more gradual,
example of Paleolithic art,
noting that some features had already appeared among archaic African
dated circa 30,000 years
Homo sapiens 300,000–200,000 years ago.[218][219][220][221][222] ago[203]
Recent evidence suggests that the Australian Aboriginal population
separated from the African population 75,000 years ago, and that they
made a 160 km (99 mi) sea journey 60,000 years ago, which may diminish the significance of the Upper
Paleolithic Revolution.[223]

Modern humans started burying their dead, making clothing from animal hides, hunting with more
sophisticated techniques (such as using pit traps or driving animals off cliffs), and cave painting.[224] As
human culture advanced, different populations innovated existing technologies: artifacts such as fish
hooks, buttons, and bone needles show signs of cultural variation, which had not been seen prior to
50,000 BP. Typically, the older H. neanderthalensis populations did not vary in their technologies,
although the Chatelperronian assemblages have been found to be Neanderthal imitations of H. sapiens
Aurignacian technologies.[225]

Recent and ongoing human evolution

Anatomically modern human populations continue to evolve, as they are affected by both natural
selection and genetic drift. Although selection pressure on some traits, such as resistance to smallpox, has
decreased in the modern age, humans are still undergoing natural selection for many other traits. Some of
these are due to specific environmental pressures, while others are related to lifestyle changes since the
development of agriculture (10,000 years ago), urbanization (5,000), and industrialization (250 years
ago). It has been argued that human evolution has accelerated since the development of agriculture
10,000 years ago and civilization some 5,000 years ago, resulting, it is claimed, in substantial genetic
differences between different current human populations,[226] and more recent research indicates that for
some traits, the developments and innovations of human culture have driven a new form of selection that
coexists with, and in some cases has largely replaced, natural selection.[227]
Particularly conspicuous is variation in superficial
characteristics, such as Afro-textured hair, or the recent
evolution of light skin and blond hair in some populations,
which are attributed to differences in climate. Particularly
strong selective pressures have resulted in high-altitude
adaptation in humans, with different ones in different isolated
populations. Studies of the genetic basis show that some
developed very recently, with Tibetans evolving over 3,000
years to have high proportions of an allele of EPAS1 that is
adaptive to high altitudes.

Other evolution is related to endemic diseases: the presence

of malaria selects for sickle cell trait (the heterozygous form
of sickle cell gene), while in the absence of malaria, the
health effects of sickle-cell anemia select against this trait.
For another 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
Reconstruction of the upper
due to the survival of immune persons.[229][230] Some Palaeolithic human Oase 2
reported trends remain unexplained and the subject of c. 40 000 years BP[228]
ongoing research in the novel field of evolutionary medicine:
polycystic ovary syndrome (PCOS) reduces fertility and thus
is expected to be subject to extremely strong negative selection, but its relative commonality in human
populations suggests a counteracting selection pressure. The identity of that pressure remains the subject
of some debate.[231]

Recent human evolution related to agriculture includes genetic resistance to infectious disease that has
appeared in human populations by crossing the species barrier from domesticated animals,[232] as well as
changes in metabolism due to changes in diet, such as lactase persistence.

Culturally-driven evolution can defy the expectations of natural selection: while human populations
experience some pressure that drives a selection for producing children at younger ages, the advent of
effective contraception, higher education, and changing social norms have driven the observed selection
in the opposite direction.[233] However, culturally-driven selection need not necessarily work counter or
in opposition to natural selection: some proposals to explain the high rate of recent human brain
expansion indicate a kind of feedback whereupon the brain's increased social learning efficiency
encourages cultural developments that in turn encourage more efficiency, which drive more complex
cultural developments that demand still-greater efficiency, and so forth.[234] Culturally-driven evolution
has an advantage in that in addition to the genetic effects, it can be observed also in the archaeological
record: the development of stone tools across the Palaeolithic period connects to culturally-driven
cognitive development in the form of skill acquisition supported by the culture and the development of
increasingly complex technologies and the cognitive ability to elaborate them.[235]

In contemporary times, since industrialization, some trends have been observed: for instance, menopause
is evolving to occur later.[236] 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.[236]

History of study

Before Darwin
The name Homo of the biological genus to which humans belong is Latin for 'human'.[e] It was chosen
originally by Carl Linnaeus in his classification system.[f] The English word human is from the Latin
humanus, the adjectival form of homo. The Latin homo derives from the Indo-European root *dhghem, or
'earth'.[237] 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.[238]

Darwin
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."[239]

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 other apes, and did so particularly in his 1863 book Evidence as to
Man's Place in Nature. 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 in his 1871 book The Descent of Man, and Selection in Relation to
Sex.[240]

First fossils
A major problem in the 19th century 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 the remains of a modern human who had suffered some kind of illness.[241] 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.[242] In 1925,
Raymond Dart described Australopithecus africanus.[243] 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 endocast 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
During the 1960s and 1970s, hundreds of fossils were found in East
Africa in the regions of the Olduvai Gorge and Lake Turkana. These
searches were carried out by the Leakey family, with Louis Leakey and
his wife Mary Leakey, and later their son Richard and daughter-in-law
Meave, fossil hunters and paleoanthropologists. From the fossil beds of
Olduvai and Lake Turkana they amassed specimens of the early
hominins: the australopithecines and Homo species, and even
H. erectus.

These finds cemented Africa as the cradle of humankind. In the late

Fossil hominid evolution
display at The Museum of
1970s and the 1980s, Ethiopia emerged as the new hot spot of
Osteology, Oklahoma City, paleoanthropology after "Lucy", the most complete fossil member of
Oklahoma, US 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.[244] 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.[245] (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.)[246] 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 A. kadabba.[247]

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.[248][249] As of September 2015, fossils of at least fifteen
individuals, amounting to 1,550 specimens, have been excavated from the cave.[249] 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 were dated close to 250,000 years ago,[250] and thus
are not ancestral to but contemporary with the first appearance of larger-brained anatomically modern
humans.[251]

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).[252] 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,[252] 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.[139][253][254] 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
chimpanzees 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
chimpanzees to an earlier period greater than 7 million years ago (Ma). Additional research with 226
offspring of wild chimpanzee populations in eight locations suggests that chimpanzees reproduce at age
26.5 years on average; which suggests the human divergence from chimpanzees occurred between 7 and
13 mya. And these data suggest that Ardipithecus (4.5 Ma), Orrorin (6 Ma) and Sahelanthropus (7 Ma)
all may be on the hominid 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-
chimpanzees" nonetheless occurred regularly enough to change certain genes in the new gene pool:

A new comparison of the human and chimpanzee genomes suggests that after the two
lineages separated, they may have begun interbreeding... A principal finding is that the X
chromosomes of humans and chimpanzees 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 chimpanzee 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.[255]

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 4.2 million years ago.

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 7.2 million years ago, and which
Brunet argued was a bipedal, and therefore a hominid—that is, a hominin (cf Hominidae; terms
"hominids" and hominins).

Human dispersal

A global mapping model of human migration, based A "trellis" (as Milford H. Wolpoff called
from divergence of the mitochondrial DNA (which it) that emphasizes back-and-forth
indicates the matrilineage).[256][257][258] Timescale gene flow among geographic
(ka) indicated by colours. regions[259]
Different models for the beginning of the present human species

Anthropologists in the 1980s were divided regarding some details of reproductive barriers and migratory
dispersals of the genus Homo. Subsequently, genetics has been used to investigate and resolve these
issues. According to the Sahara pump theory evidence suggests that the genus Homo have migrated out of
Africa at least three and possibly four times (e.g. Homo erectus, Homo heidelbergensis and two or three
times for Homo sapiens). Recent evidence suggests these dispersals are closely related to fluctuating
periods of climate change.[260]

Recent evidence suggests that humans may have left Africa half a million years earlier than previously
thought. A joint Franco-Indian team has found human artifacts in the Siwalk Hills north of New Delhi
dating back at least 2.6 million years. This is earlier than the previous earliest finding of genus Homo at
Dmanisi, in Georgia, dating to 1.85 million years. Although controversial, tools found at a Chinese cave
strengthen the case that humans used tools as far back as 2.48 million years ago.[261] This suggests that
the Asian "Chopper" tool tradition, found in Java and northern China may have left Africa before the
appearance of the Acheulian hand axe.

Dispersal of modern Homo sapiens

Up until the genetic evidence became available, there were two dominant models for the dispersal of
modern humans. The multiregional hypothesis proposed that the genus Homo 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 of million years. This model was proposed in 1988 by
Milford H. Wolpoff.[262][263] In contrast, 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 the nearly complete replacement of other Homo species. This model has been
developed by Chris Stringer and Peter Andrews.[264][265]
Sequencing mtDNA and Y-DNA sampled from a wide range of indigenous populations revealed ancestral
information relating to both male and female genetic heritage, and strengthened the "out of Africa" theory
and weakened the views of multiregional evolutionism.[266] Aligned in genetic tree differences were
interpreted as supportive of a recent single origin.[267]

"Out of Africa" has thus gained much 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.[268]

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 a possible origin of modern human migration in
southwestern Africa, near the coastal border of Namibia and Angola.[269] The fossil evidence was
insufficient for archaeologist Richard Leakey to resolve the debate about exactly where in Africa modern
humans first appeared.[270] Studies of haplogroups in Y-chromosomal DNA and mitochondrial DNA have
largely supported a recent African origin.[271] All the evidence from autosomal DNA also predominantly
supports a Recent African origin. However, evidence for archaic admixture in modern humans, both in
Africa and later, throughout Eurasia has recently been suggested by a number of studies.[272]

Recent sequencing of Neanderthal[104] and Denisovan[50] genomes shows that some admixture with these
populations has occurred. All modern human groups outside Africa have 1–4% or (according to more
recent research) about 1.5–2.6% Neanderthal alleles in their genome,[105] 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, although they make the situation more complex. After recovery from
a genetic bottleneck that some researchers speculate might be linked to the Toba supervolcano
catastrophe, a fairly small group left Africa and interbred with Neanderthals, probably in the Middle East,
on the Eurasian steppe or even in North Africa before their departure. Their still predominantly African
descendants spread to populate the world. A fraction in turn interbred with Denisovans, probably in
southeastern Asia, before populating Melanesia.[114] HLA haplotypes of Neanderthal and Denisova origin
have been identified in modern Eurasian and Oceanian populations.[52] The Denisovan EPAS1 gene has
also been found in Tibetan populations.[273] Studies of the human genome using machine learning have
identified additional genetic contributions in Eurasians from an "unknown" ancestral population
potentially related to the Neanderthal-Denisovan lineage.[274]

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,[275][276][277] 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 crossing the Bab el Mandib to A map of early human migrations
Yemen at a lower sea level 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.[275] This group seems to have been
dependent upon marine resources for their survival.

Stephen Oppenheimer has proposed a second wave of humans may have later dispersed through the
Persian Gulf oases, and the Zagros mountains into the Middle East. Alternatively it may have come
across the Sinai Peninsula into Asia, from shortly after 50,000 yrs BP, resulting in the bulk of the human
populations of Eurasia. It has been suggested that this second group possibly possessed a more
sophisticated "big game hunting" 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.[275] 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 L3 lineages, which support a single migration out of
Africa that gave rise to all non-African populations.[278]

On the basis of the early date of Badoshan Iranian Aurignacian, Oppenheimer suggests that this second
dispersal may have occurred with a pluvial period about 50,000 years before the present, with modern
human big-game hunting cultures spreading up the Zagros Mountains, carrying modern human genomes
from Oman, throughout the Persian Gulf, northward into Armenia and Anatolia, with a variant travelling
south into Israel and to Cyrenicia.[214]

Recent genetic evidence suggests that all modern non-African populations, including those of Eurasia and
Oceania, are descended from a single wave that left Africa between 65,000 and 50,000 years
ago.[279][280][281]

Evidence
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.[282]

Evidence from genetics

The closest living relatives of humans are bonobos and chimpanzees (both genus Pan) and gorillas (genus
Gorilla).[283] With the sequencing of both the human and chimpanzee genome, as of 2012 estimates of
the similarity between their DNA sequences range between 95% and 99%.[283][284][34] It is also
noteworthy that mice share around 97.5% of their working DNA with humans.[285] 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 the orangutans (genus Pongo) were
the first groups to split from the line
leading to the hominins, including
humans—followed by gorillas (genus
Gorilla), and, ultimately, by the
chimpanzees (genus Pan). The splitting
Family tree showing the extant hominoids: humans (genus
date between hominin and chimpanzee
Homo), chimpanzees and bonobos (genus Pan), gorillas
lineages is placed by some between
(genus Gorilla), orangutans (genus Pongo), and gibbons (four
4 to 8 million years ago, that is, during genera of the family Hylobatidae: Hylobates, Hoolock,
the .[286][287][288][289]
Late Miocene Nomascus, and Symphalangus). All except gibbons are
Speciation, however, appears to have hominids.
been unusually drawn out. Initial
divergence occurred sometime between
7 to 13 million years ago, but ongoing hybridization blurred the separation and delayed complete
separation during several millions of years. Patterson (2006) dated the final divergence at
5 to 6 million years ago.[290]

Genetic evidence has also been employed to compare species within the genus Homo, investigating gene
flow between early modern humans and Neanderthals, and to enhance the 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 human genomes differ among individuals, the evolutionary
past that gave rise to them, and their current effects. Differences between genomes have anthropological,
medical and forensic implications and applications. Genetic data can provide important insight into
human evolution. Polygenic scores from ancient DNA showed directional selection for various traits
including evolution of human intelligence in some periods.[291]

In May 2023, scientists reported a more complicated pathway of human evolution than previously
understood. According to the studies, humans evolved from different places and times in Africa, instead
of from a single location and period of time.[292][293]

Evidence from the fossil record

There is little fossil evidence for the divergence of the gorilla, chimpanzee and hominin lineages.[294] The
earliest fossils that have been proposed as members of the hominin lineage are Sahelanthropus tchadensis
dating from 7 million years ago, Orrorin tugenensis dating from 5.7 million years ago, and Ardipithecus
kadabba dating to 5.6 million years ago. 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 4 million years ago and
diverged into 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 100 fossil individuals represented, found from Northern Ethiopia
Replica of fossil skull of
(such as the famous "Lucy"), to Kenya, and South Africa. Fossils of
H. habilis. Fossil number KNM
robust australopithecines such as A. robustus (or alternatively ER 1813, found at Koobi Fora,
Paranthropus robustus) and A./P. boisei are particularly abundant in Kenya.
South Africa at sites such as Kromdraai and Swartkrans, and around
Lake Turkana in Kenya.

The earliest member of the genus Homo is Homo habilis which

evolved around 2.8 million years ago.[40] H. 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 Replica of fossil skull of
as an adaptation to terrestrial living. H. ergaster (African
H. erectus). Fossil number
During the next million years, a process of encephalization began and, Khm-Heu 3733 discovered in
by the arrival (about 1.9 million years ago) of H. erectus in the fossil 1975 in Kenya.

record, cranial capacity had doubled. H. erectus were the first of the
hominins to emigrate from Africa, and, from
1.8 to 1.3 million years ago, this species spread through Africa, Asia, and Europe. One population of
H. erectus, also sometimes classified as separate species H. ergaster, remained in Africa and evolved into
H. sapiens. It is believed that H. erectus and H. ergaster were the first to use fire and complex tools. In
Eurasia, H. erectus evolved into species such as H. antecessor, H. heidelbergensis and
H. neanderthalensis. The earliest fossils of anatomically modern humans are from the Middle Paleolithic,
about 300–200,000 years ago such as the Herto and Omo remains of Ethiopia, Jebel Irhoud remains of
Morocco, and Florisbad remains of South Africa; later fossils from the Skhul Cave in Israel and Southern
Europe begin around 90,000 years ago (0.09 million years ago).

As modern humans spread out from Africa, they encountered other hominins such as H. neanderthalensis
and the Denisovans, who may have evolved from populations of H. erectus that had left Africa around
2 million years ago. 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.[295][296]

This migration out of Africa is estimated to have begun about 70–50,000 years BP 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.[297]

Inter-species breeding
The hypothesis of interbreeding, also known as
hybridization, admixture or hybrid-origin
theory, has been discussed ever since the
discovery of Neanderthal remains in the 19th
century.[298] The linear view of human
evolution began to be abandoned in the 1970s
as different species of humans were discovered
that made the linear concept increasingly
unlikely. In the 21st century with the advent of
molecular biology techniques and
computerization, whole-genome sequencing of
Neanderthal and human genome were
performed, confirming recent admixture A model of the evolution of the genus Homo over the
between different human species.[104] In 2010, last 2 million years (vertical axis). The rapid "Out of
evidence based on molecular biology was Africa" expansion of H. sapiens is indicated at the top of
published, revealing unambiguous examples of the diagram, with admixture indicated with Neanderthals,
Denisovans, and unspecified archaic African hominins.
interbreeding between archaic and modern
humans during the Middle Paleolithic and early
Upper Paleolithic. It has been demonstrated that interbreeding happened in several independent events
that included Neanderthals and Denisovans, as well as several unidentified hominins.[299] Today,
approximately 2% of DNA from all non-African populations (including Europeans, Asians, and
Oceanians) is Neanderthal,[104] with traces of Denisovan heritage.[300] Also, 4–6% of modern Melanesian
genetics are Denisovan.[300] Comparisons of the human genome to the genomes of Neanderthals,
Denisovans and apes can help identify features that set modern humans apart from other hominin species.
In a 2016 comparative genomics study, a Harvard Medical School/UCLA research team made a world
map on the distribution and made some predictions about where Denisovan and Neanderthal genes may
be impacting modern human biology.[301][302]

For example, comparative studies in the mid-2010s found several traits related to neurological,
immunological,[303] developmental, and metabolic phenotypes, that were developed by archaic humans
to European and Asian environments and inherited to modern humans through admixture with local
hominins.[304][305]

Although the narratives of human evolution are often contentious, several discoveries since 2010 show
that human evolution should not be seen as a simple linear or branched progression, but a mix of related
species.[50][6][7][8] In fact, genomic research has shown that hybridization between substantially diverged
lineages is the rule, not the exception, in human evolution.[5] Furthermore, it is argued that hybridization
was an essential creative force in the emergence of modern humans.[5]

Stone tools
Stone tools are first attested around 2.6 million years ago, when hominins in Eastern Africa used so-called
core tools, choppers made out of round cores that had been split by simple strikes.[306] 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 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.[208][307][308][309]

The period from 700,000 to 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.[306] 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). Bone tools were also made by H. sapiens in Africa by 90,000–70,000 years
ago[215][310] and are also known from early H. sapiens sites in Eurasia by about 50,000 years ago.

Species list
This list is in chronological order across the table by genus. Some species/subspecies names are well-
established, and some are less established – especially in genus Homo. Please see articles for more
information.
 Sahelanthropus Homo (humans)

S. tchadensis H. gautengensis

Orrorin H. habilis
O. tugenensis H. rudolfensis

Ardipithecus H. floresiensis

A. kadabba H. ergaster
A. ramidus H. erectus

Australopithecus • H. e. georgicus

A. anamensis H. cepranensis
A. afarensis H. antecessor

A. bahrelghazali H. heidelbergensis

A. africanus H. rhodesiensis
A. garhi H. naledi

A. sediba H. helmei

Kenyanthropus H. neanderthalensis
K. platyops H. sapiens

Paranthropus • H. s. idaltu
P. aethiopicus • H. s. sapiens (early)

P. boisei • H. s. sapiens (modern)

P. robustus

See also
Adaptive evolution in the human genome Human behavioral ecology
Amity–enmity complex Human origins
Anthropology Human vestigiality
Archaeogenetics List of human evolution fossils
Biological Anthropology Molecular paleontology
Dental anthropology Obstetrical dilemma
Dual inheritance theory Origin of language
Evolution of morality Origin of speech
Evolutionary medicine Prehistory of nakedness and clothing
Evolutionary neuroscience Sexual selection in humans
Evolutionary origin of religion Transgenerational trauma
Evolutionary psychology Timeline of human evolution

Notes
a. The conventional estimate on the age of H. habilis is at roughly 2.1 to 2.3 million
years.[42][180] Suggestions for pushing back the age to 2.8 Mya were made in 2015 based
 on the discovery of a jawbone.[181]
b. Not to be confused with Pongidae, an obsolete family which grouped together orangutans,
gorillas and chimpanzees to separate them from humans
c. There is no general agreement on the line of special descent of H. sapiens from H. erectus.
Some of the species depicted in the image may not actually represent a direct evolutionary
ancestor to H. sapiens, and may not directly derive from one another, namely:

H. heidelbergensis likely did not descend from H. antecessor.[152]

H. heidelbergensis is likely not an ancestor to H. sapiens, nor is H. antecessor.[152]
H. ergaster is often considered the next evolutionary ancestor to H. sapiens following H.
erectus, however, there is considerable uncertainty as to the accuracy of classifying it as
a separate species from H. erectus at all.[153]
d. H. erectus in the narrow sense (the Asian species) was extinct by 140,000 years ago, Homo
erectus soloensis, found in Java, is considered the latest known survival of H. erectus.
Formerly dated to as late as 50,000 to 40,000 years ago, a 2011 study pushed back the
date of its extinction of H. e. soloensis to 143,000 years ago at the latest, more likely before
550,000 years ago.[183]
e. The Latin word which refers to adult males only is vir
f. See the Binomial nomenclature and Systema Naturae articles.

