from chimpanzees to rank as a separate family.
the original restrictive sense; the scholarly
A hominid is any member of the biological
family Hominidae. These are the "great apes",
living and extinct. At present there
are humans, chimpanzees, gorillas,
and orangutans.
The word "hominid" has been used in various
ways. The classification of the great apes has
been revisedseveral times in the last few
decades. These revisions led to different uses
of the word "hominid". The original meaning of
the taxon Hominidae meant only humans and
their closest relatives. Now this group is called
the Hominina.
The primatological term hominid is easily
confused with a number of very similar words:
 A hominoid or ape is a member of the
superfamily Hominoidea: existing members
are the lesser apes(gibbons) and great
apes.
 A hominid is a member of the
family Hominidae: all of the great apes.
 A hominine is a member of the
subfamily Homininae: gorillas,
chimpanzees, humans
(excludes orangutans).
 A hominin is a member of the
tribe Hominini: including humans.
 A hominan is a little used term for a sub-
tribe. It is only used by people who put
chimapnzees in the Hominini.
 A human is a member of Homo sapiens.
The word is sometimes also used to refer
to any extinct member of the
genus Homo or members from other
hominan genera.
 A humanoid is a vaguely human-shaped
entity; the term is typically used in science
fiction.
The dispute is really about whether humans
and Australopithecines are sufficiently different
In morphology and life style they are,
in genetics they are not.
A move
from rainforest to savannah and woodland led
first to bipedalism (walking on two feet). Later,
hunting and social needs led to larger
brains brains and the making and use of tools.
These differences between human beings and
the other great apes are certainly important.
However, a genetically based taxonomic
classificationshows the genetic differences are
not enough to divide us into separate
families. Genetics, rather than morphology, is
generally more widely accepted as the critical
standard. However, many scientists,
including anthropologists, use the term
"hominid" to mean humans and their direct and
near-direct ancestors.
The Hominidae (/hɒˈmɪnɪdiː/), whose members
are known as great apes[note 1] or hominids, are
a taxonomic family of primates that includes
eight extant species in four genera: Pongo,
the Bornean,Sumatran and Tapanuli
orangutan; Gorilla, the eastern and western
gorilla; Pan, the common chimpanzee and
the bonobo; and Homo, of whom only modern
humans remain, with several extinct relatives
(e.g., the Neanderthal) and ancestors, such
as Homo erectus.[1]
Several revisions in classifying the great apes
have caused the use of the term "hominid" to
vary over time. Its original meaning referred
only to humans (Homo) and their closest extinct
relatives. That restrictive meaning has now
been largely assumed by the term "hominin",
which comprises all members of the human
clade after the split from the chimpanzees
(Pan). The current, 21st-century meaning of
"hominid" includes all the great apes including
humans. Usage still varies, however, and some
scientists and laypersons still use "hominid" in
literature generally shows the traditional usage
until around the turn of the 21st century.[4]
Within the taxon Hominidae, a number of extant
and known extinct, that is, fossil, genera are
grouped with the humans, chimpanzees, and
gorillas in the subfamily Homininae; others with
orangutans in the
subfamily Ponginae (see classification
graphic below). The most recent common
ancestor of all Hominidae lived roughly 14
million years ago,[5]when the ancestors of the
orangutans speciated from the ancestral line of
the other three genera.[6] Those ancestors of
the family Hominidae had already speciated
from the family Hylobatidae (the gibbons),
perhaps 15 million to 20 million years ago.[6][7]
In the early Miocene, about 22 million years
ago, there were many species of arboreally
adapted primitive catarrhinesfrom East Africa;
the variety suggests 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, Dendropithec
us, Limnopithecus, Nacholapithecus, Equatoriu
s, Nyanzapithecus, Afropithecus, Heliopithecus,
and Kenyapithecus, all from East Africa.
At sites far distant from East Africa, the
presence of other generalized non-
cercopithecids, that is, non-monkey primates,
of middle Miocene age—Otavipithecus from
cave deposits in Namibia,
and Pierolapithecus and Dryopithecus from
France, Spain and Austria—is further evidence
of a wide diversity of ancestral ape forms
across Africa and the Mediterranean basin
during the relatively warm and equable climatic
regimes of the early and middle Miocene. The
most recent of these far-flung Miocene apes
(hominoids) is Oreopithecus, from the fossil-
rich coal beds in northern Italy and dated to 9
million years ago.