References
1. Based on Schlebusch, C. M.; Malmström, H.; Günther, T.; Sjödin, P.; Coutinho, A.; Edlund,
H.; Munters, A. R.; Vicente, M.; Steyn, M.; Soodyall, H.; Lombard, M.; Jakobsson, M. (2017).
"Southern African ancient genomes estimate modern human divergence to 350,000 to
260,000 years ago" ([Link] Science. 358 (6363):
652–655. Bibcode:2017Sci...358..652S ([Link]
2S). doi:10.1126/science.aao6266 ([Link]
PMID 28971970 ([Link] S2CID 206663925 ([Link]
[Link]/CorpusID:206663925)., Fig. 3 ([Link]
ntent/sci/early/2017/09/27/science.aao6266/[Link]) Archived ([Link]
web/20180114130711/[Link]
nce.aao6266/[Link]) January 14, 2018, at the Wayback Machine (H. sapiens
divergence times) and Stringer, C. (2012). "What makes a modern human" ([Link]
0.1038%2F485033a). Nature. 485 (7396): 33–35. Bibcode:2012Natur.485...33S ([Link]
[Link]/abs/2012Natur.485...33S). doi:10.1038/485033a ([Link]
8%2F485033a). PMID 22552077 ([Link]
S2CID 4420496 ([Link] (archaic admixture).
2. "Human evolution" ([Link] Encyclopædia
Britannica. June 8, 2024.
3. Hall, Brian K.; Hallgrímsson, Benedikt (2011). Strickberger's Evolution ([Link]
com/books?id=CrJsNQ5wX8oC&pg=PA488). Jones & Bartlett. p. 488. ISBN 978-1-4496-
6390-2.
4. Mondal, M.; Bertranpetit, J.; Lao, O. (January 2019). "Approximate Bayesian computation
with deep learning supports a third archaic introgression in Asia and Oceania" ([Link]
[Link]/pmc/articles/PMC6335398). Nature Communications. 10 (1): 246.
Bibcode:2019NatCo..10..246M ([Link]
doi:10.1038/s41467-018-08089-7 ([Link]
PMC 6335398 ([Link] PMID 30651539
([Link]
 5. Rogers Ackermann, Rebecca; Mackay, Alex; Arnold, Michael L. (October 2015). "The Hybrid
Origin of "Modern" Humans". Evolutionary Biology. 43 (1): 1–11. doi:10.1007/s11692-015-
9348-1 ([Link] S2CID 14329491 ([Link]
[Link]/CorpusID:14329491).
6. Antrosio, Jason (August 23, 2018). "Denisovans and Neandertals: Rethinking Species
Boundaries" ([Link]
andertals-human-races/). Living Anthropologically. Archived ([Link]
200801180554/[Link]
neandertals-human-races/) from the original on August 1, 2020. Retrieved August 25, 2018.
7. Hammer, Michael F. (May 2013). "Human Hybrids" ([Link]
34550/[Link] (PDF). Scientific
American. Archived from the original ([Link]
[Link]) (PDF) on August 24, 2018.
8. Yong, Ed (July 2011). "Mosaic humans, the hybrid species". New Scientist. Vol. 211,
no. 2823. pp. 34–38. Bibcode:2011NewSc.211...34Y ([Link]
1NewSc.211...34Y). doi:10.1016/S0262-4079(11)61839-3 ([Link]
2-4079%2811%2961839-3). ISSN 0262-4079 ([Link]
9. Heng, Henry H. Q. (May 2009). "The genome-centric concept: Resynthesis of evolutionary
theory". BioEssays. 31 (5): 512–525. doi:10.1002/bies.200800182 ([Link]
2Fbies.200800182). ISSN 0265-9247 ([Link]
PMID 19334004 ([Link] S2CID 1336952 ([Link]
[Link]/CorpusID:1336952).
10. Marlowe, Frank W. (April 13, 2005). "Hunter-gatherers and human evolution". Evolutionary
Anthropology: Issues, News, and Reviews. 14 (2): 54–67. doi:10.1002/evan.20046 ([Link]
[Link]/10.1002%2Fevan.20046). S2CID 53489209 ([Link]
D:53489209).
11. Tyson, Peter (July 1, 2008). "Meet Your Ancestors" ([Link]
n/[Link]). Nova ScienceNow. WGBH Educational Foundation. Archived
([Link]
[Link]) from the original on March 8, 2021. Retrieved April 18, 2015.
12. Gibbons, Ann (June 13, 2012). "Bonobos Join Chimps as Closest Human Relatives" (https://
[Link]/content/article/bonobos-join-chimps-closest-human-relatives-rev2).
TimeTree. Archived ([Link]
content/article/bonobos-join-chimps-closest-human-relatives-rev2) from the original on
September 13, 2021. Retrieved May 19, 2018.
13. Maxwell 1984, p. 296.
14. Zhang, Rui; Wang, Yin-Qiu; Su, Bing (July 2008). "Molecular Evolution of a Primate-Specific
microRNA Family" ([Link] Molecular Biology and
Evolution. 25 (7): 1493–1502. doi:10.1093/molbev/msn094 ([Link]
ev%2Fmsn094). ISSN 0737-4038 ([Link]
PMID 18417486 ([Link]
15. Willoughby, Pamela R. (2005). "Palaeoanthropology and the Evolutionary Place of Humans
in Nature" ([Link] International Journal of Comparative
Psychology. 18 (1): 60–91. doi:10.46867/IJCP.2005.18.01.02 ([Link]
JCP.2005.18.01.02). ISSN 0889-3667 ([Link]
Archived ([Link]
669xb) from the original on January 17, 2012. Retrieved April 27, 2015.
16. Martin 2001, pp. 12032–12038.
17. 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. 416
(6882): 726–729. Bibcode:2002Natur.416..726T ([Link]
ur.416..726T). doi:10.1038/416726a ([Link] ISSN 0028-0836
([Link] PMID 11961552 ([Link]
gov/11961552). S2CID 4368374 ([Link]
18. Rose, Kenneth D. (1994). "The earliest primates". Evolutionary Anthropology: Issues, News,
and Reviews. 3 (5): 159–173. doi:10.1002/evan.1360030505 ([Link]
an.1360030505). ISSN 1060-1538 ([Link]
S2CID 85035753 ([Link]
19. Fleagle, John; Gilbert, Chris (2011). Rowe, Noel; Myers, Marc (eds.). "Primate Evolution" (ht
tp://[Link]/John_Fleagle_Public.aspx). All The World's Primates.
Charlestown, Rhode Island: Primate Conservation. Archived ([Link]
150512013618/[Link] from the
original on May 12, 2015. Retrieved April 27, 2015.
20. Roach, John (March 3, 2008). "Oldest Primate Fossil in North America Discovered" (https://
[Link]/web/20121016193314/[Link]
[Link]). National Geographic News. Washington, DC: National
Geographic Society. Archived from the original ([Link]
08/03/[Link]) on October 16, 2012. Retrieved April 27, 2015.
21. McMains, Vanessa (December 5, 2011). "Found in Wyoming: New fossils of oldest American
primate" ([Link]
n-primate/). The Gazette. Baltimore: Johns Hopkins University. Archived ([Link]
[Link]/web/20190116200750/[Link]
s-of-oldest-american-primate/) from the original on January 16, 2019. Retrieved April 27,
2015.
22. Caldwell, Sara B. (May 19, 2009). "Missing link found, early primate fossil 47 million years
old" ([Link] Digital Journal. Toronto. Archived (https://
[Link]/web/20150722061001/[Link] from
the original on July 22, 2015. Retrieved April 27, 2015.
23. Watts, Alex (May 20, 2009). "Scientists Unveil Missing Link In Evolution" ([Link]
[Link]/web/20110728005247/[Link] Sky
News Online. London: BSkyB. Archived from the original ([Link]
ews/article/15284582) on July 28, 2011. Retrieved April 27, 2015.
24. Wilford, J. N. (June 5, 2013). "Palm-size fossil resets primates' clock, scientists say" (https://
[Link]/archive/20220101/[Link]
[Link]). The New York Times. Archived from the
original ([Link]
-[Link]) on January 1, 2022. Retrieved June 5, 2013.
25. Williams, B. A.; Kay, R. F.; Kirk, E. C. (2010). "New perspectives on anthropoid origins" (http
s://[Link]/pmc/articles/PMC2841917). Proceedings of the National Academy
of Sciences. 107 (11): 4797–4804. Bibcode:2010PNAS..107.4797W ([Link]
[Link]/abs/2010PNAS..107.4797W). doi:10.1073/pnas.0908320107 ([Link]
3%2Fpnas.0908320107). PMC 2841917 ([Link]
41917). PMID 20212104 ([Link]
26. Tabuce, R.; Marivaux, L.; Lebrun, R.; Adaci, M.; Bensalah, M.; Fabre, P. -H.; Fara, E.;
Gomes Rodrigues, H.; Hautier, L.; Jaeger, J. -J.; Lazzari, V.; Mebrouk, F.; Peigne, S.; Sudre,
J.; Tafforeau, P.; Valentin, X.; Mahboubi, M. (2009). "Anthropoid versus strepsirhine status of
the African Eocene primates Algeripithecus and Azibius: Craniodental evidence" ([Link]
[Link]/pmc/articles/PMC2821352). Proceedings of the Royal Society B:
Biological Sciences. 276 (1676): 4087–4094. Bibcode:2009PBioS.276.4087T ([Link]
[Link]/abs/2009PBioS.276.4087T). doi:10.1098/rspb.2009.1339 ([Link]
0.1098%2Frspb.2009.1339). PMC 2821352 ([Link]
2821352). PMID 19740889 ([Link]

Lay summary in: "African Origin Of Anthropoid Primates Called Into Question With
New Fossil Discovery" ([Link]
[Link]). Science Daily. September 15, 2009.
27. Kordos, László; Begun, David R. (January 2001). "Primates from Rudabánya: allocation of
specimens to individuals, sex and age categories". Journal of Human Evolution. 40 (1).
Amsterdam, the Netherlands: Elsevier: 17–39. Bibcode:2001JHumE..40...17K ([Link]
[Link]/abs/2001JHumE..40...17K). doi:10.1006/jhev.2000.0437 ([Link]
1006%2Fjhev.2000.0437). PMID 11139358 ([Link]
28. Cameron 2004, p. 76 ([Link]
29. Wallace 2004, p. 240 ([Link]
30. 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. 466 (7304): 360–364. Bibcode:2010Natur.466..360Z ([Link]
abs/2010Natur.466..360Z). doi:10.1038/nature09094 ([Link]
94). ISSN 0028-0836 ([Link] PMID 20631798 (https://
[Link]/20631798). S2CID 205220837 ([Link]
rpusID:205220837).
31. Srivastava 2009, p. 87 ([Link]
32. "Scientists observe chimpanzees using human-like warfare tactic" ([Link]
science/scientists-observe-chimpanzees-using-human-like-warfare-tactic-2023-11-02/).
Reuters. November 3, 2023.
33. Clark, G.; Henneberg, M. (June 2015). "The life history of Ardipithecus ramidus: A
heterochronic model of sexual and social maturation" ([Link]
-0009). Anthropological Review. 78 (2): 109–132. doi:10.1515/anre-2015-0009 ([Link]
g/10.1515%2Fanre-2015-0009). S2CID 54900467 ([Link]
D:54900467).
34. Sayers, Ken; Raghanti, Mary Ann; Lovejoy, C. Owen (October 2012). "Human Evolution and
the Chimpanzee Referential Doctrine". Annual Review of Anthropology. 41: 119–138.
doi:10.1146/annurev-anthro-092611-145815 ([Link]
611-145815). ISSN 0084-6570 ([Link]
35. Zimmer, Carl (May 27, 2015). "The Human Family Tree Bristles With New Branches" (http
s://[Link]/archive/20220101/[Link]
[Link]). The New York Times. Archived from the original (h
ttps://[Link]/2015/06/02/science/[Link]
l) on January 1, 2022. Retrieved May 30, 2015.
36. Gardner., Elizabeth K.; Purdue University (April 1, 2015). "New instrument dates old
skeleton before 'Lucy'; 'Little Foot' 3.67 million years old". Science Daily. Retrieved April 3,
2015.
37. Andrea Elyse Messer (March 4, 2015). "Earliest known fossil of the genus Homo dates to
2.8 to 2.75 million years ago" ([Link]
l-genus-homo-dates-28-275-million-years-ago). Penn State.
38. Smithsonian Institution. "The oldest fossil of our genus" ([Link]
h/whats-hot-human-origins/oldest-fossil-our-genus). [Link].
39. Brandon Bryn. "Science: Oldest Fossil of Homo Genus Found in Ethiopia" ([Link]
[Link]/news/science-oldest-fossil-homo-genus-found-ethiopia). AAAS.
40. Ghosh, Pallab (March 4, 2015). " 'First human' discovered in Ethiopia" ([Link]
m/news/science-environment-31718336). BBC News. London. Archived ([Link]
[Link]/web/20150418032919/[Link]
from the original on April 18, 2015. Retrieved April 19, 2015.
41. Swisher, Curtis & Lewin 2001.
42. Stringer 1994, p. 242.
43. McHenry 2009, p. 265.
44. "Out of Africa Revisited" ([Link] Science
(This Week in Science). 308 (5724): 921. May 13, 2005.
doi:10.1126/[Link] ([Link]
[Link]). ISSN 0036-8075 ([Link] S2CID 220100436 (h
ttps://[Link]/CorpusID:220100436).
45. Stringer, Chris (June 12, 2003). "Human evolution: Out of Ethiopia". Nature. 423 (6941):
692–695. Bibcode:2003Natur.423..692S ([Link]
692S). doi:10.1038/423692a ([Link] ISSN 0028-0836 (http
s://[Link]/issn/0028-0836). PMID 12802315 ([Link]
12802315). S2CID 26693109 ([Link]
46. Johanson, Donald (May 2001). "Origins of Modern Humans: Multiregional or Out of Africa?"
([Link]
[Link]). actionbioscience. Washington, DC: American Institute of Biological Sciences.
Archived from the original ([Link] on
November 14, 2010. Retrieved November 23, 2009.
47. Mixon, Bobbie; Ehardt, Carolyn; Hammer, Michael (September 6, 2011). "Evolution's Past Is
Modern Human's Present" ([Link]
21603&preview=false) (Press release). National Science Foundation. Press Release 11-
181. Archived ([Link]
_summ.jsp?org=NSF&cntn_id=121603&preview=false) from the original on December 17,
2014. Retrieved April 20, 2015.
48. O'Neil, Dennis. "Early Modern Homo sapiens" ([Link]
7/[Link] Evolution of Modern Humans: A
Survey of the Biological and Cultural Evolution of Archaic and Modern Homo sapiens
(Tutorial). San Marcos, California: Palomar College. Archived from the original ([Link]
[Link]/homo2/mod_homo_4.htm) on April 30, 2015. Retrieved April 20, 2015.
49. "Fossil Reanalysis Pushes Back Origin of Homo sapiens" ([Link]
m/article/fossil-reanalysis-pushes/). Scientific American. February 17, 2005. ISSN 0036-
8733 ([Link] Archived ([Link]
160115000603/[Link] from the
original on January 15, 2016. Retrieved April 20, 2015.
50. Reich, David; Green, Richard E.; Kircher, Martin; et al. (December 23, 2010). "Genetic
history of an archaic hominin group from Denisova Cave in Siberia" ([Link]
[Link]/pmc/articles/PMC4306417). Nature. 468 (7327): 1053–1060.
Bibcode:2010Natur.468.1053R ([Link]
doi:10.1038/nature09710 ([Link] hdl:10230/25596 (http
s://[Link]/10230%2F25596). ISSN 0028-0836 ([Link]
-0836). PMC 4306417 ([Link]
PMID 21179161 ([Link]
51. Noonan, James P. (May 2010). "Neanderthal genomics and the evolution of modern
humans" ([Link] Genome Research. 20
(5): 547–553. doi:10.1101/gr.076000.108 ([Link]
ISSN 1088-9051 ([Link] PMC 2860157 ([Link]
[Link]/pmc/articles/PMC2860157). PMID 20439435 ([Link]
ov/20439435).
52. 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" ([Link] Science. 334 (6052):
89–94. Bibcode:2011Sci...334...89A ([Link]
doi:10.1126/science.1209202 ([Link] ISSN 0036-
8075 ([Link] PMC 3677943 ([Link]
gov/pmc/articles/PMC3677943). PMID 21868630 ([Link]
0).
53. Mellars, Paul (June 20, 2006). "Why did modern human populations disperse from Africa ca.
60,000 years ago? A new model" ([Link]
Proceedings of the National Academy of Sciences of the United States of America. 103 (25):
9381–9386. Bibcode:2006PNAS..103.9381M ([Link]
S..103.9381M). doi:10.1073/pnas.0510792103 ([Link]
3). ISSN 0027-8424 ([Link] PMC 1480416 ([Link]
[Link]/pmc/articles/PMC1480416). PMID 16772383 ([Link]
[Link]/16772383).
54. 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. 39 (5):
453–563. Bibcode:2000JHumE..39..453M ([Link]
39..453M). doi:10.1006/jhev.2000.0435 ([Link]
ISSN 0047-2484 ([Link] PMID 11102266 ([Link]
[Link]/11102266). S2CID 42968840 ([Link]
D:42968840).
55. Strait, David S.; Grine, Frederick E.; Moniz, Marc A. (January 1997). "A reappraisal of early
hominid phylogeny". Journal of Human Evolution. 32 (1): 17–82.
Bibcode:1997JHumE..32...17S ([Link]
doi:10.1006/jhev.1996.0097 ([Link] ISSN 0047-2484
([Link] PMID 9034954 ([Link]
ov/9034954). S2CID 37754799 ([Link]
56. Bryson 2004, pp. 522–543.
57. Zimmer, Carl (August 13, 2015). "For Evolving Brains, a 'Paleo' Diet Full of Carbs" ([Link]
[Link]/archive/20220101/[Link]
-[Link]). The New York Times. Archived from the original (http
s://[Link]/2015/08/13/science/[Link])
on January 1, 2022. Retrieved August 14, 2015.
58. Leonard, William R.; Snodgrass, J. Josh; Robertson, Marcia L. (August 2007). "Effects of
brain evolution on human nutrition and metabolism". Annual Review of Nutrition. 27: 311–
327. doi:10.1146/[Link].27.061406.093659 ([Link]
7.061406.093659). ISSN 0199-9885 ([Link]
PMID 17439362 ([Link] S2CID 18869516 ([Link]
[Link]/CorpusID:18869516).
59. Walker 2007, pp. 3–10 ([Link]
60. Ungar & Teaford 2002.
61. Bogin 1997, pp. 96–142 ([Link]
[Link]/ANTAO1/Projects/[Link]).
62. Barnicot, Nigel A. (April–June 2005). "Human nutrition: Evolutionary perspectives".
Integrative Physiological & Behavioral Science. 40 (2): 114–117. doi:10.1007/BF02734246
([Link] ISSN 1932-4502 ([Link]
n/1932-4502). PMID 17393680 ([Link]
S2CID 39549910 ([Link]
63. "The new batch – 150,000 years ago" ([Link]
[Link]/sn/prehistoric_life/human/human_evolution/new_batch1.shtml). The
evolution of man. London: BBC Science & Nature. Archived from the original ([Link]
[Link]/sn/prehistoric_life/human/human_evolution/new_batch1.shtml) on January 18,
2006. Retrieved April 28, 2015.
64. Whitehouse, David (June 9, 2003). "When humans faced extinction" ([Link]
2/hi/science/nature/[Link]). BBC News. London: BBC. Archived ([Link]
org/web/20100904071921/[Link] from the
original on September 4, 2010. Retrieved January 5, 2007.
65. "Modern humans flourished through ancient supervolcano eruption 74,000 years ago:
Modern humans flourished through ancient supervolcano eruption" ([Link]
[Link]/releases/2018/03/[Link]). ScienceDaily. Archived ([Link]
g/web/20190124152235/[Link]
m) from the original on January 24, 2019. Retrieved January 24, 2019.
66. Zimmer, Carl (August 31, 2023). "Humanity's Ancestors Nearly Died Out, Genetic Study
Suggests - The population crashed following climate change about 930,000 years ago,
scientists concluded. Other experts aren't convinced by the analysis" ([Link]
om/2023/08/31/science/[Link]). the New York Times. Retrieved
September 2, 2023.
67. Hu, Wangjie; et al. (August 31, 2023). "Genomic inference of a severe human bottleneck
during the Early to Middle Pleistocene transition" ([Link]
e.abq7487). Science. 381 (6661): 979–984. Bibcode:2023Sci...381..979H ([Link]
[Link]/abs/2023Sci...381..979H). doi:10.1126/science.abq7487 ([Link]
6%2Fscience.abq7487). PMID 37651513 ([Link]
S2CID 261396309 ([Link] Retrieved
September 2, 2023.
68. Wood, Bernard; Collard, Mark (1999). "The changing face of Genus Homo". Evolutionary
Anthropology: Issues, News, and Reviews. 8 (6): 195–207. doi:10.1002/(SICI)1520-
6505(1999)8:6<195::AID-EVAN1>[Link];2-2 ([Link]
-6505%281999%298%3A6%3C195%3A%3AAID-EVAN1%[Link]%3B2-2). ISSN 1060-
1538 ([Link] S2CID 86768101 ([Link]
[Link]/CorpusID:86768101).
69. Viegas, Jennifer (May 21, 2010). "Toothy Tree-Swinger May Be Earliest Human" ([Link]
[Link]/web/20150509101320/[Link]
[Link]). Discovery News. Silver Spring, Maryland: Discovery
Communications. Archived from the original ([Link]
[Link]) on May 9, 2015. Retrieved April 28, 2015.
70. Wood, Bernard A. (January 1999). "Homo rudolfensis Alexeev, 1986 – fact or phantom?".
Journal of Human Evolution. 36 (1): 115–118. Bibcode:1999JHumE..36..115W ([Link]
[Link]/abs/1999JHumE..36..115W). doi:10.1006/jhev.1998.0246 ([Link]
0.1006%2Fjhev.1998.0246). ISSN 0047-2484 ([Link]
PMID 9924136 ([Link]
71. Antón, Susan C.; Middleton, Emily R. (June 2023). "Making meaning from fragmentary
fossils: Early Homo in the Early to early Middle Pleistocene" ([Link]
retrieve/pii/S0047248422001671). Journal of Human Evolution. 179 103307.
Bibcode:2023JHumE.17903307A ([Link]
A). doi:10.1016/[Link].2022.103307 ([Link]
PMID 37030994 ([Link]
72. 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). 1
(4): 243–253. Bibcode:2002CRPal...1..243G ([Link]
l...1..243G). doi:10.1016/S1631-0683(02)00032-5 ([Link]
3%2802%2900032-5). ISSN 1631-0683 ([Link]
73. Lordkipanidze, David; Vekua, Abesalom; Ferring, Reid; et al. (November 2006). "A fourth
hominin skull from Dmanisi, Georgia" ([Link] The
Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology.
288A (11): 1146–1157. doi:10.1002/ar.a.20379 ([Link]
ISSN 1552-4884 ([Link] PMID 17031841 ([Link]
[Link]/17031841).
74. 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" ([Link] Journal
of Anatomy and Physiology. 29 (Pt 3): 424–445. PMC 1328414 ([Link]
v/pmc/articles/PMC1328414). PMID 17232143 ([Link]
75. Weidenreich, Franz (July 1940). "Some Problems Dealing with Ancient Man" ([Link]
10.1525%2Faa.1940.42.3.02a00010). American Anthropologist. 42 (3): 375–383.
doi:10.1525/aa.1940.42.3.02a00010 ([Link]
ISSN 0002-7294 ([Link]
76. Grine, Frederick E.; Fleagle, John G. (2009), "The First Humans: A Summary Perspective
on the Origin and Early Evolution of the Genus Homo", The First Humans – Origin and Early
Evolution of the Genus Homo, "Vertebrate Paleobiology and Paleoanthropology" series,
Springer Netherlands, pp. 197–207, doi:10.1007/978-1-4020-9980-9_17 ([Link]
007%2F978-1-4020-9980-9_17), ISBN 978-1-4020-9979-3
77. Rizal, Y.; Westaway, K. E.; Zaim, Y.; van den Bergh; Bettis III, E. A.; Morwood, M. J.;
Huffman, O. F.; Grün, R.; Joannes-Boyau, R.; Bailey, R. M.; Sidarto; Westaway, M. C.;
Kurniawan, I.; Moore, M. W.; Storey, M.; Aziz, F.; Suminto; Zhao, J.; Aswan; Sipola, M. E.;
Larick, R.; Zonneveld, J.-P.; Scott, R.; Putt, S.; Ciochon, R. L. (2020). "Last appearance of
Homo erectus at Ngandong, Java, 117,000–108,000 years ago". Nature. 577 (7790): 381–
385. Bibcode:2020Natur.577..381R ([Link]
R). doi:10.1038/s41586-019-1863-2 ([Link]
hdl:10072/397335 ([Link] ISSN 0028-0836 ([Link]
[Link]/issn/0028-0836). PMID 31853068 ([Link]
8). S2CID 209410644 ([Link]
78. Louys, J.; Roberts, P. (2020). "Environmental drivers of megafauna and hominin extinction
in Southeast Asia". Nature. 586 (7829): 402–406. Bibcode:2020Natur.586..402L ([Link]
[Link]/abs/2020Natur.586..402L). doi:10.1038/s41586-020-2810-y ([Link]
rg/10.1038%2Fs41586-020-2810-y). hdl:10072/402368 ([Link]
2368). PMID 33029012 ([Link] S2CID 222217295 (htt
ps://[Link]/CorpusID:222217295).
79. Spoor, Fred; Wood, Bernard A.; Zonneveld, Frans (June 23, 1994). "Implications of early
hominid labyrinthine morphology for evolution of human bipedal locomotion". Nature. 369
(6482): 645–648. Bibcode:1994Natur.369..645S ([Link]
ur.369..645S). doi:10.1038/369645a0 ([Link] ISSN 0028-
0836 ([Link] PMID 8208290 ([Link]
[Link]/8208290). S2CID 4344784 ([Link]
80. Ings, Simon (October 4, 2009). "Catching Fire: How Cooking Made Us Human by Richard
Wrangham: Review" ([Link]
[Link]). Archived ([Link]
[Link]/archive/20220111/[Link]
-[Link]) from the original on
January 11, 2022. Retrieved February 23, 2016.
81. Wrangham, Richard (2011). Catching Fire: How cooking made us human.
82. 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. 276 (5317): 1392–1395.
doi:10.1126/science.276.5317.1392 ([Link]
ISSN 0036-8075 ([Link] PMID 9162001 ([Link]
[Link]/9162001).
83. Carbonell, Eudald; Bermúdez de Castro, José María; Parés, Josep M.; et al. (March 27,
2008). "The first hominin of Europe". Nature. 452 (7186): 465–469.
Bibcode:2008Natur.452..465C ([Link]
doi:10.1038/nature06815 ([Link] hdl:2027.42/62855 (http
s://[Link]/2027.42%2F62855). ISSN 0028-0836 ([Link]
28-0836). PMID 18368116 ([Link] S2CID 4401629 (htt
ps://[Link]/CorpusID:4401629).
84. García-Martínez, Daniel; Green, David J.; Bermúdez de Castro, José María (February 18,
2021). "Evolutionary development of the Homo antecessor scapulae (Gran Dolina site,
Atapuerca) suggests a modern-like development for Lower Pleistocene Homo" ([Link]
[Link]/pmc/articles/PMC7892855). Scientific Reports. 11 (1) 4102.
Bibcode:2021NatSR..11.4102G ([Link]
doi:10.1038/s41598-021-83039-w ([Link]
ISSN 2045-2322 ([Link] PMC 7892855 ([Link]
[Link]/pmc/articles/PMC7892855). PMID 33602966 ([Link]
ov/33602966).
85. Welker, Frido; Ramos-Madrigal, Jazmín; Gutenbrunner, Petra; Mackie, Meaghan; Tiwary,
Shivani; Rakownikow Jersie-Christensen, Rosa; Chiva, Cristina; Dickinson, Marc R.;
Kuhlwilm, Martin; de Manuel, Marc; Gelabert, Pere; Martinón-Torres, María; Margvelashvili,
Ann; Arsuaga, Juan Luis; Carbonell, Eudald (April 9, 2020). "The dental proteome of Homo
antecessor" ([Link] Nature. 580 (7802):
235–238. Bibcode:2020Natur.580..235W ([Link]
0..235W). doi:10.1038/s41586-020-2153-8 ([Link]
ISSN 0028-0836 ([Link] PMC 7582224 ([Link]
[Link]/pmc/articles/PMC7582224). PMID 32269345 ([Link]
ov/32269345).
86. Manzi, Giorgio; Mallegni, Francesco; Ascenzi, Antonio (August 14, 2001). "A cranium for the
earliest Europeans: Phylogenetic position of the hominid from Ceprano, Italy" ([Link]
[Link]/pmc/articles/PMC55569). Proceedings of the National Academy of Sciences
of the United States of America. 98 (17): 10011–10016. Bibcode:2001PNAS...9810011M (htt
ps://[Link]/abs/2001PNAS...9810011M). doi:10.1073/pnas.151259998 (http
s://[Link]/10.1073%2Fpnas.151259998). ISSN 0027-8424 ([Link]
0027-8424). PMC 55569 ([Link]
PMID 11504953 ([Link]
87. Mallegni, Francesco; Carnieri, Emiliano; Bisconti, Michelangelo; Tartarelli, Giandonato;
Ricci, Stefano; Biddittu, Italo; Segre, Aldo (March 2003). "Homo cepranensis sp. nov. and
the evolution of African-European Middle Pleistocene hominids" ([Link]
Comptes Rendus Palevol. 2 (2): 153–159. Bibcode:2003CRPal...2..153M ([Link]
[Link]/abs/2003CRPal...2..153M). doi:10.1016/S1631-0683(03)00015-0 ([Link]
g/10.1016%2FS1631-0683%2803%2900015-0). hdl:11384/160463 ([Link]
384%2F160463).
88. Czarnetzki, Alfred; Jakob, Tina; Pusch, Carsten M. (April 2003). "Palaeopathological and
variant conditions of the Homo heidelbergensis type specimen (Mauer, Germany)" ([Link]
[Link]/output/1579530). Journal of Human Evolution. 44 (4): 479–
495. Bibcode:2003JHumE..44..479C ([Link]
9C). doi:10.1016/S0047-2484(03)00029-0 ([Link]
2900029-0). ISSN 0047-2484 ([Link] PMID 12727464
([Link]
89. Mounier, Aurélien; Marchal, François; Condemi, Silvana (March 2009). "Is Homo
heidelbergensis a distinct species? New insight on the Mauer mandible" ([Link]
[Link]/retrieve/pii/S0047248409000025). Journal of Human Evolution. 56 (3): 219–246.
Bibcode:2009JHumE..56..219M ([Link]
doi:10.1016/[Link].2008.12.006 ([Link]
PMID 19249816 ([Link]
90. Grün, Rainer; Pike, Alistair; McDermott, Frank; Eggins, Stephen; Mortimer, Graham; Aubert,
Maxime; Kinsley, Lesley; Joannes-Boyau, Renaud; Rumsey, Michael; Denys, Christiane;
Brink, James; Clark, Tara; Stringer, Chris (April 16, 2020). "Dating the skull from Broken Hill,
Zambia, and its position in human evolution" ([Link]
165-4). Nature. 580 (7803): 372–375. Bibcode:2020Natur.580..372G ([Link]
[Link]/abs/2020Natur.580..372G). doi:10.1038/s41586-020-2165-4 ([Link]
8%2Fs41586-020-2165-4). ISSN 0028-0836 ([Link]
PMID 32296179 ([Link]
91. Grün, Rainer; Pike, Alistair; McDermott, Frank; Eggins, Stephen; Mortimer, Graham; Aubert,
Maxime; Kinsley, Lesley; Joannes-Boyau, Renaud; Rumsey, Michael; Denys, Christiane;
Brink, James; Clark, Tara; Stringer, Chris (April 16, 2020). "Dating the skull from Broken Hill,
Zambia, and its position in human evolution" ([Link]
165-4). Nature. 580 (7803): 372–375. Bibcode:2020Natur.580..372G ([Link]
[Link]/abs/2020Natur.580..372G). doi:10.1038/s41586-020-2165-4 ([Link]
8%2Fs41586-020-2165-4). ISSN 0028-0836 ([Link]
PMID 32296179 ([Link]
92. Semaw, Sileshi; Toth, Nicholas; Schick, Kathy; et al. (March 27, 2006). "Scientists discover
hominid cranium in Ethiopia" ([Link] (Press
release). Bloomington: Indiana University. Archived ([Link]
61617/[Link] from the original on November
15, 2006. Retrieved November 26, 2006.
93. Harvati, Katerina (January 2003). "The Neanderthal taxonomic position: Models of intra- and
inter-specific craniofacial variation". Journal of Human Evolution. 44 (1): 107–132.
Bibcode:2003JHumE..44..107H ([Link]
doi:10.1016/S0047-2484(02)00208-7 ([Link]
208-7). ISSN 0047-2484 ([Link] PMID 12604307 (http
s://[Link]/12604307).
94. 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 ([Link]
x). PMID 19391204 ([Link] S2CID 25787484 ([Link]
[Link]/CorpusID:25787484).
95. Finlayson, C.; Giles Pacheco, F.; Rodríguez-Vidal, J.; et al. (2006). "Late survival of
Neanderthals at the southernmost extreme of Europe". Nature. 443 (7113): 850–853.
Bibcode:2006Natur.443..850F ([Link]
doi:10.1038/nature05195 ([Link] hdl:10261/18685 (http
s://[Link]/10261%2F18685). PMID 16971951 ([Link]
71951). S2CID 4411186 ([Link]
Full list of authors
Clive Finlayson
Francisco Giles Pacheco
Joaquín Rodríguez-Vidal
Darren A. Fa
José María Gutierrez López
Antonio Santiago Pérez
Geraldine Finlayson
Ethel Allue
Javier Baena Preysler
Isabel Cáceres
José S. Carrión
Yolanda Fernández-Jalvo
Christopher P. Gleed-Owen
Francisco J. Jimenez-Espejo
Pilar López Martínez
José Antonio López Sáez
José Antonio Riquelme Cantal
Antonio Sánchez Marco
Francisco Giles Guzman
Kimberly Brown
Noemí Fuentes
Claire A. Valarino
Antonio Villalpando
Christopher B. Stringer
Francisca Martinez Ruiz
Tatsuhiko Sakamoto
96. Pearce, Eiluned; Stringer, Chris; Dunbar, R. I. M. (2013). "New insights into differences in
brain organization between Neanderthals and anatomically modern humans" ([Link]
[Link]/pmc/articles/PMC3619466). Proceedings of the Royal Society of London B:
Biological Sciences. 280 (1758) 20130168. doi:10.1098/rspb.2013.0168 ([Link]
098%2Frspb.2013.0168). PMC 3619466 ([Link]
19466). PMID 23486442 ([Link]
97. Bocquet-Appel, Jean-Pierre; Degioanni, Anna (December 2013). "Neanderthal Demographic
Estimates" ([Link] Current
Anthropology. 54 (S8): S202–S213. Bibcode:2013CurrA..54S.202B ([Link]
[Link]/abs/2013CurrA..54S.202B). doi:10.1086/673725 ([Link]
 98. Krings, Matthias; Stone, Anne; Schmitz, Ralf W.; et al. (July 11, 1997). "Neandertal DNA
sequences and the origin of modern humans". Cell. 90 (1): 19–30. doi:10.1016/S0092-
8674(00)80310-4 ([Link]
hdl:11858/00-001M-0000-0025-0960-8 ([Link]
5-0960-8). ISSN 0092-8674 ([Link] PMID 9230299 (ht
tps://[Link]/9230299). S2CID 13581775 ([Link]
CorpusID:13581775).
99. Green, Richard E.; Malaspinas, Anna-Sapfo; Krause, Johannes; et al. (August 8, 2008). "A
Complete Neandertal Mitochondrial Genome Sequence Determined by High-Throughput
Sequencing" ([Link] Cell. 134 (3): 416–
426. doi:10.1016/[Link].2008.06.021 ([Link]
ISSN 0092-8674 ([Link] PMC 2602844 ([Link]
[Link]/pmc/articles/PMC2602844). PMID 18692465 ([Link]
ov/18692465).
100. Serre, David; Langaney, André; Chech, Mario; et al. (March 2004). "No Evidence of
Neandertal mtDNA Contribution to Early Modern Humans" ([Link]
c/articles/PMC368159). PLOS Biology. 2 (3) e57. doi:10.1371/[Link].0020057 (https://
[Link]/10.1371%[Link].0020057). ISSN 1545-7885 ([Link]
n/1545-7885). PMC 368159 ([Link]
PMID 15024415 ([Link]
101. Viegas, Jennifer (May 6, 2010). "Neanderthals, Humans Interbred, DNA Proves" ([Link]
[Link]/web/20150508221757/[Link]
[Link]). Discovery News. Silver Spring, Maryland: Discovery
Communications. Archived from the original ([Link]
[Link]) on May 8, 2015. Retrieved April 30, 2015.
102. Calloway, Ewan (May 13, 2015). "Early European may have had Neanderthal great-great-
grandparent" ([Link]
at-great-grandparent-1.17534). Nature. doi:10.1038/nature.2015.17534 ([Link]
038%2Fnature.2015.17534). S2CID 181973496 ([Link]
81973496). Archived ([Link]
news/early-european-may-have-had-neanderthal-great-great-grandparent-1.17534) from
the original on January 15, 2019. Retrieved January 23, 2019.
103. Sample, Ian (June 22, 2015). "My Neanderthal sex secret: Modern European's great-great
grandparent link" ([Link]
cret-modern-europeans-great-great-grandparent-link). The Guardian. Archived ([Link]
[Link]/web/20160923235758/[Link]
anderthal-sex-secret-modern-europeans-great-great-grandparent-link) from the original on
September 23, 2016. Retrieved July 27, 2018.
104. Green, R. E.; Krause, J.; Briggs, A. W.; Maricic, T.; et al. (May 7, 2010). "A Draft Sequence
of the Neandertal Genome" ([Link]
Science. 328 (5979): 710–722. Bibcode:2010Sci...328..710G ([Link]
abs/2010Sci...328..710G). doi:10.1126/science.1188021 ([Link]
e.1188021). ISSN 0036-8075 ([Link] PMC 5100745
([Link] PMID 20448178 ([Link]
[Link]/20448178).
Full list of authors
Richard E. Green
Johannes Krause
Adrian W. Briggs
Tomislav Maricic
Udo Stenzel
Martin Kircher
Nick Patterson
Heng Li
Weiwei Zhai
Markus Hsi-Yang Fritz
Nancy F. Hansen
Eric Y. Durand
Anna-Sapfo Malaspinas
Jeffrey D. Jensen
Tomas Marques-Bonet
Can Alkan
Kay Prüfer
Matthias Meyer
Hernán A. Burbano
Jeffrey M. Good
Rigo Schultz
Ayinuer Aximu-Petri
Anne Butthof
Barbara Höber
Barbara Höffner
Madlen Siegemund
Antje Weihmann
Chad Nusbaum
Eric S. Lander
Carsten Russ
Nathaniel Novod
Jason Affourtit
Michael Egholm
Christine Verna
Pavao Rudan
Dejana Brajkovic
Željko Kucan
Ivan Gušic
 Vladimir B. Doronichev
Liubov V. Golovanova
Carles Lalueza-Fox
Marco de la Rasilla
Javier Fortea
Antonio Rosas
Ralf W. Schmitz
Philip L. F. Johnson
Evan E. Eichler
Daniel Falush
Ewan Birney
James C. Mullikin
Montgomery Slatkin
Rasmus Nielsen
Janet Kelso
Michael Lachmann
David Reich
Svante Pääbo
105. Prüfer, K.; de Filippo, C.; Grote, S.; Mafessoni, F.; Korlević, P.; Hajdinjak, M.; et al. (2017).
"A high-coverage Neandertal genome from Vindija Cave in Croatia" ([Link]
[Link]/pmc/articles/PMC6185897). Science. 358 (6363): 655–658.
Bibcode:2017Sci...358..655P ([Link]
doi:10.1126/science.aao1887 ([Link] PMC 6185897
([Link] PMID 28982794 ([Link]
[Link]/28982794).
106. Gutiérrez, Gabriel; Sánchez, Diego; Marín, Antonio (August 2002). "A Reanalysis of the
Ancient Mitochondrial DNA Sequences Recovered from Neandertal Bones" ([Link]
0.1093%[Link].a004197). Molecular Biology and Evolution. 19 (8): 1359–
1366. doi:10.1093/[Link].a004197 ([Link]
[Link].a004197). ISSN 0737-4038 ([Link]
PMID 12140248 ([Link]
107. Hebsgaard, M. B.; Wiuf, C.; Gilbert, M. T.; Glenner, H.; Willerslev, E. (January 2007).
"Evaluating Neanderthal Genetics and Phylogeny" ([Link]
01041826/[Link] (PDF). Journal of
Molecular Evolution. 64 (1): 50–60. Bibcode:2007JMolE..64...50H ([Link]
[Link]/abs/2007JMolE..64...50H). CiteSeerX [Link].8969 ([Link]
ewdoc/summary?doi=[Link].8969). doi:10.1007/s00239-006-0017-y ([Link]
1007%2Fs00239-006-0017-y). ISSN 0022-2844 ([Link]
4). PMID 17146600 ([Link] S2CID 2746487 ([Link]
[Link]/CorpusID:2746487). Archived from the original ([Link]
k/%7Ewiuf/journalWiuf/[Link]) (PDF) on April 1, 2011. Retrieved October 24, 2017.
{{cite journal}}: Cite uses deprecated parameter |citeseerx= (help)
108. Mellars, Paul; French, Jennifer C. (July 29, 2011). "Tenfold Population Increase in Western
Europe at the Neandertal–to–Modern Human Transition Paul". Science. 333 (6042): 623–
627. Bibcode:2011Sci...333..623M ([Link]
doi:10.1126/science.1206930 ([Link] ISSN 0036-
8075 ([Link] PMID 21798948 ([Link]
[Link]/21798948). S2CID 28256970 ([Link]
0).
109. Brown, Terence A. (April 8, 2010). "Human evolution: Stranger from Siberia" ([Link]
10.1038%2F464838a). Nature. 464 (7290): 838–839. Bibcode:2010Natur.464..838B (https://
[Link]/abs/2010Natur.464..838B). doi:10.1038/464838a ([Link]
038%2F464838a). ISSN 0028-0836 ([Link]
PMID 20376137 ([Link] S2CID 4320406 ([Link]
[Link]/CorpusID:4320406).
110. Krause, Johannes; Fu, Qiaomei; Good, Jeffrey M.; et al. (April 8, 2010). "The complete
mitochondrial DNA genome of an unknown hominin from southern Siberia" ([Link]
[Link]/pmc/articles/PMC10152974). Nature. 464 (7290): 894–897.
Bibcode:2010Natur.464..894K ([Link]
doi:10.1038/nature08976 ([Link] ISSN 0028-0836 (http
s://[Link]/issn/0028-0836). PMC 10152974 ([Link]
articles/PMC10152974). PMID 20336068 ([Link]
S2CID 4415601 ([Link]
111. Katsnelson, Alla (March 24, 2010). "New hominin found via mtDNA" ([Link]
[Link]/?[Link]/articleNo/28876/title/New-hominin-found-via-mtDNA/#ixzz0j820ioz1).
The Nutshell (Blog). Philadelphia: The Scientist. ISSN 0890-3670 ([Link]
g/issn/0890-3670). Archived ([Link]
[Link]/?[Link]/articleNo/28876/title/New-hominin-found-via-mtDNA/#ixzz0j820io
z1) from the original on July 2, 2015. Retrieved May 1, 2015.
112. "Kaufman, Danial (2002), "Comparisons and the Case for Interaction among Neanderthals
and Early Modern Humans in the Levant" (Oxford Journal of Anthropology)
113. Bokma, Folmer; van den Brink, Valentijn; Stadler, Tanja (September 2012). "Unexpectedly
many extinct hominins" ([Link] Evolution.
66 (9): 2969–2974. Bibcode:2012Evolu..66.2969B ([Link]
volu..66.2969B). doi:10.1111/j.1558-5646.2012.01660.x ([Link]
646.2012.01660.x). ISSN 0014-3820 ([Link]
PMID 22946817 ([Link] S2CID 13145359 ([Link]
[Link]/CorpusID:13145359).
114. Reich, David; Patterson, Nick; Kircher, Martin; et al. (October 7, 2011). "Denisova Admixture
and the First Modern Human Dispersals into Southeast Asia and Oceania" ([Link]
[Link]/pmc/articles/PMC3188841). American Journal of Human Genetics. 89 (4):
516–528. doi:10.1016/[Link].2011.09.005 ([Link]
ISSN 0002-9297 ([Link] PMC 3188841 ([Link]
[Link]/pmc/articles/PMC3188841). PMID 21944045 ([Link]
ov/21944045).
115. 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. 60
(2): 251–255. Bibcode:2011JHumE..60..251M ([Link]
E..60..251M). doi:10.1016/[Link].2010.10.005 ([Link]
005). ISSN 0047-2484 ([Link] PMID 21129766 (http
s://[Link]/21129766).
116. Simonti, C. N.; Vernot, B.; Bastarache, L.; Bottinger, E.; Carrell, D. S.; Chisholm, R. L.;
Crosslin, D. R.; Hebbring, S. J.; Jarvik, G. P.; Kullo, I. J.; Li, R.; Pathak, J.; Ritchie, M. D.;
Roden, D. M.; Verma, S. S.; Tromp, G.; Prato, J. D.; Bush, W. S.; Akey, J. M.; Denny, J. C.;
Capra, J. A. (2016). "The phenotypic legacy of admixture between modern humans and
Neandertals" ([Link] Science. 351
(6274): 737–741. Bibcode:2016Sci...351..737S ([Link]
i...351..737S). doi:10.1126/science.aad2149 ([Link]
PMC 4849557 ([Link] PMID 26912863
([Link]
117. Kuhlwilm, M.; Gronau, I.; Hubisz, M. J.; de Filippo, C.; Prado-Martinez, J.; Kircher, M.; Fu,
Q.; Burbano, H. A.; Lalueza-Fox, C.; de la Rasilla, M.; Rosas, A.; Rudan, P.; Brajkovic, D.;
Kucan, Ž.; Gušic, I.; Marques-Bonet, T.; Andrés, A. M.; Viola, B.; Pääbo, S.; Meyer, M.;
Siepel, A.; Castellano, S. (2016). "Ancient gene flow from early modern humans into Eastern
Neanderthals" ([Link] Nature. 530
(7591): 429–433. Bibcode:2016Natur.530..429K ([Link]
ur.530..429K). doi:10.1038/nature16544 ([Link]
PMC 4933530 ([Link] PMID 26886800
([Link]
118. Dean, MC, Stringer, CB et al, (1986) "Age at death of the Neanderthal child from Devil's
Tower, Gibraltar and the implications for studies of general growth and development in
Neanderthals" (American Journal of Physical Anthropology, Vol 70 Issue 3, July 1986)
119. Brown, P.; Sutikna, T.; Morwood, M. J.; Soejono, R. P.; Jatmiko, A.; Wayhu, S. E.; Awe Due,
R. (October 28, 2004). "A new small-bodied hominin from the Late Pleistocene of Flores,
Indonesia" ([Link] (PDF). Nature. 431 (7012):
1055–1061. Bibcode:2004Natur.431.1055B ([Link]
31.1055B). doi:10.1038/nature02999 ([Link] ISSN 0028-
0836 ([Link] PMID 15514638 ([Link]
[Link]/15514638). S2CID 26441 ([Link]
Archived ([Link]
s/PAL_E2586.pdf) (PDF) from the original on January 3, 2023. Retrieved January 3, 2023.
120. Argue, Debbie; Donlon, Denise; Groves, Colin; Wright, Richard (October 2006). "Homo
floresiensis: Microcephalic, pygmoid, Australopithecus, or Homo?". Journal of Human
Evolution. 51 (4): 360–374. Bibcode:2006JHumE..51..360A ([Link]
bs/2006JHumE..51..360A). doi:10.1016/[Link].2006.04.013 ([Link]
evol.2006.04.013). ISSN 0047-2484 ([Link]
PMID 16919706 ([Link]
121. Martin, Robert D.; Maclarnon, Ann M.; Phillips, James L.; Dobyns, William B. (November
2006). "Flores hominid: New species or microcephalic dwarf?" ([Link]
r.a.20389). The Anatomical Record Part A: Discoveries in Molecular, Cellular, and
Evolutionary Biology. 288A (11): 1123–1145. doi:10.1002/ar.a.20389 ([Link]