Molecular evidence indicates that the lineage
of gibbons (family Hylobatidae), the lesser
apes, diverged from that of the great apes
some 18–12 million years ago, and that
of orangutans (subfamily Ponginae) diverged
from the other great apes at about 12 million
years. There are no fossils that clearly
document the ancestry of gibbons, which may
have originated in a still-unknown South East
Asian hominoid population; but fossil proto-
orangutans, dated to around 10 million years
ago, may be represented by Sivapithecus from
India and Griphopithecus from Turkey.[8]
Species close to the last common ancestor of
gorillas, chimpanzees and humans may be
represented by Nakalipithecus fossils found in
Kenya and Ouranopithecusfound in Greece.
Molecular evidence suggests that between 8
and 4 million years ago, first the gorillas
(genus Gorilla), 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).[9] 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 most to
this problem.
Other hominins probably adapted to the drier
environments outside the African equatorial
belt; and there they encountered antelope,
hyenas, elephants and other forms becoming
adapted to surviving in the East
African savannas, particularly the regions of
the Sahel and the Serengeti. The wet
equatorial belt contracted after about 8 million
years ago, and there is very little fossil
evidence for the divergence of the hominin
lineage from that of gorillas and chimpanzees—
which split was thought to have occurred
around that time. 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.
 A hominoid, commonly called an ape, is a
member of the superfamily Hominoidea:
extant members are the gibbons (lesser
apes, family Hylobatidae) and the
hominids.
 A hominid is a member of the family
Hominidae, the great
apes: orangutans, gorillas, chimpanzees,
and humans.
 A hominine is a member of the
subfamily Homininae: gorillas,
chimpanzees, and humans
(excludes orangutans).
 A hominin is a member of the
tribe Hominini: chimpanzees and
humans..[10]
 A homininan, following a suggestion by
Wood and Richmond (2000), would be a
member of the subtribe Hominina of the
tribe Hominini: that is, modern humans and
their closest relatives,
including Australopithecina, but excluding
chimpanzees.[11]
 A human is a member of the genus Homo,
of which Homo sapiens is the only extant
species, and within that Homo sapiens
sapiens is the only surviving subspecies.
 The great apes are large, tailless
primates, with the smallest living
species being the bonobo at 30–40
kilograms in weight, and the largest
being the eastern gorillas, with males
weighing 140–180 kilograms. In all
great apes, the males are, on average,
larger and stronger than the females,
although the degree of sexual
dimorphismvaries greatly among
species. Although most living species
are predominantly quadrupedal, they
are all able to use their hands for
gathering food or nesting materials,
and, in some cases, for tool use.[32]
 Most species are omnivorous,[citation
needed]
but fruit is the preferred food
among all but some human groups.
Chimpanzees and orangutans primarily
eat fruit. When gorillas run short of fruit
at certain times of the year or in certain
regions, they resort to eating shoots
and leaves, often of bamboo, a type of
grass. Gorillas have extreme
adaptations for chewing and digesting
such low-quality forage, but they still
prefer fruit when it is available, often
going miles out of their way to find
especially preferred fruits. Humans,
since the neolithic revolution, consume
mostly cereals and other starchy foods,
including increasingly highly processed
foods, as well as many
other domesticated plants (including
fruits) and meat. Hominid teeth are
similar to those of the Old World
monkeys and gibbons, although they
are especially large in gorillas.
The dental formula is 2.1.2.32.1.2.3 .
Human teeth and jaws are markedly
smaller for their size than those of other
apes, which may be an adaptation to
eating cooked food since the end of
the Pleistocene.[33][34]
 Gorilla
 Gestation in great apes lasts 8–9
months, and results in the birth of a
single offspring, or, rarely, twins. The
young are born helpless, and require
care for long periods of time. Compared
 with most other mammals, great apes
have a remarkably long adolescence,
not being weaned for several years,
and not becoming fully mature for eight
to thirteen years in most species
(longer in humans). As a result, females
typically give birth only once every few
years. There is no distinct breeding
season.[32]
 The gorillas and the common
chimpanzee live in family groups of
around five to ten individuals, although
much larger groups are sometimes
noted. Chimpanzees live in larger
groups that break up into smaller
groups when fruit becomes less
available. When small groups of female
chimpanzees go off in separate
directions to forage for fruit, the
dominant males can no longer control
them and the females often mate with
other subordinate males. In contrast,
groups of gorillas stay together
regardless of the availability of fruit.