2%2Far.a.20389). ISSN 1552-4884 ([Link]
PMID 17031806 ([Link]
122. Callaway, E. (June 8, 2016). " 'Hobbit' relatives found after ten-year hunt" ([Link]
1038%2F534164a). Nature. 534 (7606): 164–165. Bibcode:2016Natur.534Q.164C ([Link]
[Link]/abs/2016Natur.534Q.164C). doi:10.1038/534164a ([Link]
038%2F534164a). PMID 27279191 ([Link]
123. Brumm, A.; van den Bergh, G. D.; Storey, M.; Kurniawan, I.; et al. (June 8, 2016). "Age and
context of the oldest known hominin fossils from Flores" ([Link]
0924/Brumm_et_al_2016_Age_stratigraphic_context_hominin_fossils_Flores_NAture_DRA
FT_MS.pdf) (PDF). Nature. 534 (7606): 249–253. Bibcode:2016Natur.534..249B ([Link]
[Link]/abs/2016Natur.534..249B). doi:10.1038/nature17663 ([Link]
038%2Fnature17663). PMID 27279222 ([Link]
S2CID 28608179 ([Link] Archived ([Link]
[Link]/web/20200306112537/[Link]
016_Age_stratigraphic_context_hominin_fossils_Flores_NAture_DRAFT_MS.pdf) (PDF)
from the original on March 6, 2020. Retrieved November 11, 2021.
Full list of authors
Adam Brumm
Gerrit D. van den Bergh
Michael Storey
Iwan Kurniawan
Brent V. Alloway
Ruly Setiawan
Erick Setiyabudi
Rainer Grün
Mark W. Moore
Dida Yurnaldi
Mika R. Puspaningrum
Unggul P. Wibowo
Halmi Insani
Indra Sutisna
John A. Westgate
Nick J. G. Pearce
Mathieu Duval
Hanneke J. M. Meijer
Fachroel Aziz
Thomas Sutikna
Sander van der Kaars
Stephanie Flude
Michael J. Morwood
124. van den Bergh, G. D.; Kaifu, Y.; Kurniawan, I.; Kono, R. T.; Brumm, A.; Setiyabudi, E.; Aziz,
F.; Morwood, M. J. (June 8, 2016). "Homo floresiensis-like fossils from the early Middle
Pleistocene of Flores" ([Link] Nature.
534 (7606): 245–248. Bibcode:2016Natur.534..245V ([Link]
6Natur.534..245V). doi:10.1038/nature17999 ([Link]
PMID 27279221 ([Link] S2CID 205249218 ([Link]
[Link]/CorpusID:205249218).
125. Détroit, F.; Mijares, A. S.; Corny, J.; Daver, G.; Zanolli, C.; Dizon, E.; Robles, E.; Grün, R. &
Piper, P. J. (2019). "A new species of Homo from the Late Pleistocene of the Philippines" (htt
ps://[Link]/hal-02296712/file/Detroit_%26_al_2019_Nature_postprint.pdf)
(PDF). Nature. 568 (7751): 181–186. Bibcode:2019Natur.568..181D ([Link]
[Link]/abs/2019Natur.568..181D). doi:10.1038/s41586-019-1067-9 ([Link]
8%2Fs41586-019-1067-9). hdl:10072/386785 ([Link]
PMID 30971845 ([Link] S2CID 106411053 ([Link]
[Link]/CorpusID:106411053). Archived ([Link]
3114830/[Link]
[Link]) (PDF) from the original on October 13, 2022. Retrieved May 11, 2021.
126. Schlebusch, Carina M.; Malmström, Helena; Günther, Torsten; Sjödin, Per; Coutinho,
Alexandra; Edlund, Hanna; Munters, Arielle R.; Vicente, Mário; Steyn, Maryna; Soodyall,
Himla; Lombard, Marlize; Jakobsson, Mattias (2017). "Southern African ancient genomes
estimate modern human divergence to 350,000 to 260,000 years ago" ([Link]
26%2Fscience.aao6266). Science. 358 (6363): 652–655. Bibcode:2017Sci...358..652S (http
s://[Link]/abs/2017Sci...358..652S). doi:10.1126/science.aao6266 ([Link]
[Link]/10.1126%2Fscience.aao6266). PMID 28971970 ([Link]
71970). S2CID 206663925 ([Link]
127. Sample, Ian (June 7, 2017). "Oldest Homo sapiens bones ever found shake foundations of
the human story" ([Link]
ones-ever-found-shake-foundations-of-the-human-story). The Guardian. Archived ([Link]
[Link]/web/20191031005024/[Link]
est-homo-sapiens-bones-ever-found-shake-foundations-of-the-human-story) from the
original on October 31, 2019. Retrieved June 7, 2017.
128. Zimmer, Carl (September 10, 2019). "Scientists Find the Skull of Humanity's Ancestor — on
a Computer: By comparing fossils and CT scans, researchers say they have reconstructed
the skull of the last common forebear of modern humans" ([Link]
0220101/[Link]
l). The New York Times. Archived from the original ([Link]
ence/[Link]) on January 1, 2022. Retrieved September 10,
2019.
129. Mounier, Aurélien; Lahr, Marta (2019). "Deciphering African late middle Pleistocene hominin
diversity and the origin of our species" ([Link]
881). Nature Communications. 10 (1): 3406. Bibcode:2019NatCo..10.3406M ([Link]
[Link]/abs/2019NatCo..10.3406M). doi:10.1038/s41467-019-11213-w ([Link]
g/10.1038%2Fs41467-019-11213-w). PMC 6736881 ([Link]
les/PMC6736881). PMID 31506422 ([Link]
130. Ambrose, Stanley H. (June 1998). "Late Pleistocene human population bottlenecks, volcanic
winter, and differentiation of modern humans". Journal of Human Evolution. 34 (6): 623–651.
Bibcode:1998JHumE..34..623A ([Link]
doi:10.1006/jhev.1998.0219 ([Link] ISSN 0047-2484
([Link] PMID 9650103 ([Link]
ov/9650103). S2CID 33122717 ([Link]
131. Huff, Chad D.; Xing, Jinchuan; Rogers, Alan R.; Witherspoon, David; Jorde, Lynn B.
(January 19, 2010). "Mobile Elements Reveal Small Population Size in the Ancient
Ancestors of Homo sapiens" ([Link]
[Link]+html). Proceedings of the National Academy of Sciences of the United States of
America. 107 (5): 2147–2152. Bibcode:2010PNAS..107.2147H ([Link]
u/abs/2010PNAS..107.2147H). doi:10.1073/pnas.0909000107 ([Link]
nas.0909000107). PMC 2836654 ([Link]
PMID 20133859 ([Link]
132. Clark, Jamie L. (September 2011). "The evolution of human culture during the later
Pleistocene: Using fauna to test models on the emergence and nature of "modern" human
behavior" ([Link] Journal of
Anthropological Archaeology. 30 (3): 273–291. Bibcode:2011JAnAr..30..273C ([Link]
[Link]/abs/2011JAnAr..30..273C). doi:10.1016/[Link].2011.04.002 ([Link]
0.1016%[Link].2011.04.002). Archived ([Link]
s://[Link]/retrieve/pii/S0278416511000225) from the original on May 25,
2021. Retrieved October 27, 2021.
133. Boyd & Silk 2003, p. .
134. Brues & Snow 1965, pp. 1–39 ([Link]
A1).
135. Barham, Lawrence; Mitchell, Peter (June 23, 2008). The First Africans: African Archaeology
from the Earliest Toolmakers to Most Recent Foragers ([Link]
duct/identifier/9780511817830/type/book) (1 ed.). Cambridge University Press.
doi:10.1017/cbo9780511817830 ([Link]
ISBN 978-0-521-61265-4.
136. Brunet, M.; Guy, F.; Pilbeam, D.; Mackaye, H.; Likius, A.; Ahounta, D.; et al. (July 11, 2002).
"A new hominid from the Upper Miocene of Chad, Central Africa" ([Link]
3388/files/PAL_E190.pdf) (PDF). Nature. 418 (6894): 145–151.
Bibcode:2002Natur.418..145B ([Link]
doi:10.1038/nature00879 ([Link] ISSN 0028-0836 (http
s://[Link]/issn/0028-0836). PMID 12110880 ([Link]
2110880). S2CID 1316969 ([Link] Archived (ht
tps://[Link]/web/20230225204437/[Link]
[Link]) (PDF) from the original on February 25, 2023. Retrieved February 20, 2023.
Full list of authors
Michel Brunet
Franck Guy
David Pilbeam
Hassane Taisso Mackaye
Andossa Likius
Djimdoumalbaye Ahounta
Alain Beauvilain
Cécile Blondel
Hervé Bocherens
Jean-Renaud Boisserie
Louis De Bonis
Yves Coppens
Jean Dejax
Christiane Denys
Philippe Duringer
Véra Eisenmann
Gongdibé Fanone
Pierre Fronty
Denis Geraads
Thomas Lehmann
Fabrice Lihoreau
Antoine Louchart
Adoum Mahamat
Gildas Merceron
Guy Mouchelin
Olga Otero
Pablo Pelaez Campomanes
Marcia Ponce De Leon
Jean-Claude Rage
Michel Sapanet
Mathieu Schuster
Jean Sudre
Pascal Tassy
Xavier Valentin
Patrick Vignaud
Laurent Viriot
 Antoine Zazzo
Christoph Zollikofer
137. White, T. D.; Asfaw, B.; Beyene, Y.; Haile-Selassie, Y.; Lovejoy, C. O.; Suwa, G.;
Woldegabriel, G. (2009). "Ardipithecus ramidus and the Paleobiology of Early Hominids".
Science. 326 (5949): 75–86. Bibcode:2009Sci...326...75W ([Link]
s/2009Sci...326...75W). doi:10.1126/science.1175802 ([Link]
175802). PMID 19810190 ([Link] S2CID 20189444 (h
ttps://[Link]/CorpusID:20189444).
138. Kwang Hyun, Ko (2015). "Origins of Bipedalism". Brazilian Archives of Biology and
Technology. 58 (6): 929–934. arXiv:1508.02739 ([Link]
Bibcode:2015arXiv150802739K ([Link]
doi:10.1590/S1516-89132015060399 ([Link]
S2CID 761213 ([Link]
139. DeSalle & Tattersall 2008, p. 146 ([Link]
A146).
140. Curry 2008, pp. 106–109.
141. "Study Identifies Energy Efficiency as Reason for Evolution of Upright Walking" ([Link]
[Link]/releases/2007/07/[Link]). ScienceDaily. July 17, 2007.
Archived ([Link]
es/2007/07/[Link]) from the original on May 4, 2015. Retrieved April 9, 2015.
"Study identifies energy efficiency as reason for evolution of upright walking" ([Link]
[Link]/web/20220703195556/[Link]
son-evolution-upright-walking). UANews. Tucson: University of Arizona. July 16, 2007.
Archived from the original on July 3, 2022. Retrieved April 23, 2015.
142. Sockol, Michael D.; Raichlen, David A.; Pontzer, Herman (July 24, 2007). "Chimpanzee
locomotor energetics and the origin of human bipedalism" ([Link]
c/articles/PMC1941460). Proceedings of the National Academy of Sciences of the United
States of America. 104 (30): 12265–12269. Bibcode:2007PNAS..10412265S ([Link]
[Link]/abs/2007PNAS..10412265S). doi:10.1073/pnas.0703267104 ([Link]
10.1073%2Fpnas.0703267104). ISSN 0027-8424 ([Link]
24). PMC 1941460 ([Link]
PMID 17636134 ([Link]
143. David-Barrett, T.; Dunbar, R. I. M. (2016). "Bipedality and Hair-loss Revisited: The Impact of
Altitude and Activity Scheduling" ([Link]
Journal of Human Evolution. 94: 72–82. doi:10.1016/[Link].2016.02.006 ([Link]
1016%[Link].2016.02.006). PMC 4874949 ([Link]
MC4874949). PMID 27178459 ([Link]
144. Aiello & Dean 1990.
145. Kondo 1985.
146. Strickberger 2000, pp. 475–476 ([Link]
476).
147. Trevathan 2011, p. 20 ([Link]
148. Zuk, Marlene (2014). Paleofantasy: What Evolution Really Tells Us About Sex, Diet, and
How We Live. W.W. Norton & Company. ISBN 978-0-393-34792-0. OCLC 846889455 (http
s://[Link]/oclc/846889455).
149. Hrdy, Sarah Blaffer (2011). Mothers and Others: The Evolutionary Origins of Mutual
Understanding. Harvard University Press. ISBN 978-0-674-06032-6. OCLC 940575388 (http
s://[Link]/oclc/940575388).
150. Wayman, Erin (August 19, 2013). "Killer whales, grandmas and what men want:
Evolutionary biologists consider menopause" ([Link]
les-grandmas-and-what-men-want-evolutionary-biologists-consider-menopause). Science
News. ISSN 0036-8423 ([Link] Archived ([Link]
[Link]/web/20141106194849/[Link]
as-and-what-men-want-evolutionary-biologists-consider-menopause) from the original on
November 6, 2014. Retrieved April 24, 2015.
151. Blell, Mwenza (September 29, 2017). "Grandmother Hypothesis, Grandmother Effect, and
Residence Patterns". The International Encyclopedia of Anthropology. pp. 1–5.
doi:10.1002/9781118924396.wbiea2162 ([Link]
2162). ISBN 978-1-118-92439-6.
152. Welker, F.; Ramos-Madrigal, J.; Gutenbrunner, P.; et al. (April 1, 2020). "The dental
proteome of Homo antecessor" ([Link]
n1_AAM.docx). Nature. 580 (7802): 235–238. Bibcode:2020Natur.580..235W ([Link]
[Link]/abs/2020Natur.580..235W). doi:10.1038/s41586-020-2153-8 ([Link]
g/10.1038%2Fs41586-020-2153-8). ISSN 1476-4687 ([Link]
4687). PMC 7582224 ([Link]
PMID 32269345 ([Link] S2CID 214736611 ([Link]
[Link]/CorpusID:214736611). Archived ([Link]
0200345/[Link] from
the original on November 10, 2021. Retrieved November 10, 2021.
Full list of authors
Frido Welker
Jazmín Ramos-Madrigal
Petra Gutenbrunner
Meaghan Mackie
Shivani Tiwary
Rosa Rakownikow Jersie-Christensen
Cristina Chiva
Marc R. Dickinson
Martin Kuhlwilm
Marc de Manuel
Pere Gelabert
María Martinón-Torres
Ann Margvelashvili
Juan Luis Arsuaga
Eudald Carbonell
Tomas Marques-Bonet
Kirsty Penkman
Eduard Sabidó
Jürgen Cox
Jesper V. Olsen
David Lordkipanidze
Fernando Racimo
Carles Lalueza-Fox
José María Bermúdez de Castro
Eske Willerslev
Enrico Cappellini
153. Dennell, Robin; Roebroeks, Wil (2005). "An Asian perspective on early human dispersal
from Africa" ([Link] Nature. 438 (7071): 1099–1104.
Bibcode:2005Natur.438.1099D ([Link]
doi:10.1038/nature04259 ([Link] PMID 16371999 (http
s://[Link]/16371999). S2CID 4405913 ([Link]
orpusID:4405913). Archived ([Link]
[Link]/articles/nature04259) from the original on October 31, 2021. Retrieved
November 10, 2021.
154. Schoenemann, P. Thomas (October 2006). "Evolution of the Size and Functional Areas of
the Human Brain". Annual Review of Anthropology. 35 (1): 379–406.
Bibcode:2006ARAnt..35..379S ([Link]
doi:10.1146/[Link].35.081705.123210 ([Link]
5.081705.123210). ISSN 0084-6570 ([Link]
S2CID 7611321 ([Link]
155. Brown, Graham; Fairfax, Stephanie; Sarao, Nidhi. "Tree of Life Web Project: Human
Evolution" ([Link] [Link]. Archived (ht
tps://[Link]/web/20200606091921/[Link]
10) from the original on June 6, 2020. Retrieved August 24, 2015.
156. Kaplan, Hillard; Hill, Kim; Lancaster, Jane; Hurtado, Magdelena (August 16, 2000). "A
Theory of Human Life History Evolution: Diet, Intelligence, and Longevity" ([Link]
edu/~hkaplan/KaplanHillLancasterHurtado_2000_LHEvolution.pdf) (PDF). Evolutionary
Anthropology. 9 (4): 156–185. doi:10.1002/1520-6505(2000)9:4<156::AID-
EVAN5>[Link];2-7 ([Link]
A%3AAID-EVAN5%[Link]%3B2-7). S2CID 2363289 ([Link]
pusID:2363289). Archived ([Link]
du/~hkaplan/KaplanHillLancasterHurtado_2000_LHEvolution.pdf) (PDF) from the original on
February 15, 2020. Retrieved September 20, 2019.
157. Park, M. S.; Nguyen, A. D.; Aryan, H. E.; U, H. S.; Levy, M. L.; Semendeferi, K. (March
2007). "Evolution of the human brain: Changing brain size and the fossil record".
Neurosurgery. 60 (3): 555–562. doi:10.1227/[Link].0000249284.54137.32 ([Link]
10.1227%[Link].0000249284.54137.32). ISSN 0148-396X ([Link]
ssn/0148-396X). PMID 17327801 ([Link]
S2CID 19610624 ([Link]
158. Bruner, Emiliano (December 2007). "Cranial shape and size variation in human evolution:
Structural and functional perspectives". Child's Nervous System. 23 (12): 1357–1365.
CiteSeerX [Link].288 ([Link]
88). doi:10.1007/s00381-007-0434-2 ([Link]
ISSN 0256-7040 ([Link] PMID 17680251 ([Link]
[Link]/17680251). S2CID 16163137 ([Link]
D:16163137). {{cite journal}}: Cite uses deprecated parameter |citeseerx= (help)
159. Potts, Richard (October 2012). "Evolution and Environmental Change in Early Human
Prehistory". Annual Review of Anthropology. 41 (1): 151–167.
Bibcode:2012ARAnt..41..151P ([Link]
doi:10.1146/annurev-anthro-092611-145754 ([Link]
611-145754). ISSN 0084-6570 ([Link]
160. Neubauer, Simon; Hublin, Jean-Jacques; Gunz, Philipp (January 5, 2018). "The evolution of
modern human brain shape - Science" ([Link]
678). Science Advances. 4 (1) eaao5961. Bibcode:2018SciA....4.5961N ([Link]
[Link]/abs/2018SciA....4.5961N). doi:10.1126/sciadv.aao5961 ([Link]
2Fsciadv.aao5961). PMC 5783678 ([Link]
8). PMID 29376123 ([Link]
161. Mann, Neil (September 2007). "Meat in the human diet: An anthropological perspective" (htt
ps://[Link]/10.1111%2Fj.1747-0080.2007.00194.x). Nutrition & Dietetics. 64 (Supplement
4): S102–S107. doi:10.1111/j.1747-0080.2007.00194.x ([Link]
80.2007.00194.x). ISSN 1747-0080 ([Link]
162. McBroom, Patricia (June 14, 1999). "Meat-eating was essential for human evolution, says
UC Berkeley anthropologist specializing in diet" ([Link]
es/99legacy/[Link]) (Press release). Berkeley: University of California Press.
Archived ([Link]
a/releases/99legacy/[Link]) from the original on April 20, 2015. Retrieved April 25,
2015.
163. Barr, W. Andrew; Pobiner, Briana; Rowan, John; Du, Andrew; Faith, J. Tyler (February 1,
2022). "No sustained increase in zooarchaeological evidence for carnivory after the
appearance of Homo erectus" ([Link]
Proceedings of the National Academy of Sciences of the United States of America. 119 (5)
e2115540119. Bibcode:2022PNAS..11915540B ([Link]
AS..11915540B). doi:10.1073/pnas.2115540119 ([Link]
19). ISSN 0027-8424 ([Link] PMC 8812535 ([Link]
[Link]/pmc/articles/PMC8812535). PMID 35074877 ([Link]
[Link]/35074877).
164. Organ, Chris; Nunn, Charles L.; Machanda, Zarin; Wrangham, Richard W. (August 30,
2011). "Phylogenetic rate shifts in feeding time during the evolution of Homo" ([Link]
[Link]/pmc/articles/PMC3167533). Proceedings of the National Academy of
Sciences of the United States of America. 108 (35): 14555–14559.
Bibcode:2011PNAS..10814555O ([Link]
O). doi:10.1073/pnas.1107806108 ([Link]
ISSN 0027-8424 ([Link] PMC 3167533 ([Link]
[Link]/pmc/articles/PMC3167533). PMID 21873223 ([Link]
ov/21873223).
165. David-Barrett, T.; Dunbar, R. I. M. (2013). "Processing Power Limits Social Group Size:
Computational Evidence for the Cognitive Costs of Sociality" ([Link]
mc/articles/PMC3712454). Proceedings of the Royal Society of London B: Biological
Sciences. 280 (1765) 20131151. doi:10.1098/rspb.2013.1151 ([Link]
pb.2013.1151). PMC 3712454 ([Link]
PMID 23804623 ([Link]
166. Bown & Rose 1987.
167. Barton, Robert A.; Venditti, Chris (October 20, 2014). "Rapid Evolution of the Cerebellum in
Humans and Other Great Apes" ([Link] Current
Biology. 24 (20): 2440–2444. Bibcode:2014CBio...24.2440B ([Link]
abs/2014CBio...24.2440B). doi:10.1016/[Link].2014.08.056 ([Link]
2014.08.056). ISSN 0960-9822 ([Link]
PMID 25283776 ([Link] S2CID 5041106 ([Link]
[Link]/CorpusID:5041106).
168. Starowicz-Filip, Anna; Milczarek, Olga; Kwiatkowski, Stanisław; Bętkowska-Korpała,
Barbara; Prochwicz, Katarzyna (2013). "Cerebellar cognitive affective syndrome CCAS – a
case report" ([Link] Archives of Psychiatry and
Psychotherapy. 15 (3): 57–64. doi:10.12740/APP/18666 ([Link]
2F18666).
169. Yu, Feng; Jiang, Qing-jun; Sun, Xi-yan; Zhang, Rong-wei (August 22, 2014). "A new case of
complete primary cerebellar agenesis: Clinical and imaging findings in a living patient" (http
s://[Link]/pmc/articles/PMC4614135). Brain. 138 (Pt 6): e353.
doi:10.1093/brain/awu239 ([Link] ISSN 1460-2156
([Link] PMC 4614135 ([Link]
mc/articles/PMC4614135). PMID 25149410 ([Link]
170. Weaver, Anne H. (March 8, 2005). "Reciprocal evolution of the cerebellum and neocortex in
fossil humans" ([Link] Proceedings of the
National Academy of Sciences of the United States of America. 102 (10): 3576–3580.
Bibcode:2005PNAS..102.3576W ([Link]
W). doi:10.1073/pnas.0500692102 ([Link]
ISSN 0027-8424 ([Link] PMC 553338 ([Link]
[Link]/pmc/articles/PMC553338). PMID 15731345 ([Link]
15731345).
171. Klein, Stefan (2014). Survival of the Nicest. The Experiment. ISBN 978-1-61519-090-4.
172. Sallet, J.; Mars, R. B.; Noonan, M. P.; Andersson, J. L.; O'Reilly, J. K.; Jbabdi, S.; Croxson,
P. L.; Jenkinson, M.; Miller, K. L. (November 4, 2011). "Social Network Size Affects Neural
Circuits in Macaques". Science. 334 (6056): 697–700. Bibcode:2011Sci...334..697S (https://
[Link]/abs/2011Sci...334..697S). doi:10.1126/science.1210027 ([Link]
rg/10.1126%2Fscience.1210027). ISSN 0036-8075 ([Link]
075). PMID 22053054 ([Link] S2CID 206536017 (http
s://[Link]/CorpusID:206536017).
173. Dunbar, R. I. M. (1992). "Neocortex size as a constraint on group size in primates" ([Link]
[Link]/10.1016%2F0047-2484%2892%2990081-j). Journal of Human Evolution. 22 (6): 469–
493. Bibcode:1992JHumE..22..469D ([Link]
9D). doi:10.1016/0047-2484(92)90081-j ([Link]
0081-j).
174. Shultz, Susanne; Dunbar, Robin (December 14, 2010). "Encephalization is not a universal
macroevolutionary phenomenon in mammals but is associated with sociality" ([Link]
[Link]/pmc/articles/PMC3003036). Proceedings of the National Academy of
Sciences of the United States of America. 107 (50): 21582–21586.
Bibcode:2010PNAS..10721582S ([Link]
S). doi:10.1073/pnas.1005246107 ([Link]
ISSN 0027-8424 ([Link] PMC 3003036 ([Link]
[Link]/pmc/articles/PMC3003036). PMID 21098277 ([Link]
ov/21098277).
175. Richard, Potts (1998). "Environmental Hypotheses of Hominin Evolution". American Journal
of Physical Anthropology. 107 (S27): 93–136. doi:10.1002/(sici)1096-8644(1998)107:27+
<93::aid-ajpa5>[Link];2-x ([Link]
107%3A27%2B%3C93%3A%3Aaid-ajpa5%[Link]%3B2-x). PMID 9881524 ([Link]
[Link]/9881524).
176. Richerson, Peter J. (2006). Not by Genes Alone: How Culture Transformed Human
Evolution. University of Chicago Press. ISBN 0-226-71212-5. OCLC 642503808 ([Link]
[Link]/oclc/642503808).
177. Mesoudi, Alex; Whiten, Andrew; Laland, Kevin N. (August 2006). "Towards a unified science
of cultural evolution". Behavioral and Brain Sciences. 29 (4): 329–347.
doi:10.1017/s0140525x06009083 ([Link]
ISSN 0140-525X ([Link] PMID 17094820 ([Link]
[Link]/17094820).
178. Berger, L. R.; Hawks, J.; Dirks, P. H.; Elliott, M.; Roberts, E. M. (May 9, 2017). "Homo naledi
and Pleistocene hominin evolution in subequatorial Africa" ([Link]
c/articles/PMC5423770). eLife. 6 e24234. doi:10.7554/elife.24234 ([Link]
2Felife.24234). PMC 5423770 ([Link]
PMID 28483041 ([Link]
179. Strait D, Grine F, Fleagle J (2015). Analyzing Hominin Hominin Phylogeny: Cladistic
Approach ([Link] Springer. pp. 1989–2014
(cladogram p. 2006). ISBN 978-3-642-39978-7..
180. Schrenk, F.; Kullmer, O.; Bromage, T. (2007). "Chapter 9: The Earliest Putative Homo
Fossils". In Henke, W.; Tattersall, I. (eds.). Handbook of Paleoanthropology. pp. 1611–1631.
doi:10.1007/978-3-540-33761-4_52 ([Link]
181. Spoor, F.; Gunz, P.; Neubauer, S.; Stelzer, S.; Scott, N.; Kwekason, A.; Dean, M. C. (March
2015). "Reconstructed Homo habilis type OH 7 suggests deep-rooted species diversity in
early Homo". Nature. 519 (7541): 83–6. Bibcode:2015Natur.519...83S ([Link]
[Link]/abs/2015Natur.519...83S). doi:10.1038/nature14224 ([Link]
ature14224). PMID 25739632 ([Link] S2CID 4470282
([Link]
182. Schuster, A. M. (1997). "Earliest Remains of Genus Homo" ([Link]
9701/newsbriefs/[Link]). Archaeology. 50 (1). Archived ([Link]
150317002614/[Link] from the original
on March 17, 2015. Retrieved March 5, 2015. The line to the earliest members of Homo was
derived from Australopithecus, a genus which had separated from the Chimpanzee–human
last common ancestor by late Miocene or early Pliocene times.
183. Indriati, E.; Swisher, C. C.; Lepre, C.; Quinn, R. L.; Suriyanto, R. A.; et al. (2011). "The age
of the 20 meter Solo River terrace, Java, Indonesia and the survival of Homo erectus in
Asia" ([Link] PLoS One. 6 (6) e21562.
Bibcode:2011PLoSO...621562I ([Link]
doi:10.1371/[Link].0021562 ([Link]
PMC 3126814 ([Link] PMID 21738710
([Link]
Full list of authors
Etty Indriati
Carl C. Swisher III
Christopher Lepre
Rhonda L. Quinn
Rusyad A. Suriyanto
Agus T. Hascaryo
Rainer Grün
Craig S. Feibel
Briana L. Pobiner
Maxime Aubert
Wendy Lees
Susan C. Antón
184. Zeitoun, V. (September 2003). "High occurrence of a basicranial feature in Homo erectus:
Anatomical description of the preglenoid tubercle" ([Link]
The Anatomical Record Part B: The New Anatomist. 274 (1): 148–56.
doi:10.1002/ar.b.10028 ([Link] PMID 12964205 ([Link]
[Link]/12964205).
185. Dembo, M.; Matzke, N. J.; Mooers, A. Ø.; Collard, M. (August 2015). "Bayesian analysis of a
morphological supermatrix sheds light on controversial fossil hominin relationships" (https://
[Link]/pmc/articles/PMC4528516). Proceedings. Biological Sciences. 282
(1812) 20150943. Bibcode:2015PBioS.28250943D ([Link]
PBioS.28250943D). doi:10.1098/rspb.2015.0943 ([Link]
3). PMC 4528516 ([Link]
PMID 26202999 ([Link]
186. Dembo, M.; Radovčić, D.; et al. (August 2016). "The evolutionary relationships and age of
Homo naledi: An assessment using dated Bayesian phylogenetic methods". Journal of
Human Evolution. 97: 17–26. Bibcode:2016JHumE..97...17D ([Link]
abs/2016JHumE..97...17D). doi:10.1016/[Link].2016.04.008 ([Link]
hevol.2016.04.008). hdl:2164/8796 ([Link] PMID 27457542
([Link]
Full list of authors
Mana Dembo
Davorka Radovčić
Heather M. Garvinc
Myra F. Laird
Lauren Schroeder
Jill E. Scott
Juliet Brophy
Rebecca R. Ackermann
Chares M. Musiba
Darryl J. de Ruiter
Arne Ø. Mooers
Mark Collard
187. Mounier, A.; Caparros, M. (2015). "The phylogenetic status of Homo heidelbergensis – a
cladistic study of Middle Pleistocene hominins". BMSAP (in French). 27 (3–4): 110–134.
doi:10.1007/s13219-015-0127-4 ([Link]
ISSN 0037-8984 ([Link] S2CID 17449909 ([Link]
[Link]/CorpusID:17449909).
188. Harrison, N. (May 1, 2019). The Origins of Europeans and Their Pre-Historic Innovations
from 6 Million to 10,000 BCE: From 6 Million to 10,000 BCE ([Link]
s?id=bL2XDwAAQBAJ&q=Sahelanthropus+tchadensis&pg=PA7). Algora Publishing.
ISBN 978-1-62894-379-5.
189. Rogers, A. R.; Harris, N. S.; Achenbach, A. A. (February 2020). "Neanderthal-Denisovan
ancestors interbred with a distantly related hominin" ([Link]
es/PMC7032934). Science Advances. 6 (8) eaay5483. Bibcode:2020SciA....6.5483R (http
s://[Link]/abs/2020SciA....6.5483R). doi:10.1126/sciadv.aay5483 ([Link]
[Link]/10.1126%2Fsciadv.aay5483). PMC 7032934 ([Link]
s/PMC7032934). PMID 32128408 ([Link]
190. Tanner, Nancy Makepeace (1981). On Becoming Human ([Link]
d=_j45AAAAIAAJ). CUP Archive. p. 202 ([Link]
J&pg=PA202). ISBN 978-0-521-28028-0.
191. Reno, Philip L.; Lovejoy, C. Owen (April 28, 2015). "From Lucy to Kadanuumuu: Balanced
analyses of Australopithecus afarensis assemblages confirm only moderate skeletal
dimorphism" ([Link] PeerJ. 3 e925.
Bibcode:2015PeerJ...3.e925R ([Link]
doi:10.7717/peerj.925 ([Link] ISSN 2167-8359 ([Link]
[Link]/issn/2167-8359). PMC 4419524 ([Link]
MC4419524). PMID 25945314 ([Link]
192. Lovejoy, C. Owen (October 2, 2009). "Reexamining Human Origins in Light of Ardipithecus
ramidus" ([Link] (PDF). Science. 326
(5949): 74–74e8. Bibcode:2009Sci...326...74L ([Link]
326...74L). doi:10.1126/science.1175834 ([Link]
ISSN 0036-8075 ([Link] PMID 19810200 ([Link]
[Link]/19810200). S2CID 42790876 ([Link]
D:42790876). Archived ([Link]
ord/211449/files/PAL_E4439.pdf) (PDF) from the original on February 24, 2021. Retrieved
December 6, 2019.
193. Young, Richard W. (January 2003). "Evolution of the human hand: The role of throwing and
clubbing" ([Link] Journal of Anatomy.
202 (1): 165–174. doi:10.1046/j.1469-7580.2003.00144.x ([Link]
-7580.2003.00144.x). ISSN 0021-8782 ([Link]
PMC 1571064 ([Link] PMID 12587931
([Link]
194. Ittyerah, Miriam (2013). Hand Preference and Hand Ability: Evidence from studies in Haptic
Cognition. John Benjamins Publishing. pp. 37–38. ISBN 978-90-272-7164-8.
195. Wilson, Frank R. "The Hand How Its Use Shapes the Brain, Language, and Human Culture"
([Link] The New York Times. Archived (h
ttps://[Link]/web/20170828024550/[Link]
[Link]) from the original on August 28, 2017. Retrieved July 2, 2017.
196. Kuzawa, Christopher W.; Chugani, Harry T.; Grossman, Lawrence I.; Lipovich, Leonard;
Muzik, Otto; Hof, Patrick R.; Wildman, Derek E.; Sherwood, Chet C.; Leonard, William R.;
Lange, Nicholas (August 25, 2014). "Metabolic costs and evolutionary implications of human
brain development" ([Link] Proceedings
of the National Academy of Sciences of the United States of America. 111 (36): 13010–
13015. Bibcode:2014PNAS..11113010K ([Link]
3010K). doi:10.1073/pnas.1323099111 ([Link]
PMC 4246958 ([Link] PMID 25157149
([Link]
197. Pontzer, Herman; Brown, Mary H.; Raichlen, David A.; Dunsworth, Holly; Hare, Brian;
Walker, Kara; Luke, Amy; Dugas, Lara R.; Durazo-Arvizu, Ramon; Schoeller, Dale; Plange-
Rhule, Jacob; Bovet, Pascal; Forrester, Terrence E.; Lambert, Estelle V.; Thompson, Melissa
Emery; Shumaker, Robert W.; Ross, Stephen P. (May 4, 2016). "Metabolic acceleration and
the evolution of human brain size and life history" ([Link]
s/PMC4942851). Nature. 533 (7603): 390–392. Bibcode:2016Natur.533..390P ([Link]
[Link]/abs/2016Natur.533..390P). doi:10.1038/nature17654 ([Link]
38%2Fnature17654). PMC 4942851 ([Link]
1). PMID 27144364 ([Link]
198. Rantala, Markus J. (August 20, 2007). "Evolution of nakedness in Homo sapiens" ([Link]
[Link]/doi/full/10.1111/j.1469-7998.2007.00295.x). Journal of
Zoology. 273 (1): 1–7. doi:10.1111/j.1469-7998.2007.00295.x ([Link]
469-7998.2007.00295.x). Archived ([Link]
[Link]/doi/full/10.1111/j.1469-7998.2007.00295.x) from the
original on November 5, 2022. Retrieved November 5, 2022.
199. Differential modularity of the mammalian Engrailed 1 enhancer network directs sweat gland
development ([Link]
4)
200. Humans Evolved to Be the Water-Saving Ape | Duke Today ([Link]
humans-evolved-be-water-saving-ape)
201. Why Humans Sleep Less Than Their Primate Relatives ([Link]
cience-nature/why-humans-sleep-less-than-their-primate-relatives-180980018/)
202. Potter, Lisa Marie (September 8, 2015). "Shouldering the Burden of Evolution" ([Link]
[Link]/news/2015/09/131526/shouldering-burden-evolution). UCSF News Center.
University of California San Francisco. Archived ([Link]
36/[Link] from the
original on May 8, 2023. Retrieved May 8, 2023.
203. Weber, G.W.; Lukeneder, A.; Harzhauser, M. (February 28, 2022). "The microstructure and
origin of the Venus of Willendorf" ([Link]
Scientific Reports. 12 (2926). Nature: 2926. doi:10.1038/s41598-022-06799-z ([Link]
g/10.1038%2Fs41598-022-06799-z). PMC 8885675 ([Link]
es/PMC8885675). PMID 35228605 ([Link]
204. Ko, Kwang Hyun (2016). "Origins of human intelligence: The chain of tool-making and brain
evolution" ([Link]
XXII_1_Ko.pdf) (PDF). Anthropological Notebooks. 22 (1): 5–22. Archived ([Link]
[Link]/web/20160817210720/[Link]
ogical_Notebooks_XXII_1_Ko.pdf) (PDF) from the original on August 17, 2016. Retrieved
August 8, 2016.
205. Jabr, Ferris (July 18, 2012). "Does Thinking Really Hard Burn More Calories?" ([Link]
[Link]/article/thinking-hard-calories/). Scientific American. ISSN 0036-8733
([Link] Archived ([Link]
03113428/[Link] from the original
on May 3, 2015. Retrieved May 3, 2015.
206. Gibbons, Ann (May 29, 1998). "Solving the Brain's Energy Crisis". Science. 280 (5368):
1345–1347. doi:10.1126/science.280.5368.1345 ([Link]
68.1345). ISSN 0036-8075 ([Link] PMID 9634409 (htt
ps://[Link]/9634409). S2CID 28464775 ([Link]
CorpusID:28464775).
207. Robinson 2008, p. 398 ([Link]
208. Harmand, Sonia; Lewis, Jason E.; Feibel, Craig S.; Lepre, Christopher J.; Prat, Sandrine;
Lenoble, Arnaud; Boës, Xavier; Quinn, Rhonda L.; Brenet, Michel (May 20, 2015). "3.3-
million-year-old stone tools from Lomekwi 3, West Turkana, Kenya" ([Link]
1159189). Nature. 521 (7552): 310–315. Bibcode:2015Natur.521..310H ([Link]
[Link]/abs/2015Natur.521..310H). doi:10.1038/nature14464 ([Link]
nature14464). ISSN 0028-0836 ([Link]
PMID 25993961 ([Link] S2CID 1207285 ([Link]
[Link]/CorpusID:1207285).
209. Semaw, Sileshi; Rogers, Michael J.; Quade, Jay; Renne, Paul R.; Butler, Robert F.;
Dominguez-Rodrigo, Manuel; Stout, Dietrich; Hart, William S.; Pickering, Travis; Simpson,
Scott W. (August 1, 2003). "2.6-Million-year-old stone tools and associated bones from
OGS-6 and OGS-7, Gona, Afar, Ethiopia". Journal of Human Evolution. 45 (2): 169–177.
Bibcode:2003JHumE..45..169S ([Link]
doi:10.1016/S0047-2484(03)00093-9 ([Link]
093-9). ISSN 0047-2484 ([Link] PMID 14529651 (http
s://[Link]/14529651).
210. Freeman & Herron 2007, pp. 786–788.
211. 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" ([Link] Proceedings of the
National Academy of Sciences of the United States of America. 111 (1): 121–124.
Bibcode:2014PNAS..111..121W ([Link]
doi:10.1073/pnas.1316014110 ([Link] ISSN 0027-
8424 ([Link] PMC 3890866 ([Link]
gov/pmc/articles/PMC3890866). PMID 24344276 ([Link]
6).
212. Diamond 1999, p. 39.
213. Bar-Yosef, Ofer (October 2002). "The Upper Paleolithic Revolution". Annual Review of
Anthropology. 31: 363–393. doi:10.1146/[Link].31.040402.085416 ([Link]
0.1146%[Link].31.040402.085416). ISSN 0084-6570 ([Link]
g/issn/0084-6570).
214. Oppenheimer, Stephen (2012). Out of Eden: The Peopling of the World (New ed.).
Robinson.
215. Henshilwood, C. S.; d'Errico, F.; Yates, R.; et al. (2002). "Emergence of Modern Human
Behavior: Middle Stone Age Engravings from South Africa". Science. 295 (5558): 1278–
1280. Bibcode:2002Sci...295.1278H ([Link]
H). doi:10.1126/science.1067575 ([Link]
PMID 11786608 ([Link] S2CID 31169551 ([Link]
[Link]/CorpusID:31169551).
216. Henshilwood, Christopher; Marean, Curtis (2003). "The Origin of Modern Human Behavior:
Critique of the Models and Their Test Implications". Current Anthropology. 44 (5): 627–651.
doi:10.1086/377665 ([Link] PMID 14971366 ([Link]
[Link]/14971366). S2CID 11081605 ([Link]
081605).
217. Backwell, L.; d'Errico, F.; Wadley, L. (2008). "Middle Stone Age bone tools from the
Howiesons Poort layers, Sibudu Cave, South Africa". Journal of Archaeological Science. 35
(6): 1566–1580. Bibcode:2008JArSc..35.1566B ([Link]
Sc..35.1566B). doi:10.1016/[Link].2007.11.006 ([Link]
218. Nowell, April (October 2010). "Defining Behavioral Modernity in the Context of Neandertal
and Anatomically Modern Human Populations". Annual Review of Anthropology. 39 (1):
437–452. Bibcode:2010ARAnt..39..437N ([Link]
9..437N). doi:10.1146/[Link].012809.105113 ([Link]
nthro.012809.105113). ISSN 0084-6570 ([Link]
219. d'Errico, Francesco; Stringer, Chris B. (April 12, 2011). "Evolution, revolution or saltation
scenario for the emergence of modern cultures?" ([Link]
PMC3049097). Philosophical Transactions of the Royal Society B. 366 (1567): 1060–1069.
doi:10.1098/rstb.2010.0340 ([Link] ISSN 0962-8436
([Link] PMC 3049097 ([Link]
mc/articles/PMC3049097). PMID 21357228 ([Link]
220. Chatterjee, Rhitu (March 15, 2018). "Scientists Are Amazed By Stone Age Tools They Dug
Up In Kenya" ([Link]
are-amazed-by-stone-age-tools-they-dug-up-in-kenya). NPR. Archived ([Link]
org/web/20180315193655/[Link]
96/scientists-are-amazed-by-stone-age-tools-they-dug-up-in-kenya) from the original on
March 15, 2018. Retrieved March 15, 2018.
221. Yong, Ed (March 15, 2018). "A Cultural Leap at the Dawn of Humanity – New finds from
Kenya suggest that humans used long-distance trade networks, sophisticated tools, and
symbolic pigments right from the dawn of our species" ([Link]
archive/2018/03/a-deeper-origin-of-complex-human-cultures/555674/). The Atlantic.
Archived ([Link]
e/archive/2018/03/a-deeper-origin-of-complex-human-cultures/555674/) from the original on
November 17, 2020. Retrieved March 15, 2018.
222. Brooks, A. S.; Yellen, J. E.; Potts, R.; Behrensmeyer, A. K.; Deino, A. L.; Leslie, D. E.;
Ambrose, S. H.; Ferguson, J. R.; d'Errico, F.; Zipkin, A. M.; Whittaker, S.; Post, J.; Veatch, E.
G.; Foecke, K.; Clark, J. B. (2018). "Long-distance stone transport and pigment use in the
earliest Middle Stone Age" ([Link] Science. 360
(6384): 90–94. Bibcode:2018Sci...360...90B ([Link]
0...90B). doi:10.1126/science.aao2646 ([Link]
PMID 29545508 ([Link] S2CID 14051717 ([Link]
[Link]/CorpusID:14051717).
223. Rasmussen, Morten; Guo, Xiaosen; Wang, Yong; Lohmueller, Kirk E.; Rasmussen, Simon;
Albrechtsen, Anders; et al. (2011). "An Aboriginal Australian Genome Reveals Separate
Human Dispersals into Asia" ([Link]
Science. 334 (6052): 94–98. Bibcode:2011Sci...334...94R ([Link]
s/2011Sci...334...94R). doi:10.1126/science.1211177 ([Link]
11177). PMC 3991479 ([Link]
PMID 21940856 ([Link]
224. Ambrose, Stanley H. (March 2, 2001). "Paleolithic Technology and Human Evolution".
Science. 291 (5509): 1748–1753. Bibcode:2001Sci...291.1748A ([Link]
du/abs/2001Sci...291.1748A). doi:10.1126/science.1059487 ([Link]
nce.1059487). ISSN 0036-8075 ([Link]
PMID 11249821 ([Link] S2CID 6170692 ([Link]
[Link]/CorpusID:6170692).
225. Mellars, P. (2010). "Neanderthal symbolism and ornament manufacture: The bursting of a
bubble?" ([Link] Proc Natl Acad Sci
USA. 107 (47): 20147–20148. Bibcode:2010PNAS..10720147M ([Link]
du/abs/2010PNAS..10720147M). doi:10.1073/pnas.1014588107 ([Link]
Fpnas.1014588107). PMC 2996706 ([Link]
6). PMID 21078972 ([Link]
226. Cochran & Harpending 2009.
227. Bender, Andrea (August 8, 2019). "The Role of Culture and Evolution for Human Cognition"
([Link] Topics in Cognitive Science. 12 (4): 1403–1420.
doi:10.1111/tops.12449 ([Link] hdl:1956/22031 ([Link]
[Link]/1956%2F22031). ISSN 1756-8757 ([Link]
PMID 31392845 ([Link]
228. Fu, Q.; Hajdinjak, M.; Moldovan, O. T.; Constantin, S.; Mallick, S.; Skoglund, Pontus;
Patterson, N.; Rohland, N.; Lazaridis, I.; Nickel, B.; Viola, B.; Prüfer, Kay; Meyer, M.; Kelso,
J.; Reich, D.; Pääbo, S. (2015). "An early modern human from Romania with a recent
Neanderthal ancestor" ([Link] Nature.
524 (7564): 216–219. Bibcode:2015Natur.524..216F ([Link]
5Natur.524..216F). doi:10.1038/nature14558 ([Link]
PMC 4537386 ([Link] PMID 26098372
([Link]
229. Medical Research Council [UK] (November 21, 2009). "Brain Disease 'Resistance Gene'
evolves in Papua New Guinea community; could offer insights into CJD" ([Link]
[Link]/releases/2009/11/[Link]). ScienceDaily. Archived ([Link]
[Link]/web/20190608194846/[Link]
[Link]) from the original on June 8, 2019. Retrieved November 22, 2009.
230. Mead, S.; Whitfield, J.; Poulter, M.; et al. (2009). "A Novel Protective Prion Protein Variant
that Colocalizes with Kuru Exposure" ([Link]
[Link]) (PDF). The New England Journal of Medicine. 361 (21): 2056–2065.
doi:10.1056/NEJMoa0809716 ([Link]
PMID 19923577 ([Link] Archived ([Link]
org/web/20180720215808/[Link]
(PDF) from the original on July 20, 2018. Retrieved August 20, 2019.
Full list of authors
Simon Mead
Jerome Whitfield
Mark Poulter
Paresh Shah
James Uphill
Tracy Campbell
Huda Al-Dujaily
Holger Hummerich
Jon Beck
Charles A. Mein
Claudio Verzilli
John Whittaker
Michael P. Alpers
John Collinge
231. Corbett, Stephen; Morin-Papunen, Laure (July 2013). "The Polycystic Ovary Syndrome and
recent human evolution" ([Link]
Molecular and Cellular Endocrinology. 373 (1–2): 39–50. doi:10.1016/[Link].2013.01.001 (htt
ps://[Link]/10.1016%[Link].2013.01.001). PMID 23352610 ([Link]
gov/23352610). S2CID 13078403 ([Link]
Archived ([Link]
eve/pii/S030372071300004X) from the original on May 25, 2021. Retrieved October 8,
2020.
232. Diamond 1999.
233. Courtiol, Alexandre; Tropf, Felix C.; Mills, Melinda C. (July 12, 2016). "When genes and
environment disagree: Making sense of trends in recent human evolution" ([Link]
[Link]/pmc/articles/PMC4948334). Proceedings of the National Academy of Sciences
of the United States of America. 113 (28): 7693–7695. Bibcode:2016PNAS..113.7693C (http
s://[Link]/abs/2016PNAS..113.7693C). doi:10.1073/pnas.1608532113 (http
s://[Link]/10.1073%2Fpnas.1608532113). ISSN 0027-8424 ([Link]
n/0027-8424). PMC 4948334 ([Link]
PMID 27402758 ([Link]
234. Markov, Alexander V.; Markov, Mikhail A. (June 2020). "Runaway brain-culture coevolution
as a reason for larger brains: Exploring the "cultural drive" hypothesis by computer
modeling" ([Link] Ecology and Evolution.
10 (12): 6059–6077. Bibcode:2020EcoEv..10.6059M ([Link]
0EcoEv..10.6059M). doi:10.1002/ece3.6350 ([Link]
ISSN 2045-7758 ([Link] PMC 7319167 ([Link]
[Link]/pmc/articles/PMC7319167). PMID 32607213 ([Link]
ov/32607213).
235. Stout, Dietrich (April 12, 2011). "Stone toolmaking and the evolution of human culture and
cognition" ([Link] Philosophical
Transactions of the Royal Society B: Biological Sciences. 366 (1567): 1050–1059.
doi:10.1098/rstb.2010.0369 ([Link] ISSN 0962-8436
([Link] PMC 3049103 ([Link]
mc/articles/PMC3049103). PMID 21357227 ([Link]
236. Byars, S. G.; Ewbank, D.; Govindaraju, D. R.; Stearns, S. C. (2009). "Natural selection in a
contemporary human population" ([Link]
Proceedings of the National Academy of Sciences of the United States of America. 107
(suppl_1): 1787–1792. Bibcode:2010PNAS..107.1787B ([Link]
010PNAS..107.1787B). doi:10.1073/pnas.0906199106 ([Link]
06199106). PMC 2868295 ([Link]
PMID 19858476 ([Link]
237. Editors of the American Heritage Dictionaries, ed. (2006). More Word Histories and
Mysteries: From Aardvark to Zombie ([Link]
Boston: Houghton Mifflin. ISBN 978-0-618-71681-4. LCCN 2006020835 ([Link]
v/2006020835). OCLC 70199867 ([Link]
238. "Nested Hierarchies, the Order of Nature: Carolus Linnaeus" ([Link]
evolibrary/article/0_0_0/history_05). Understanding Evolution: The History of Evolutionary
Thought. The University of California at Berkeley. Archived ([Link]
90802105432/[Link] from the
original on August 2, 2019. Retrieved August 2, 2019.
239. Darwin 1859, pp. 367–404 ([Link]
ype=text&pageseq=506).
240. Darwin 1981.
241. Montgomery 1988, pp. 95–96.
242. Shipman, Pat; Storm, Paul (2002). "Missing links: Eugène Dubois and the origins of
paleoanthropology". Evolutionary Anthropology: Issues, News, and Reviews. 11 (3): 108–
116. doi:10.1002/evan.10021 ([Link] ISSN 1520-6505 (htt
ps://[Link]/issn/1520-6505). S2CID 84889438 ([Link]
CorpusID:84889438).
243. Dart, Raymond (February 7, 1925). "Australopithecus africanus: The Man-Ape of South
Africa" ([Link] Nature.
115 (2884): 195–199. Bibcode:1925Natur.115..195D ([Link]
5Natur.115..195D). doi:10.1038/115195a0 ([Link]
ISSN 0028-0836 ([Link] S2CID 4142569 ([Link]
[Link]/CorpusID:4142569).
244. Johanson & Edey 1981, pp. 20–22, 184–185.
245. Cartmill, Matt; Smith, Fred H.; Brown, Kaye B. (2009). The Human Lineage. Wiley-
Blackwell. p. 151 ([Link]
ISBN 978-0-471-21491-5.
246. Johanson & Edey 1981, p. 22.
247. Shreeve, Jamie (July 2010). "The Evolutionary Road" ([Link]
6024510/[Link] National
Geographic. ISSN 0027-9358 ([Link] Archived from
the original ([Link] on
January 26, 2016. Retrieved May 28, 2015.
248. Berger, L. R.; Hawks, J.; de Ruiter, D. J.; Churchill, S. E.; et al. (September 10, 2015).
"Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa"
([Link] eLife. 4 e09560.
Bibcode:2015eLife...409560B ([Link]
doi:10.7554/eLife.09560 ([Link] PMC 4559886 ([Link]
[Link]/pmc/articles/PMC4559886). PMID 26354291 ([Link]
[Link]/26354291).
Full list of authors
Lee R Berger
John Hawks
Darryl J de Ruiter
Steven E Churchill
Peter Schmid
Lucas K Delezene
Tracy L Kivell
Heather M Garvin
Scott A Williams
Jeremy M DeSilva
Matthew M Skinner
Charles M Musiba
Noel Cameron
Trenton W Holliday
William Harcourt-Smith
Rebecca R Ackermann
Markus Bastir
Barry Bogin
Debra Bolter
Juliet Brophy
Zachary D Cofran
Kimberly A Congdon
Andrew S Deane
Mana Dembo
Michelle Drapeau
Marina C Elliott
Elen M Feuerriegel
Daniel Garcia-Martinez
David J Green
Alia Gurtov
Joel D Irish
Ashley Kruger
Myra F Laird
Damiano Marchi
Marc R Meyer
Shahed Nalla
Enquye W Negash
Caley M Orr
 Davorka Radovcic
Lauren Schroeder
Jill E Scott
Zachary Throckmorton
Matthew W Tocheri
Caroline VanSickle
Christopher S Walker
Pianpian Wei
Bernhard Zipfel
249. Shreeve, Jamie (September 10, 2015). "This Face Changes the Human Story. But How?" (h
ttps://[Link]/web/20180414054102/[Link]
150910-human-evolution-change/). National Geographic News. Archived from the original (h
ttp://[Link]/2015/09/150910-human-evolution-change/) on April 14,
2018. Retrieved September 10, 2015.
250. Dirks et al. (2017): between 335 and 236 ka. The lower limit of 236 ka is due to optically
stimulated luminescence dating of sediments with U-Th and palaeomagnetic analyses of
flowstones; the upper limit of 335 ka is due to U-series and electron spin resonance (US-
+82
ESR) dating of two H. naledi teeth, to 253 −70 ka, for an estimated age of the fossils of
+82
253 −17 ka.
251. Dirks PH, Roberts EM, Hilbert-Wolf H, Kramers JD, et al. (May 9, 2017). "The age of Homo
naledi and associated sediments in the Rising Star Cave, South Africa" ([Link]
[Link]/pmc/articles/PMC5423772). eLife. 6 e24231. Bibcode:2017eLife...624231D (http
s://[Link]/abs/2017eLife...624231D). doi:10.7554/eLife.24231 ([Link]
g/10.7554%2FeLife.24231). PMC 5423772 ([Link]
5423772). PMID 28483040 ([Link]
Full list of authors
Paul H.G.M. Dirks
Eric M. Roberts
Hannah Hilbert-Wolf
Jan D. Kramers
John Hawks
Anthony Dosseto
Mathieu Duval
Marina Elliott
Mary Evans
Rainer Grün
John Hellstrom
Andy I.R. Herries
Renaud Joannes-Boyau
Tebogo V. Makhubela
Christa J. Placzek
Jessie Robbins
Carl Spandler
Jelle Wiersma
Jon Woodhead
Lee R. Berger
252. Sarich, V. M.; Wilson, A. C. (1967). "Immunological time scale for hominid evolution".
Science. 158 (3805): 1200–1204. Bibcode:1967Sci...158.1200S ([Link]
du/abs/1967Sci...158.1200S). doi:10.1126/science.158.3805.1200 ([Link]
2Fscience.158.3805.1200). PMID 4964406 ([Link]
S2CID 7349579 ([Link]
253. M'charek 2005, p. 96 ([Link]
254. Trent 2005, p. 6 ([Link]
255. Wade, Nicholas (May 18, 2006). "Two Splits Between Human and Chimp Lines Suggested"
([Link] The New York Times.
Archived ([Link]
8/science/[Link]) from the original on May 18, 2022. Retrieved April 20, 2015.
256. Behar, D. M.; Villems, R.; Soodyall, H.; et al. (May 9, 2008). "The Dawn of Human
Matrilineal Diversity" ([Link] American
Journal of Human Genetics. 82 (5): 1130–1140. Bibcode:2008AmJHG..82.1130B ([Link]
[Link]/abs/2008AmJHG..82.1130B). doi:10.1016/[Link].2008.04.002 ([Link]
[Link]/10.1016%[Link].2008.04.002). ISSN 0002-9297 ([Link]
02-9297). PMC 2427203 ([Link]
PMID 18439549 ([Link]
Full list of authors
Doron M. Behar
Richard Villems
Himla Soodyall
Jason Blue-Smith
Luisa Pereira
Ene Metspalu
Rosaria Scozzari
Heeran Makkan
Shay Tzur
David Comas
Jaume Bertranpetit
Lluis Quintana-Murci
Chris Tyler-Smith
R. Spencer Wells
Saharon Rosset
The Genographic Consortium
257. Gonder, M. K.; Mortensen, H. M.; Reed, F. A.; de Sousa, A.; Tishkoff, S. A. (March 2007).
"Whole-mtDNA Genome Sequence Analysis of Ancient African Lineages" ([Link]
1093%2Fmolbev%2Fmsl209). Molecular Biology and Evolution. 24 (3): 757–768.
doi:10.1093/molbev/msl209 ([Link] ISSN 0737-
4038 ([Link] PMID 17194802 ([Link]
[Link]/17194802).
258. Tishkoff, S. A.; Reed, F. A.; Friedlaender, F. R.; et al. (2009). "The Genetic Structure and
History of Africans and African Americans" ([Link]
947357). Science. 324 (5930) (published May 22, 2009): 1035–1044.
Bibcode:2009Sci...324.1035T ([Link]
doi:10.1126/science.1172257 ([Link] PMC 2947357
([Link] PMID 19407144 ([Link]
[Link]/19407144).
Full list of authors
Sarah A. Tishkoff
Floyd A. Reed
Françoise R. Friedlaender
Christopher Ehret
Alessia Ranciaro
Alain Froment
Jibril B. Hirbo
Agnes A. Awomoyi
Jean-Marie Bodo
Ogobara Doumbo
Muntaser Ibrahim
Abdalla T. Juma
Maritha J. Kotze
Godfrey Lema
Jason H. Moore
Holly Mortensen
Thomas B. Nyambo
Sabah A. Omar
Kweli Powell
Gideon S. Pretorius
Michael W. Smith
Mahamadou A. Thera
Charles Wambebe
James L. Weber
Scott M. Williams
259. Templeton, Alan R. (2005). "Haplotype Trees and Modern Human Origins". American
Journal of Physical Anthropology. 128 (Supplement 41: Yearbook of Physical Anthropology):
33–59. Bibcode:2005AJPA..128S..33T ([Link]
33T). doi:10.1002/ajpa.20351 ([Link] ISSN 0002-9483 (htt
ps://[Link]/issn/0002-9483). PMID 16369961 ([Link]
v/16369961).
260. deMenocal, Peter B. (2016). "Climate Shocks". Scientific American. Vol. 25, no. 4.
261. Barras, Colin (February 6, 2016). "Stone tools hint humans reached Asia much earlier". New
Scientist.
262. Wolpoff, Milford H.; Hawks, John; Caspari, Rachel (May 2000). "Multiregional, Not Multiple
Origins" ([Link] (PDF).
American Journal of Physical Anthropology. 112 (1): 129–136. doi:10.1002/(SICI)1096-
8644(200005)112:1<129::AID-AJPA11>[Link];2-K ([Link]
91096-8644%28200005%29112%3A1%3C129%3A%3AAID-AJPA11%[Link]%3B2-K).
hdl:2027.42/34270 ([Link] ISSN 0002-9483 ([Link]
[Link]/issn/0002-9483). PMID 10766948 ([Link]
48). Archived ([Link]
u/~wolpoff/Papers/[Link]) (PDF) from the original on January 25, 2021.
Retrieved May 12, 2013.
263. Wolpoff, M. H.; Spuhler, J. N.; Smith, F. H.; Radovcic, J.; Pope, G.; Frayer, D. W.; Eckhardt,
R.; Clark, G. (August 12, 1988). "Modern Human Origins". Science. 241 (4867): 772–774.
Bibcode:1988Sci...241..772W ([Link]
doi:10.1126/science.3136545 ([Link] ISSN 0036-
8075 ([Link] PMID 3136545 ([Link]
[Link]/3136545). S2CID 5223638 ([Link]
264. Owen, James (July 18, 2007). "Modern Humans Came Out of Africa, 'Definitive' Study Says"
([Link]
007/07/[Link]). National Geographic News. Washington, DC: National
Geographic Society. Archived from the original ([Link]
07/07/[Link]) on June 27, 2018. Retrieved May 14, 2011.
265. Stringer, Chris B.; Andrews, Peter (March 11, 1988). "Genetic and fossil evidence for the
origin of modern humans". Science. 239 (4845): 1263–1268. Bibcode:1988Sci...239.1263S
([Link] doi:10.1126/science.3125610 (htt
ps://[Link]/10.1126%2Fscience.3125610). ISSN 0036-8075 ([Link]
n/0036-8075). PMID 3125610 ([Link] S2CID 39200956
([Link]
266. Webster 2010, p. 53 ([Link]
267. Ramachandran et al. 2010, p. 606 ([Link]
PA606).
268. Cann, Rebecca L.; Stoneking, Mark; Wilson, Allan C. (January 1, 1987). "Mitochondrial DNA
and human evolution" ([Link]
~landc/html/cann/). Nature. 325 (6099): 31–36. Bibcode:1987Natur.325...31C ([Link]
[Link]/abs/1987Natur.325...31C). doi:10.1038/325031a0 ([Link]
8%2F325031a0). ISSN 0028-0836 ([Link]
PMID 3025745 ([Link] S2CID 4285418 ([Link]
[Link]/CorpusID:4285418). Archived from the original ([Link]
c/html/cann/) on August 13, 2010. Retrieved April 21, 2015.
269. Gill, Victoria (May 1, 2009). "Africa's genetic secrets unlocked" ([Link]
ience/nature/[Link]). BBC News. London: BBC. Archived ([Link]
b/20200514095952/[Link] from the original
on May 14, 2020. Retrieved June 8, 2011.
270. Leakey 1994, pp. 87–89.
271. Jorde, Lynn B.; Bamshad, Michael; Rogers, Alan R. (February 1998). "Using mitochondrial
and nuclear DNA markers to reconstruct human evolution". BioEssays. 20 (2): 126–136.
doi:10.1002/(SICI)1521-1878(199802)20:2<126::AID-BIES5>[Link];2-R ([Link]
1002%2F%28SICI%291521-1878%28199802%2920%3A2%3C126%3A%3AAID-BIES5%3
[Link]%3B2-R). ISSN 0265-9247 ([Link]
PMID 9631658 ([Link] S2CID 17203268 ([Link]
[Link]/CorpusID:17203268).
272. Wall, Jeffrey D.; Lohmueller, Kirk E.; Plagnol, Vincent (August 2009). "Detecting Ancient
Admixture and Estimating Demographic Parameters in Multiple Human Populations" (https://
[Link]/pmc/articles/PMC2734152). Molecular Biology and Evolution. 26 (8):
1823–1827. doi:10.1093/molbev/msp096 ([Link]
ISSN 0737-4038 ([Link] PMC 2734152 ([Link]
[Link]/pmc/articles/PMC2734152). PMID 19420049 ([Link]
ov/19420049).
273. Huertha Sanchez, Emilia et al. (2014), "Altitude adaptation in Tibetans caused by
introgression of Denisovan-like DNA" (Nature Vol 512, August 14, 2014)
274. "Artificial intelligence applied to the genome identifies an unknown human ancestor" (https://
[Link]/releases/2019/01/[Link]). ScienceDaily. Archived (http
s://[Link]/web/20190116213803/[Link]
[Link]) from the original on January 16, 2019. Retrieved January 17, 2019.
275. Lahr, Marta Mirazón; Petraglia, Mike; Stokes, Stephen; et al. "Searching for traces of the
Southern Dispersal" ([Link]
[Link]/Projects/sdispersal/[Link]). Leverhulme Centre for Human Evolutionary
Studies. University of Cambridge. Archived from the original ([Link]
[Link]/Projects/sdispersal/[Link]) on May 10, 2012. Retrieved April 21, 2015.
276. Simpson, Colin. DNA evidence suggests the whole world is a little bit Arab ([Link]
[Link]/uae/science/dna-evidence-suggests-the-whole-world-is-a-little-bit-arab-1.4
08619). The National. 16, June 2020. Archived at The Wayback Machine ([Link]
[Link]/uae/science/dna-evidence-suggests-the-whole-world-is-a-little-bit-arab-1.40
8619)
277. Fernandes, V.; Alshamali, F.; Alves, M.; et al. (2012). "The Arabian cradle: Mitochondrial
relicts of the first steps along the southern route out of Africa" ([Link]
pmc/articles/PMC3276663). American Journal of Human Genetics. 90 (2): 347–355.
Bibcode:2012AmJHG..90..347F ([Link]
doi:10.1016/[Link].2011.12.010 ([Link]
PMC 3276663 ([Link] PMID 22284828
([Link]
278. Macaulay, Vincent; Hill, Catherine; Achilli, Alessandro; et al. (May 13, 2005). "Single, Rapid
Coastal Settlement of Asia Revealed by Analysis of Complete Mitochondrial Genomes" (htt
p://[Link]/id/eprint/40255/1/Single%2C%20rapid%20coastal%20settlement%20
of%20Asia%20revealed%20by%20analysis%20of%20complete%20mitochondrial%20geno
[Link]) (PDF). Science. 308 (5724): 1034–1036. Bibcode:2005Sci...308.1034M ([Link]
[Link]/abs/2005Sci...308.1034M). doi:10.1126/science.1109792 ([Link]
g/10.1126%2Fscience.1109792). ISSN 0036-8075 ([Link]
75). PMID 15890885 ([Link] S2CID 31243109 (http
s://[Link]/CorpusID:31243109). Archived ([Link]
20828110206/[Link]
20settlement%20of%20Asia%20revealed%20by%20analysis%20of%20complete%20mitoc
hondrial%[Link]) (PDF) from the original on August 28, 2022. Retrieved August 20,
2019.
279. Posth, C.; Renaud, G.; Mittnik, M.; Drucker, D. G.; Rougier, H.; Cupillard, C.; et al. (2016).
"Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and
a Late Glacial Population Turnover in Europe" ([Link]
7). Current Biology. 26 (6): 827–833. Bibcode:2016CBio...26..827P ([Link]
[Link]/abs/2016CBio...26..827P). doi:10.1016/[Link].2016.01.037 ([Link]
[Link].2016.01.037). hdl:2440/114930 ([Link]
PMID 26853362 ([Link] S2CID 140098861 ([Link]
[Link]/CorpusID:140098861).
280. Karmin, M.; Saag, L.; Vicente, M.; Wilson Sayres, M. A.; Järve, M.; Talas, U. G.; et al. (April
2015). "A recent bottleneck of Y chromosome diversity coincides with a global change in
culture" ([Link] Genome Research. 25
(4): 459–66. doi:10.1101/gr.186684.114 ([Link]
PMC 4381518 ([Link] PMID 25770088
([Link]
281. Haber, M.; Jones, A. L.; Connell, B. A.; Arciero, E.; Yang, H.; Thomas, M. G.; et al. (August
2019). "A Rare Deep-Rooting D0 African Y-Chromosomal Haplogroup and Its Implications
for the Expansion of Modern Humans Out of Africa" ([Link]
es/PMC6707464). Genetics. 212 (4): 1421–1428. doi:10.1534/genetics.119.302368 (https://
[Link]/10.1534%2Fgenetics.119.302368). PMC 6707464 ([Link]
c/articles/PMC6707464). PMID 31196864 ([Link]
282. Stanford, Allen & Antón 2009.
283. Wood, Bernard A.; Richmond, Brian G. (July 2000). "Human evolution: Taxonomy and
paleobiology" ([Link] Journal of
Anatomy. 197 (1): 19–60. doi:10.1046/j.1469-7580.2000.19710019.x ([Link]
6%2Fj.1469-7580.2000.19710019.x). ISSN 1469-7580 ([Link]
9-7580). PMC 1468107 ([Link]
PMID 10999270 ([Link]
284. Ajit, Varki; Nelson, David L. (October 2007). "Genomic Comparisons of Humans and
Chimpanzees" ([Link] (PDF). Annual
Review of Anthropology. 36: 191–209. doi:10.1146/[Link].36.081406.094339 (http
s://[Link]/10.1146%[Link].36.081406.094339). ISSN 0084-6570 ([Link]
[Link]/issn/0084-6570). Archived ([Link]
p://[Link]/_files/fall2008/[Link]) (PDF) from the original on
September 23, 2015. Retrieved April 26, 2015. "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%."
285. Coghlan, Andy (May 30, 2002). "Just 2.5% of DNA turns mice into men" ([Link]
[Link]/article/dn2352-just-2-5-of-dna-turns-mice-into-men/#:~:text=Mice%20and%20me
n%20share%20about%2097.5%20per,mouse-human%20differences%20as%20high%20a
s%2015%20per.). New Scientist. Retrieved August 18, 2024.
286. Dawkins 2004.
287. "Find Time of Divergence: Hominidae versus Hylobatidae" ([Link]
50418222205/[Link]
&submit=Search). TimeTree. Archived from the original ([Link]
axon_a=Hominidae&taxon_b=Hylobatidae&submit=Search) on April 18, 2015. Retrieved
April 18, 2015.
288. Ruvolo, Maryellen (October 1997). "Genetic Diversity in Hominoid Primates". Annual Review
of Anthropology. 26: 515–540. doi:10.1146/[Link].26.1.515 ([Link]
6%[Link].26.1.515). ISSN 0084-6570 ([Link]
0).
289. Ruvolo, Maryellen (March 1997). "Molecular Phylogeny of the Hominoids: Inferences from
Multiple Independent DNA Sequence Data Sets" ([Link]
molbev.a025761). Molecular Biology and Evolution. 14 (3): 248–265.
doi:10.1093/[Link].a025761 ([Link]
ev.a025761). ISSN 0737-4038 ([Link] PMID 9066793
([Link]
290. Patterson, N.; Richter, D. J.; Gnerre, S.; Lander, E. S.; Reich, D. (2006). "Genetic evidence
for complex speciation of humans and chimpanzees". Nature. 441 (7097): 1103–1108.
Bibcode:2006Natur.441.1103P ([Link]
doi:10.1038/nature04789 ([Link] PMID 16710306 (http
s://[Link]/16710306). S2CID 2325560 ([Link]
orpusID:2325560).
291. Woodley, Michael A.; Younuskunju, Shameem; Balan, Bipin; Piffer, Davide (2017).
"Holocene Selection for Variants Associated With General Cognitive Ability: Comparing
Ancient and Modern Genomes" ([Link]
7417000378/type/journal_article). Twin Research and Human Genetics. 20 (4): 271–280.
doi:10.1017/thg.2017.37 ([Link] ISSN 1832-4274 (https://
[Link]/issn/1832-4274). PMID 28724479 ([Link]
24479). Retrieved April 17, 2026.
292. Zimmer, Carl (May 17, 2023). "Study Offers New Twist in How the First Humans Evolved - A
new genetic analysis of 290 people suggests that humans emerged at various times and
places in Africa" ([Link]
The New York Times. Retrieved May 18, 2023.
293. Ragsdale, Aaron P.; et al. (May 17, 2023). "A weakly structured stem for human origins in
Africa" ([Link] Nature. 167 (7962): 755–
763. Bibcode:2023Natur.617..755R ([Link]
R). doi:10.1038/s41586-023-06055-y ([Link]
PMC 10208968 ([Link]
PMID 37198480 ([Link]
294. Begun, David R. (October 2010). "Miocene Hominids and the Origins of the African Apes
and Humans". Annual Review of Anthropology. 39: 67–84.
doi:10.1146/[Link].012809.105047 ([Link]
809.105047). ISSN 0084-6570 ([Link]
295. Wood 2009, pp. 17–27 ([Link]
#page-1).
296. Mitchell, Alanna (January 30, 2012). "DNA Turning Human Story Into a Tell-All" ([Link]
[Link]/archive/20220101/[Link]
[Link]). The New York Times. Archived from the
original ([Link]
[Link]) on January 1, 2022. Retrieved February 13, 2012.
297. Wood, Bernard A. (December 1996). "Human evolution". BioEssays. 18 (12): 945–954.
doi:10.1002/bies.950181204 ([Link] ISSN 0265-9247
([Link] PMID 8976151 ([Link]
ov/8976151). S2CID 221464189 ([Link]
298. Huxley, T. (1890). "The Aryan Question and Pre-Historic Man" ([Link]
ey/CE7/[Link]). Collected Essays: Volume VII, Man's Place in Nature. Archived (https://
[Link]/web/20110726231311/[Link] from
the original on July 26, 2011. Retrieved August 25, 2018.
299. Coghlan, Andy (March 15, 2018). "Our ancestors mated with the mystery 'Denisovan' people
– twice" ([Link]
ry-denisovan-people-twice/). New Scientist. Archived ([Link]
035916/[Link]
y-denisovan-people-twice) from the original on August 5, 2018. Retrieved August 25, 2018.
300. Wei-Haas, Maya (August 22, 2018). "Ancient Girl's Parents Were Two Different Human
Species – Born 90,000 years ago, the child is the first direct evidence of interbreeding
among Neanderthals and their cousins the Denisovans" ([Link]
614092015/[Link]
hal-hominin-hybrid-ancient-human/). National Geographic. Archived from the original (http
s://[Link]/science/2018/08/news-denisovan-neanderthal-hominin-hyb
rid-ancient-human/) on June 14, 2019. Retrieved August 22, 2018.
301. Cell Press (March 28, 2016). "A world map of Neanderthal and Denisovan ancestry in
modern humans" ([Link]
ScienceDaily. Archived ([Link]
[Link]/releases/2016/03/[Link]) from the original on August 26, 2018.
Retrieved August 25, 2018.
302. Sankararaman, S.; Mallick, S.; Patterson, N.; Reich, D. (March 28, 2016). "The Combined
Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans" ([Link]
[Link]/pmc/articles/PMC4864120). Current Biology. 26 (9): 1241–1247.
Bibcode:2016CBio...26.1241S ([Link]
doi:10.1016/[Link].2016.03.037 ([Link]
PMC 4864120 ([Link] PMID 27032491
([Link]
303. "Human-Neandertal Comparisons" ([Link]
[Link]/site/special/neandertal/feature/[Link]). the Neandertal Genome.
American Association for the Advancement of Science. 2010. Archived from the original (htt
p://[Link]/site/special/neandertal/feature/[Link]) on May 28, 2020.
304. Dannemann, M.; Andrés, A. M.; Kelso, J. (2016). "Introgression of Neandertal- and
Denisovan-like Haplotypes Contributes to Adaptive Variation in Human Toll-like Receptors"
([Link] American Journal of Human
Genetics. 98 (1): 22–33. Bibcode:2016AmJHG..98...22D ([Link]
2016AmJHG..98...22D). doi:10.1016/[Link].2015.11.015 ([Link]
015.11.015). PMC 4716682 ([Link]
PMID 26748514 ([Link]
305. Gittelman, Rachel M.; Schraiber, Joshua G.; Vernot, Benjamin; Mikacenic, Carmen; Wurfel,
Mark M.; Akey, Joshua M. (2016). "Archaic Hominin Admixture Facilitated Adaptation to Out-
of-Africa Environments" ([Link] Current
Biology. 26 (24): 3375–3382. Bibcode:2016CBio...26.3375G ([Link]
abs/2016CBio...26.3375G). doi:10.1016/[Link].2016.10.041 ([Link]
b.2016.10.041). PMC 6764441 ([Link]
PMID 27839976 ([Link]
306. Plummer, Thomas (2004). "Flaked stones and old bones: Biological and cultural evolution at
the dawn of technology" ([Link] American Journal of
Physical Anthropology. 125 (Supplement 39: Yearbook of Physical Anthropology): 118–164.
Bibcode:2004AJPA..125S.118P ([Link]
doi:10.1002/ajpa.20157 ([Link] ISSN 0002-9483 ([Link]
[Link]/issn/0002-9483). PMID 15605391 ([Link]
5391).
307. Wong, Kate (April 15, 2015). "Archaeologists Take Wrong Turn, Find World's Oldest Stone
Tools" ([Link]
ng-turn-find-worlds-oldest-stone-tools/). Scientific American (Blog). ISSN 0036-8733 (https://
[Link]/issn/0036-8733). Archived ([Link]
44/[Link]
urn-find-worlds-oldest-stone-tools/) from the original on May 8, 2015. Retrieved May 3,
2015.
308. Balter, Michael (April 14, 2015). "World's oldest stone tools discovered in Kenya" ([Link]
[Link]/content/article/world-s-oldest-stone-tools-discovered-kenya). Science (News).
doi:10.1126/science.aab2487 ([Link] ISSN 0036-
8075 ([Link] Archived ([Link]
211020055503/[Link]
-kenya) from the original on October 20, 2021. Retrieved May 3, 2015.
309. Drake, Nadia (April 16, 2015). "Oldest Stone Tools Discovered in Kenya" ([Link]
[Link]/web/20150423222706/[Link]
arts-spill-tuna-gulf-bp-deepwater-exxon-alaska/150416-oldest-stone-tools-archaeology-keny
a-human-origins-evolution/). National Geographic News. Washington, DC: National
Geographic Society. Archived from the original ([Link]
4/150416-oil-fish-hearts-spill-tuna-gulf-bp-deepwater-exxon-alaska/150416-oldest-stone-tool
s-archaeology-kenya-human-origins-evolution/) on April 23, 2015. Retrieved May 3, 2015.
310. Yellen, J. E.; Brooks, A. S.; Cornelissen, E.; Mehlman, M. J.; Stewart, K. (April 28, 1995). "A
middle stone age worked bone industry from Katanda, Upper Semliki Valley, Zaire".
Science. 268 (5210): 553–556. Bibcode:1995Sci...268..553Y ([Link]
abs/1995Sci...268..553Y). doi:10.1126/science.7725100 ([Link]
e.7725100). PMID 7725100 ([Link]