When fruit is hard to find, they resort to
eating leaves and shoots. Because
gorilla groups stay together, the male is
able to monopolize the females in his
group. This fact is related to gorillas'
greater sexual dimorphism relative to
that of chimpanzees; that is, the
difference in size between male and
female gorillas is much larger than that
of male and female chimpanzees. This
enables gorilla males to physically
dominate female gorillas more easily. In
both chimpanzees and gorillas, the
groups include at least one dominant
male, and females leave the group at
maturity.
Homo habilis is a proposed archaic species
of Homo, which lived between roughly 2.1 and
1.5 million years ago, during the Gelasian and sophisticated Homo ergaster, which in turn
early Calabrian stages of gave rise to the more human-appearing
the Pleistocene geological epoch.[1] species, Homo erectus. Debates continue over
whether all of the known fossils are properly
The type specimen is OH 7, discovered in 1960
attributed to the species, and some
at Olduvai Gorge in Tanzania, associated with
paleoanthropologists regard the taxon as
the Oldowan lithic industry; the fossils were
invalid, made up of fossil specimens
identified as a separate species of Homo with
of Australopithecus and Homo.[9] New findings
the proposed binomial name of H.
in 2007 seemed to confirm the view that H.
habilis ("handy man") in 1964.[2] In its
habilisand H. erectus coexisted, representing
appearance and morphology, H. habilis is
separate lineages from a common ancestor
intermediate between Australopithecus and the
instead of H. erectus being descended from H.
somewhat younger Homo erectus and its
habilis.[10] An alternative explanation would be
classification in the genus Homo has been the
that any ancestral relationship from H.
subject of controversial debate since its original
habilisto H. erectus would have to have
proposal.[3] A main argument for its
been cladogenetic rather
classification as the first Homo("human")
than anagenetic (meaning that if an isolated
species was its use of flaked stone tools.
subgroup population of H. habilis became the
However, evidence for earlier tool use (3.39
ancestor of H. erectus, other subgroups
million years ago) by undisputed members
remained as unchanged H. habilis until their
of Australopithecus has been found in the
much later extinction).[11]
1990s.[4][5]
Discoveries at Dmanisi, Georgia, which had
Louis Leakey (father of Richard Leakey), the
diverse physical traits and differences in tooth
British-Kenyan paleoanthropologist who was
wear, suggest to some scholars that all the
the first to suggest the existence of H. habilis,
contemporary groups of early Homo in Africa,
and his wife, Mary Leakey, found the first trace
including Homo ergaster, Homo habilis,
of H. habilis in 1955: two hominin teeth. These
and Homo rudolfensis are of the same species
were later classified as "milk teeth", and
and should be assigned to Homo erectus, with
therefore considered difficult to link to taxa,
the implication that variation between these
unlike permanent teeth.
"species" represents the prolonged evolution of
H. habilis was short and had disproportionately one lineage, rather than interspecific
long arms compared to modern humans; differences.[12][13][14][15]
however, it had a less protruding face than
the australopithecines from which it is thought
to have descended. H. habilishad a cranial Morphology[edit]
capacity slightly less than half of the size of H. habilis brain size has been shown to range
modern humans. Despite the ape-like from 550 cm3 (34 cu in) to 687 cm3 (41.9 cu in),
morphology of the bodies, H. habilisremains rather than from 363 cm3 (22.2 cu in) to
are often accompanied by primitive stone 600 cm3 (37 cu in) as
tools (e.g. Olduvai Gorge, Tanzania and Lake previously[year needed] thought.[7][16]
Turkana, Kenya).
A virtual reconstruction published in 2015
Homo habilis has often been thought to be the estimated the endocranial volume at between
ancestor of the more gracile and 729 ml (25.7 imp fl oz; 24.7 US fl oz) and
824 ml (29.0 imp fl oz; 27.9 US fl oz), larger
than any previously published value.[17]
H. habilis' brain capacity of around
640 cm3 (39 cu in) was on average 50% larger
than australopithecines, but considerably
smaller than the 1,350 cm3 (82 cu in) to
1,450 cm3 (88 cu in) range of modern Homo
sapiens. These hominins were smaller than
modern humans, on average standing no more
than 1.3 m (4 ft 3 in).