Sources
Aiello, Leslie; Dean, Christopher (1990). An Introduction to Human Evolutionary Anatomy.
London / San Diego: Elsevier Academic Press. ISBN 978-0-12-045591-1. LCCN 95185095
([Link] OCLC 33408268 ([Link]
8).
Bogin, Barry (1997). "The Evolution of Human Nutrition". In Romanucci-Ross, Lola;
Moerman, Daniel E.; Tancredi, Laurence R. (eds.). The Anthropology of Medicine: From
Culture to Method (3rd ed.). Westport, Connecticut: Bergin & Garvey. ISBN 978-0-89789-
516-3. LCCN 96053993 ([Link] OCLC 36165190 ([Link]
[Link]/oclc/36165190).
Bown, Thomas M.; Rose, Kenneth D. (1987). Patterns of Dental Evolution in Early Eocene
Anaptomorphine Primates (Omomyidae) From the Bighorn Basin, Wyoming ([Link]
[Link]/cgi/[Link]?article=1217&context=usgsstaffpub) (PDF). Memoir
(Paleontological Society). Vol. 23. Tulsa, Oklahoma: Paleontological Society.
OCLC 16997265 ([Link] Archived ([Link]
org/web/20161111062501/[Link]
ontext=usgsstaffpub) from the original on November 11, 2016. Retrieved April 25, 2015.
Boyd, Robert; Silk, Joan B. (2003). How Humans Evolved ([Link]
mansevolved03edboyd) (3rd ed.). New York: W. W. Norton & Co. ISBN 978-0-393-97854-4.
LCCN 2002075336 ([Link] OCLC 49959461 ([Link]
[Link]/oclc/49959461).
Boyd, Robert; Silk, Joan B. (2021). How Humans Evolved (9th ed.). New York: W. W. Norton
& Co. ISBN 978-0-393-53315-6. OCLC 1282924892 ([Link]
24892).
Brues, Alice M.; Snow, Clyde C. (1965). "Physical Anthropology". In Siegel, Bernard J. (ed.).
Biennial Review of Anthropology 1965. Biennial Review of Anthropology. Vol. 4. Stanford
University Press. ISBN 978-0-8047-1746-5. ISSN 0067-8503 ([Link]
n/0067-8503). LCCN 59012726 ([Link] OCLC 01532912 ([Link]
[Link]/oclc/01532912).
Bryson, Bill (2004) [Originally published 2003]. "The Mysterious Biped". A Short History of
Nearly Everything. Toronto: Anchor Canada. ISBN 978-0-385-66004-4. LCCN 2003046006
([Link] OCLC 55016591 ([Link]
91).
Cameron, David W. (2004). Hominid Adaptations and Extinctions. Sydney: University of
New South Wales Press. ISBN 978-0-86840-716-6. LCCN 2004353026 ([Link]
2004353026). OCLC 57077633 ([Link]
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 ([Link] OCLC 191926088 ([Link]
[Link]/oclc/191926088).
Curry, James R. (2008). Children of God: Children of Earth. Bloomington, Indiana:
AuthorHouse. ISBN 978-1-4389-1846-4. OCLC 421466369 ([Link]
c/421466369).
Darwin, Charles (1859). On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life ([Link]
ages/images/nytint/docs/charles-darwin-on-the-origin-of-species/[Link]) (PDF).
Archived ([Link]
ges/images/nytint/docs/charles-darwin-on-the-origin-of-species/[Link]) (PDF) from the
original on March 4, 2016. Retrieved May 13, 2017.
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 University Press. ISBN 978-0-691-02369-4. LCCN 80008679
([Link] OCLC 7197127 ([Link]
Dawkins, Richard (2004). The Ancestor's Tale: A Pilgrimage to the Dawn of Evolution.
Boston: Houghton Mifflin. ISBN 978-0-618-00583-3. LCCN 2004059864 ([Link]
v/2004059864). OCLC 56617123 ([Link]
DeSalle, Rob; Tattersall, Ian (2008). Human Origins: What Bones and Genomes Tell Us
About Ourselves ([Link] Texas A&M
University Anthropology Series. Vol. 13. College Station: Texas A&M University Press.
ISBN 978-1-58544-567-7. OCLC 144520427 ([Link]
Diamond, Jared (1999). Guns, Germs, and Steel: The Fates of Human Societies. New York:
W. W. Norton & Co. ISBN 978-0-393-31755-8. LCCN 2005284124 ([Link]
284124). OCLC 35792200 ([Link]
Freeman, Scott; Herron, Jon C. (2007). Evolutionary Analysis (4th ed.). Upper Saddle River,
New Jersey: Pearson Prentice Hall. ISBN 978-0-13-227584-2. LCCN 2006034384 ([Link]
[Link]/2006034384). OCLC 73502978 ([Link]
Johanson, Donald; Edey, Maitland (1981). Lucy, the Beginnings of Humankind. St Albans:
Granada. ISBN 978-0-586-08437-3.
Montgomery, William M. (1988) [Originally published 1974]. "Germany". In Glick, Thomas F.
(ed.). The Comparative Reception of Darwinisms. University of Chicago Press. ISBN 978-0-
226-29977-8. LCCN 87035814 ([Link] OCLC 17328115 ([Link]
[Link]/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". American Journal of Physical Anthropology. 70 (2). University of Tokyo Press:
278–279. doi:10.1002/ajpa.1330700214 ([Link]
ISBN 978-4-13-066093-8. LCCN 85173489 ([Link]
OCLC 12352830 ([Link]
Leakey, Richard E. (1994). The Origin of Humankind ([Link]
ankin00leak). Science Masters Series. New York: Basic Books. ISBN 978-0-465-03135-1.
LCCN 94003617 ([Link] OCLC 30739453 ([Link]
org/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
University Press. ISBN 978-0-521-83222-9. LCCN 2004052648 ([Link]
2648). OCLC 55600894 ([Link]
Martin, Robert D. (2001). "Primates, Evolution of". In Smelser, Neil J.; Baltes, Paul B. (eds.).
International Encyclopedia of the Social & Behavioral Sciences. Amsterdam / New York:
Elsevier. pp. 12032–12038. doi:10.1016/B0-08-043076-7/03083-7 ([Link]
2FB0-08-043076-7%2F03083-7). ISBN 978-0-08-043076-8. LCCN 2001044791 ([Link]
[Link]/2001044791). OCLC 47869490 ([Link]
Maxwell, Mary (1984). Human Evolution: A Philosophical Anthropology. New York: Columbia
University Press. ISBN 978-0-231-05946-6. LCCN 83024005 ([Link]
5). OCLC 10163036 ([Link]
McHenry, Henry M. (2009). "Human Evolution" ([Link]
mich). In Ruse, Michael; Travis, Joseph (eds.). Evolution: The First Four Billion Years.
Foreword by Edward O. Wilson. Cambridge, Massachusetts: Belknap Press of Harvard
University. ISBN 978-0-674-03175-3. LCCN 2008030270 ([Link]
OCLC 225874308 ([Link]
Ramachandran, Sohini; Tang, Hua; Gutenkunst, Ryan N.; Bustamante, Carlos D. (2010).
"Genetics and Genomics of Human Population Structure". In Speicher, Michael R.;
Antonarakis, Stylianos E.; Motulsky, Arno G. (eds.). Vogel and Motulsky's Human Genetics:
Problems and Approaches (4th ed.). Heidelberg / London / New York: Springer.
doi:10.1007/978-3-540-37654-5 ([Link] ISBN 978-
3-540-37653-8. LCCN 2009931325 ([Link] OCLC 549541244 (ht
tps://[Link]/oclc/549541244).
Robinson, J. T. (2008) [Originally published 1963; Chicago, Illinois: Aldine Transaction].
"Adaptive Radiation in the Australopithecines and the Origin of Man". In Howell, F. Clark;
Bourlière, François (eds.). African Ecology and Human Evolution. New Brunswick, New
Jersey: Transaction Publishers. ISBN 978-0-202-36136-9. LCCN 2007024716 ([Link]
[Link]/2007024716). OCLC 144770218 ([Link]
Srivastava, R. P. (2009). Morphology of the Primates and Human Evolution. New Delhi: PHI
Learning. ISBN 978-81-203-3656-8. OCLC 423293609 ([Link]
293609).
Stanford, Craig; Allen, John S.; Antón, Susan C. (2009). Biological Anthropology: The
Natural History of Humankind ([Link] (2nd ed.).
Upper Saddle River, New Jersey: Pearson Prentice Hall. ISBN 978-0-13-601160-6.
LCCN 2007052429 ([Link] OCLC 187548835 ([Link]
[Link]/oclc/187548835).
Strickberger, Monroe W. (2000). Evolution (3rd ed.). Sudbury, Massachusetts: Jones &
Bartlett Publishers. ISBN 978-0-7637-1066-8. LCCN 99032072 ([Link]
72). OCLC 41431683 ([Link]
Stringer, Chris B. (1994) [1992]. "Evolution of Early Humans". In Jones, Steve; Martin,
Robert D.; Pilbeam, David (eds.). The Cambridge Encyclopedia of Human Evolution (https://
[Link]/details/cambridgeencyclo00step). Cambridge University Press. ISBN 978-0-521-
32370-3. LCCN 92018037 ([Link] OCLC 444512451 ([Link]
[Link]/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.
University of Chicago Press. ISBN 978-0-226-78734-3. LCCN 2001037337 ([Link]
gov/2001037337). OCLC 48066180 ([Link]
Trent, Ronald J. (2005). Molecular Medicine: An Introductory Text (3rd ed.). Burlington,
Massachusetts: Elsevier Academic Press. ISBN 978-0-12-699057-7. LCCN 2004028087 (htt
ps://[Link]/2004028087). OCLC 162577235 ([Link]
35).
 Trevathan, Wenda R. (2011) [Originally published 1987; New York: Aldine De Gruyter].
Human Birth: An Evolutionary Perspective. New Brunswick, New Jersey: Transaction
Publishers. ISBN 978-1-4128-1502-4. LCCN 2010038249 ([Link]
OCLC 669122326 ([Link]
Ungar, Peter S.; Teaford, Mark F., eds. (2002). Human Diet: Its Origin and Evolution.
Westport, Connecticut: Bergin & Garvey. ISBN 978-0-89789-736-5. LCCN 2001043790 (http
s://[Link]/2001043790). OCLC 537239907 ([Link]
7).
Walker, Alan (2007). "Early Hominin Diets: Overview and Historical Perspectives". In Ungar,
Peter (ed.). Evolution of the Human Diet: The Known, the Unknown, and the Unknowable.
Human Evolution Series. Oxford University Press. ISBN 978-0-19-518346-7.
LCCN 2005036120 ([Link] OCLC 132816551 ([Link]
[Link]/oclc/132816551).
Wallace, David Rains (2004). Beasts of Eden: Walking Whales, Dawn Horses, and Other
Enigmas of Mammal Evolution. Berkeley: University of California Press. ISBN 978-0-520-
24684-3. LCCN 2003022857 ([Link] OCLC 53254011 ([Link]
[Link]/oclc/53254011).
Webster, Donovan (2010). Meeting the Family: One Man's Journey Through His Human
Ancestry. Foreword by Spencer Wells. Washington, DC: National Geographic Society.
ISBN 978-1-4262-0573-6. LCCN 2009050471 ([Link]
OCLC 429022321 ([Link]
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. (eds.). The First
Humans – Origin and Early Evolution of the Genus Homo ([Link]
mansorigi00grin). Vertebrate Paleobiology and Paleoanthropology. Dordrecht: Springer
Netherlands. pp. 17 ([Link]
doi:10.1007/978-1-4020-9980-9_3 ([Link]
ISBN 978-1-4020-9979-3. ISSN 1877-9077 ([Link]
LCCN 2009927083 ([Link] OCLC 310400980 ([Link]
[Link]/oclc/310400980). "Contributions from the Third Stony Brook Human Evolution
Symposium and Workshop October 3–7, 2006."

Further reading
Alexander, Richard D. (1990). How Did Humans Evolve? Reflections on the Uniquely
Unique Species ([Link]
y/LahtiSites/RDAlexander/Pubs/[Link]) (PDF). Special Publication. Ann Arbor:
Museum of Zoology, University of Michigan. pp. 1–38. LCCN 90623893 ([Link]
90623893). OCLC 22860997 ([Link] Archived from the
original ([Link]
(PDF) on March 4, 2016. Retrieved May 6, 2015.
Barton, Nicholas H.; Briggs, Derek E. G.; Eisen, Jonathan A.; et al. (2007). Evolution. Cold
Spring Harbor Laboratory Press. ISBN 978-0-87969-684-9. LCCN 2007010767 ([Link]
[Link]/2007010767). OCLC 86090399 ([Link]
Bellwood, Peter (2022). The Five-Million-Year Odyssey: The Human Journey from Ape to
Agriculture. Princeton: Princeton University Press. ISBN 978-0-691-19757-9.
Enard, Wolfgang; Przeworski, Molly; Fisher, Simon E.; et al. (August 22, 2002). "Molecular
evolution of FOXP2, a gene involved in speech and language". Nature. 418 (6900): 869–
872. Bibcode:2002Natur.418..869E ([Link]
E). doi:10.1038/nature01025 ([Link] hdl:11858/00-001M-
0000-0012-CB89-A ([Link]
ISSN 0028-0836 ([Link] PMID 12192408 ([Link]
[Link]/12192408). S2CID 4416233 ([Link]
D:4416233).
Flinn, Mark V.; Geary, David C.; Ward, Carol V. (2005). "Ecological dominance, social
competition, and coalitionary arms races: Why humans evolved extraordinary intelligence"
([Link] (PDF). Evolution and Human Behavior.
26 (1): 10–46. Bibcode:2005EHumB..26...10F ([Link]
mB..26...10F). doi:10.1016/[Link].2004.08.005 ([Link]
umbehav.2004.08.005). ISSN 1090-5138 ([Link]
Archived ([Link]
[Link]) (PDF) from the original on June 4, 2007. Retrieved May 5, 2015.
Galinon-Melenec, Béatrice (2015). "From "Traces" and "Human Trace" to "Human–Trace
Paradigm" ". In Parrend, Pierre; Bourgine, Paul; Collet, Pierre (eds.). First Complex systems
Digital Campus World E-Conference. Tempe, Arizona: Springer.
Gibbons, Ann (2006). The First Human: The Race to Discover our Earliest Ancestors (http
s://[Link]/details/firsthumanraceto00gibb). New York: Doubleday. ISBN 978-0-385-
51226-8. LCCN 2005053780 ([Link] OCLC 61652817 ([Link]
[Link]/oclc/61652817).
Hartwig, Walter C., ed. (2002). The Primate Fossil Record. Cambridge Studies in Biological
and Evolutionary Anthropology. Vol. 33. Cambridge University Press. ISBN 978-0-521-
66315-1. LCCN 2001037847 ([Link] OCLC 47254191 ([Link]
[Link]/oclc/47254191).
Heizmann, Elmar P. J.; Begun, David R. (November 2001). "The oldest Eurasian hominoid"
([Link] (PDF). Journal of Human Evolution. 41
(5): 463–481. Bibcode:2001JHumE..41..463H ([Link]
E..41..463H). doi:10.1006/jhev.2001.0495 ([Link]
ISSN 0047-2484 ([Link] PMID 11681862 ([Link]
[Link]/11681862). S2CID 21230992 ([Link]
D:21230992).
Hill, Andrew; Ward, Steven (1988). "Origin of the hominidae: The record of African large
hominoid evolution between 14 my and 4 my" ([Link]
5). American Journal of Physical Anthropology. 31 (59): 49–83.
Bibcode:1988AJPA...31S..49H ([Link]
doi:10.1002/ajpa.1330310505 ([Link] ISSN 0002-
9483 ([Link]
Hoagland, Hudson (1964). "Science and the New Humanism". Science. 143 (3602): 111–
114. Bibcode:1964Sci...143..111H ([Link]
doi:10.1126/science.143.3602.111 ([Link]
PMID 17781189 ([Link]
Ijdo, Jacob W.; Baldini, Antonio; Ward, David C.; et al. (October 15, 1991). "Origin of human
chromosome 2: An ancestral telomere-telomere fusion" ([Link]
rticles/PMC52649). Proceedings of the National Academy of Sciences of the United States
of America. 88 (20): 9051–9055. Bibcode:1991PNAS...88.9051I ([Link]
du/abs/1991PNAS...88.9051I). doi:10.1073/pnas.88.20.9051 ([Link]
as.88.20.9051). ISSN 0027-8424 ([Link] PMC 52649
([Link] PMID 1924367 ([Link]
[Link]/1924367). – 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 ([Link]
v/2010483830). OCLC 419801728 ([Link]
Jones, Steve; Martin, Robert D.; Pilbeam, David, eds. (1994) [1992]. The Cambridge
Encyclopedia of Human Evolution ([Link]
Foreword by Richard Dawkins. Cambridge University Press. ISBN 978-0-521-32370-3.
LCCN 92018037 ([Link] OCLC 444512451 ([Link]
[Link]/oclc/444512451). (This book contains very useful, information-dense chapters on
primate evolution in general, and human evolution in particular, including fossil history.)
Kawabata, Hiroshi; Kaifu, Yousuke (2020). Lost in Evolution: Exploring Humanity's Path in
Asia ([Link]
[Link]). Tokyo: Japan Publishing Industry Foundation for Culture. ISBN 978-4-
86658-133-0. Archived ([Link]
[Link]/books/science/[Link]) from the
original on October 20, 2022. Retrieved October 20, 2022.
Leakey, Richard E.; Lewin, Roger (1992). Origins Reconsidered: In Search of What Makes
us Human. New York: Doubleday. ISBN 978-0-385-41264-3. LCCN 92006661 ([Link]
[Link]/92006661). OCLC 25373161 ([Link]
Lewin, Roger (1997). Bones of Contention: Controversies in the Search for Human Origins
(2nd ed.). University of Chicago Press. ISBN 978-0-226-47651-3. LCCN 97000972 ([Link]
[Link]/97000972). OCLC 36181117 ([Link]
Morwood, Mike; van Oosterzee, Penny (2007). A New Human: The Startling Discovery and
Strange Story of the 'Hobbits' of Flores, Indonesia ([Link]
nge00morw). New York: Smithsonian Books / HarperCollins. ISBN 978-0-06-089908-0.
LCCN 2006052267 ([Link] OCLC 76481584 ([Link]
[Link]/oclc/76481584).
Oppenheimer, Stephen (2003). Out of Eden: The Peopling of the World. London: Constable
& Robinson. ISBN 978-1-84119-697-8. LCCN 2005482222 ([Link]
2). OCLC 52195607 ([Link]
Ovchinnikov, Igor V.; Götherström, Anders; Romanova, Galina P.; et al. (March 30, 2000).
"Molecular analysis of Neanderthal DNA from the Northern Caucasus". Nature. 404 (6777):
490–493. Bibcode:2000Natur.404..490O ([Link]
490O). doi:10.1038/35006625 ([Link] ISSN 0028-0836 (htt
ps://[Link]/issn/0028-0836). PMID 10761915 ([Link]
v/10761915). S2CID 3101375 ([Link]
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 (h
ttps://[Link]/oclc/310156315).
Shreeve, James (1995). The Neandertal Enigma: Solving the Mystery of Modern Human
Origins ([Link] New York: William Morrow &
Co. ISBN 978-0-688-09407-2. LCCN 95006337 ([Link]
OCLC 32088673 ([Link]
Stringer, Chris B. (2011). The Origin of Our Species. London: Allen Lane. ISBN 978-1-
84614-140-9. LCCN 2011489742 ([Link] OCLC 689522193 (http
s://[Link]/oclc/689522193).
Stringer, Chris B.; Andrews, Peter (2005). The Complete World of Human Evolution (https://
[Link]/details/completeworldofh0000stri). London / New York: Thames & Hudson.
ISBN 978-0-500-05132-0. LCCN 2004110563 ([Link]
OCLC 224377190 ([Link]
Stringer, Christopher; McKie, Robin (1997). African Exodus: The Origins of Modern
Humanity ([Link] New York: Henry Holt & Co.
ISBN 978-0-8050-2759-4. LCCN 96037718 ([Link]
OCLC 36001167 ([Link]
Sykes, Rebecca Wragg (2020). Kindred: Neanderthal Life, Love, Death and Art. London:
Bloomsbury Sigma. ISBN 978-1-4729-3749-0.
Tattersall, Ian (2008). The Fossil Trail: How We Know What We Think We Know About
Human Evolution (2nd ed.). Oxford University Press. ISBN 978-0-19-536766-9.
 LCCN 2008013654 ([Link] OCLC 218188644 ([Link]
[Link]/oclc/218188644).
van Oosterzee, Penny (1999). Dragon Bones: The Story of Peking Man. St Leonards, New
South Wales: Allen & Unwin Australia. ISBN 978-1-86508-123-6. LCCN 00300421 ([Link]
[Link]/00300421). OCLC 45853997 ([Link]
Wade, Nicholas (2006). Before the Dawn: Recovering the Lost History of Our Ancestors.
New York: Penguin. ISBN 978-1-59420-079-3. LCCN 2005055293 ([Link]
055293). OCLC 62282400 ([Link]
Walker, Alan; Shipman, Pat (1996). The Wisdom of the Bones: In Search of Human Origins.
London: Weidenfeld & Nicolson. ISBN 978-0-297-81670-6. OCLC 35202130 ([Link]
[Link]/oclc/35202130).
Weiss, Mark L.; Mann, Alan E. (1985). Human Biology and Behavior: An Anthropological
Perspective (4th ed.). Boston: Little, Brown & Co. ISBN 978-0-316-92894-6.
LCCN 85000158 ([Link] OCLC 11726796 ([Link]
rg/oclc/11726796). (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. ISBN 978-0-8129-7146-0. LCCN 2003066679 ([Link]
ov/2003066679). OCLC 53287806 ([Link]