The body proportions for H. habilis are in
accordance with craniodental evidence,
suggesting closer association with H. erectus.[18]
A 2018 study of the anatomy
of Australopithecus sediba found that A.
sediba is distinct from but closely related to
both Homo habilis and Australopithecus
africanus.[19]
Homo habilis is thought to have mastered the
Lower Paleolithic Olduwan tool set, which used
stone flakes. H. habilis used these stones to
butcher and skin animals.[23] These stone flakes
were more advanced than any tools previously
used, and gave H. habilis the edge it needed to
prosper in hostile environments previously too
formidable for primates. Whether H. habilis was
the first hominin to master stone tool
technology remains controversial,
as Australopithecus garhi, dated to 2.6 million
years ago, has been found along with stone
tool implements.
Most experts assume the intelligence and
social organization of H. habiliswere more
sophisticated than typical australopithecines
or chimpanzees. H. habilis used tools primarily
for scavenging, such as cleaving meat off
carrion, rather than defense or hunting. Yet,
despite tool usage, H. habiliswas not the
master hunter its sister species (or
descendants) proved to be, as ample fossil
evidence indicates H. habilis was a staple in
the diet of large predatory animals, such
as Dinofelis, a large scimitar-toothed
predatory cat the size of a jaguar.[24]
Homo habilis coexisted with other Homo-like
bipedal primates, such as Paranthropus boisei,
some of which prospered for many millennia.
However, H. habilis, possibly because of its
early tool innovation and a less specialized diet,
became the precursor of an entire line of new
species, whereas Paranthropus boisei and
its robust relatives disappeared from the fossil
record. H. habilis may also have coexisted
with H. erectus in Africa for a period of 500,000
years.[25]
Homo habilis, (Latin: “able man” or
“handy man”) extinctspecies of human,
the most ancient representative of the
human genus, Homo. Homo
habilis inhabited parts of sub-
Saharan Africa from roughly 2.4 to 1.5
million years ago (mya). In 1959 and
1960 the first fossils were discovered
at Olduvai Gorgein northern Tanzania.
This discovery was a turning point in the
science of paleoanthropology because
the oldest previously known human
fossils were Asian specimens of Homo
erectus. Many features of H.
habilis appear to be intermediate in terms
of evolutionary development between the
relatively primitive Australopithecus and
the more-advanced Homo species.
The first confirmed remains found at
Olduvai consist of several teeth and a
lower jaw associated with fragments of a
cranium and some hand bones. As more
specimens were unearthed at locations
such as Koobi Fora in northern Kenya,
researchers began to realize that these
hominins were anatomically different
from Australopithecus, a genus of more-
apelike creatures whose remains had been
found at many African sites. Formal
announcement of the discoveries was made
in 1964 by anthropologists Louis S.B.
Leakey, Phillip Tobias, and John Napier. As
justification for designating their new
creature Homorather than Australopithecus,
they described the increased cranial capacity
and comparatively smaller molar and
premolar teeth of the fossils, a
humanlike foot, and hand bones that
suggested an ability to manipulate objects
with precision—hence the species
name Homo habilis, or “handy man.”
Furthermore, simple stone tools were found
along with the fossils. All these
characteristics foreshadow the anatomy and
behaviour of H. erectus and later humans,
making H. habilisextremely important, even
though there are few remnants of it. Since
1964 more material has been discovered.
One intriguing specimen is OH 24, which was
also from Olduvai and dated to about 1.8
mya. This cranium is more complete than
others from Olduvai. Because some of the
bones are crushed and distorted, however,
the face and braincase are warped. OH 24
may differ from Australopithecus in brain size
and dental characteristics, but it resembles
the australopiths of southern Africa in other
features, such as the shape of the face.
Homo erectus (meaning 'upright man') is
a species of archaic humansthat lived
throughout most of the Pleistocene geological
epoch. Its earliest fossil evidence dates to 1.8
million years ago (discovered 1991
in Dmanisi, Georgia).[5] or more controversially
even older at 2.1 million years ago (discovered
2018 in the Loess Plateau, China).[6]
H. erectus has been hypothesized as a direct
ancestor of the later hominins including Homo
heidelbergensis, Homo antecessor, Homo
neanderthalensis, Homo denisova, and Homo
sapiens [7]
A debate regarding the classification, ancestry,
and progeny of H. erectus, especially in relation
to Homo ergaster, is ongoing, with two major
positions:
1) H. erectus is the same species as
African H. ergaster; or,
2) it is in fact an Asian species or
subspecies distinct from African H.
ergaster that would later undergo back-
migration to Africa, where it would
evolve eventually into modern homo
sapiens.[8][9]
Some paleoanthropologists consider H.
ergaster to be a variety, that is, the
"African" variety, of H. erectus; the
labels "Homo erectus sensu stricto"
(strict sense) for the Asian species and
"Homo erectus sensu lato" (broad
sense) have been offered for the
greater species comprising both Asian
and African populations.[10][11]
H. erectus eventually became extinct
throughout its range in Africa, Europe
and Asia, but developed into derived
species, notably Homo
heidelbergensis. As a chronospecies,
the time of its disappearance is thus a
matter of contention. The species
name proposed in 1950 defines Java
Man as the type specimen (now H. e.
erectus). Since then, there has been a
trend in palaeoanthropology of
reducing the number of proposed
species of Homo, to the point where H.
erectus includes all early (Lower ἄνθρωπος (ánthrōpos, "human")—based on
Paleolithic) forms of Homo sufficiently the proposal that the femur suggested that the
derived from H. habilis and distinct creature had been bipedal, like Homo sapiens.
from early H. heidelbergensis (in Africa
Dubois' 1891 find was the first fossil of a Homo-
also known as H. rhodesiensis).[12] In
species (or any hominin species) found as
this wider sense, H. erectus had mostly
result of a directed expedition and search (the
been replaced by H.
first recognized human fossil had been the
heidelbergensis by about 300,000
circumstantial discovery of Homo
years ago, with possible late survival in
neanderthalensis in 1856; see List of human
Java as late as 70,000 years
evolution fossils). The Java fossil from
ago.[1] The discovery of the
Indonesia aroused much public interest. It was
morphologically divergent Dmanisi
dubbed by the popular press as Java Man; but
skull 5 in 2013 has reinforced the trend
few scientists accepted Dubois' argument that
of subsuming fossils formerly given
his fossil was the transitional form—the so-
separate species names under H.
called "missing link"—between humans and
erectus considered as a wide-ranging,
nonhuman apes.[16]
polymorphous species.[13] Thus, H.
ergaster is now well within the
Poster of homo georgicus. National Archaeological
accepted morphological range of H.
erectus, and it has been suggested Museum of Spain.
that even H. rudolfensis and H.
habilis (alternatively suggested as late Most of the spectacular discoveries of H.
forms of Australopithecus rather than erectus next took place at the Zhoukoudian
early Homo) should be considered Project, now known as the Peking Man site, in
early varieties of H. erectus Zhoukoudian, China. This site was first
discovered by Johan Gunnar Andersson in
The Dutch anatomist Eugène Dubois, inspired
1921[17] and was first excavated in 1921, and
by Darwin's theory of evolution as it applied to produced two human teeth.[18] Davidson Black's
humanity, set out in 1886 for Asia (despite
initial description (1921) of a lower molar as
Darwin's theory of African origin) to find a belonging to a previously unknown species
human ancestor. In 1891–92, his team
(which he named Sinanthropus
discovered first a tooth, then a skullcap, and
pekinensis)[19] prompted widely publicized
finally a femur of a human fossil on the island interest. Extensive excavations followed, which
of Java, Dutch East Indies (now Indonesia).
altogether uncovered 200 human fossils from
Excavated from the bank of the Solo
more than 40 individuals including five nearly
River at Trinil, in East Java, he first (1893)
complete skullcaps.[20] Franz
allocated the material to a genus of fossil
Weidenreich provided much of the detailed
chimpanzees as Anthropopithecus erectus,
description of this material in several
then the following year assigned his species to
monographs published in the
a new genus as Pithecanthropus erectus (the
journal Palaeontologica Sinica (Series D).
genus name had been coined by Ernst
Haeckel in 1868 for the hypothetical link Nearly all of the original specimens were lost
between humans and fossil Apes)—from the during World War II during an attempt to
Greek πίθηκος (píthēkos, "ape") and smuggle them out of China for safekeeping;
however, authentic casts were made by
Weidenreich, which exist at the American
Museum of Natural History in New York
City and at the Institute of Vertebrate
Paleontology and Paleoanthropology in Beijing,
and are considered to be reliable evidence.