External links
"Race, Evolution and the Science of Human Origins ([Link]
cle/race-evolution-and-the-science-of-human-origins/)" by Allison Hopper, Scientific
American (July 5, 2021).
"The evolution of man" ([Link]
[Link]). BBC Science & Nature. Retrieved May 6, 2015.
"Becoming Human" ([Link] Arizona State University's Institute of
Human Origins. Retrieved May 6, 2015.
"Bones, Stones and Genes: The Origin of Modern Humans" ([Link]
150424155530/[Link]
umans-0) (Video lecture series). Howard Hughes Medical Institute. Archived from the
original ([Link]
-0) on April 24, 2015. Retrieved May 6, 2015.
"Evolution Figures: Chapter 25" ([Link]
[Link]). Cold Spring Harbor Laboratory Press. Retrieved May 6, 2015. – Illustrations from
the book Evolution (2007)
"Human Evolution" ([Link] Smithsonian Institution's Human Origins
Program. Retrieved June 24, 2013.
"Human Evolution Timeline" ([Link]
[Link]/human-evolution-timeline/). [Link]. Archived from the original (htt
p://[Link]/human-evolution-timeline/) on June 18, 2013. Retrieved June 24,
2013.
"Human Trace" video ([Link]
le_de_l_unitwin_unesco_sur_les_systemes_complexes_le_paradigme_de_l_homme_trace.
20031) (2015) Normandy University UNIHAVRE, CNRS, IDEES, [Link] on Human
Trace Unitwin Complex System Digital Campus UNESCO.
Lambert, Tim (Producer) (June 24, 2015). First Peoples ([Link]
ples/). London: Wall to Wall Television. OCLC 910115743 ([Link]
10115743). Retrieved July 18, 2015.
 Shaping Humanity Video ([Link]
2013 Yale University
Human Timeline (Interactive) ([Link]
-interactive) – Smithsonian, National Museum of Natural History (August 2016).
Human Evolution ([Link] BBC Radio 4 discussion
with Steve Jones, Fred Spoor & Margaret Clegg (In Our Time, February 16, 2006)
Evolutionary Timeline of Home Sapiens ([Link]
ssential-timeline-understanding-evolution-homo-sapiens-180976807/) − Smithsonian
(February 2021)
History of Human Evolution in the United States ([Link]
mericans-believe-in-evolution/) – Salon (August 24, 2021)

Retrieved from "[Link]