Similarities between Java Man and Peking
Man led Ernst Mayr to rename both Homo
erectus in 1950.
Throughout much of the 20th century,
anthropologists debated the role of H.
erectus in human evolution. Early in the
century, due in part to the discoveries at Java
and Zhoukoudian, the belief that modern
humans first evolved in Asia was widely
accepted. A few naturalists—Charles Darwin
most prominent among them—theorized that
humans' earliest ancestors were African:
Darwin pointed out that chimpanzees and
gorillas, humans' closest relatives, evolved and
exist only in Africa.[21]
Origin and dispersal[edit]
Further information: Early human expansions
out of Africa and Acheulean
Map of the distribution of Middle Pleistocene
(Acheulean) cleaver finds
The derivation of the
genus Homo from Australopithecina took place
in East Africa after 3 million years ago. The
inclusion of species dated to just before 2
million years ago, Homo habilis and Homo
rudolfensis, into Homo is somewhat
contentious.[22] Especially as H. habilis appears
to have coexisted with H. ergaster/erectus for a
substantial period after 2 Mya, it has been
proposed that ergaster may not be directly
derived from habilis.[23]
Homo erectus emerged about 2 million years
ago. Fossils dated close to 1.8 million years
ago have been found both in Africa and in West
Asia, so it is unclear whether H.
erectus emerged in Africa or in Asia. Ferring et
al. (2011) suggest that it was still H.
habilis which reached West Asia, and that
early H. erectusdeveloped there. Early H.
erectus would then have dispersed from West
Asia, to East Asia (Peking Man) Southeast Asia
(Java Man), back to Africa (Homo ergaster),
and to Europe (Tautavel Man).[24][25]
Africa[edit]
Main article: Homo ergaster
KNM-ER 3733 (1.6 Mya, discovered 1975 at Koobi
Fora, Kenya)
In the 1950s, archaeologists John T.
Robinson and Robert
Broom named Telanthropus
capensis;[26] Robinson had discovered a jaw
fragment in 1949 in Swartkrans, South Africa.
Later,[when?] Simonetta[who?] proposed to re-
designate it to Homo erectus, and Robinson
agreed.[27]
From the 1950s forward, numerous finds in
East Africa suggested sympatriccoexistence
for H. ergaster and H. habilis for several
hundred millennia, which tends to confirm the
hypothesis that they represent separate
lineages from a common ancestor; that is, the
ancestral relationship between them was
not anagenetic, but was cladogenetic, which
here suggests that a subgroup population of H.
habilis—or of a common ancestor of H.
habilis and H. ergaster/erectus—
became reproductively isolated from the main-
group population, eventually evolving into the
new species Homo ergaster (Homo erectus
sensu lato).[28]
In 1961, Yves Coppens discovered a skull in
northern Chad. He coined the
name Tchadanthropus uxoris for what he
considered the earliest fossil human discovered
in north Africa.[29] Although once considered to
be a specimen of H. habilis,[30] T. uxoris has
been subsumed into H. erectus but it is no
longer considered a valid taxon.[29][31] It was
reported that the fossil "had been so eroded by
wind-blown sand that it mimicked the
appearance of an australopith, a primitive type
of hominid".[32] It is probably only 10,000 years
old according
to stratigraphy, paleontology and C14
dating presented in Michel Servant's PhD as
early as 1973.[33][better source needed][clarification needed]
Eurasia[edit]
Caucasus[edit]
Main article: Dmanisi skulls
Dmanisi skull 3 (fossils skull D2700 and jaw D2735,
two of several found in Dmanisi in
the GeorgianTranscaucasus)
Reconstruction of Homo georgicus based on D2700,
by Élisabeth Daynès, Museo de la Evolución
Humana, Burgos, Spain.
Homo erectus georgicus is the subspecies
name assigned to fossil skulls and jaws found
in Dmanisi, Georgia. First proposed as a
separate species, it is now classified within H.
erectus.[34][35][36] The site was discovered in 1991
by Georgian scientist David Lordkipanidze. Five
skulls were excavated from 1991 forward,
including a "very complete" skull in 2005.
Excavations at Dmanisi have yielded 73 stone
tools for cutting and chopping and 34 bone
fragments from unidentified fauna.[37]
After their initial assessment, some scientists
were persuaded to name the Dmanisi find as a
new species, Homo georgicus, which they
posited as a descendant of African Homo
habilis and an ancestor to Asian Homo erectus.
This classification, however, was not
supported, and the fossil was instead
designated a divergent subgroup of Homo
erectus.[38][39][40][41]
The fossil skeletons present a species primitive
in its skull and upper body but with relatively
advanced spine and lower limbs, implying
greater mobility than the previous
morphology.[42] It is now thought not to be a
separate species, but to represent a stage soon
after the transition between H. habilis to H.
erectus; it has been dated at 1.8 Mya.[35][43] The
assemblage includes one of the
largest Pleistocene Homo mandibles (D2600),
one of the smallest Lower
Pleistocenemandibles (D211), a nearly
complete sub-adult (D2735), and a toothless
specimen D3444/D3900.[44]
Two of the skulls—D2700, with a brain volume
of 600 cubic centimetres (37 cu in), and D4500
or Dmanisi Skull 5, with a brain volume of about
546 centimetres—present the two smallest and
most primitive Hominina skulls from the
Pleistocene period.[14] The variation in these
skulls were compared to variations in modern
humans and within a sample group of
chimpanzees. The researchers found that,
despite appearances, the variations in the
Dmanisi skulls were no greater than those seen
among modern people and among
chimpanzees. These findings suggest that
previous fossil finds that were classified as
different species on the basis of the large
morphological variation among them—
including Homo rudolfensis, Homo
gautengensis, H. ergaster, and potentially
even H. habilis—should perhaps be re-
classified to the same lineage as Homo
erectus.[45]
East and Southeast Asia[edit]
H. erectus is attested with certainty in East and
Southeast Asia from about 0.7 Ma, with
possible early presence before 1 Ma; stone
tools from Shangchen discovered in 2018 were
even claimed to be older than 2 Mya.[46][47]
Meganthropus refers to a group of fossils found
in Java, dated to between 1.4 and 0.9 Mya,
which are tentatively grouped with H. erectus,
at least in the wider sense of the term in which
"all earlier Homo populations that are
sufficiently derived from African
early Homo belong to H. erectus",[12] although
older literature has placed the fossils outside
of Homo altogether.[48]
Java Man (H. e. erectus, the type specimen
for H. erectus), discovered on the island
of Java in 1891/2, is dated to 1.0–0.7
Mya. Lantian Man (H. e. lantianensis),
discovered in 1963 in Lantian
County, Shaanxi province, China, is roughly
contemporary with Java Man.
Peking Man (H. e. pekinensis), discovered in
1923–27 at Zhoukoudian (Chou K'ou-tien)
near Beijing, China, dates to about 0.75
Mya.[49] and a new 26Al/10Be dating suggests
they are in the range of 680,000–780,000 years
old.[50][51] Yuanmou Man (H. e. yuanmouensis),
discovered in Yuanmou County in Yunnan,
China, in 1965, is likely of similar age as Peking
Man (but with dates proposed as early as 1.7
Mya).[52]
Nanjing Man (H. e. nankinensis), discovered in
1993 in the Hulu cave on the Tangshan hills
near Nanjing, dates to about 0.6 Mya.[53][54]
Solo Man (H. e. soloensis), discovered
between 1931/1933 along the Solo River, Java,
is of uncertain age, dated between 0.25 and
0.075 Mya (the younger date would qualify Solo
Man as the latest fossil still classified as H.
erectus, even though it shows some derived
features, notably larger cranial capacity).[55]
Europe[edit]
Further information: Homo
antecessor and Homo heidelbergensis
It is conventional to label European archaic
humans contemporary with late H.
erectus under the separate species name
of Homo heidelbergensis, the immediate
predecessor of Homo neanderthalensis. H.
heidelbergensis fossils are recorded from 600
ka (Mauer 1 mandible), the oldest complete
skulls are "Tautavel Man" (Homo erectus
tautavelensis), c. 450 ka, and the Atapuerca
skull ("Miguelón"), c. 430 ka. The oldest known
human fossils found in Europe is a molar from
the Sima del Elefante site, Atapuerca
Mountains, Spain, dated c. 1.2 Mya. This is
associated with the skull fragments of the "Boy
of Gran Dolina" found nearby, dated 0.9 Mya
and classified as Homo antecessor by the
discoverers. The relationship of these fossils
to H. erectus is open to debate, as no complete
skull has been found. H. e.
bilzingslebenensis (Vlcek 1978) refers to skull
fragments found at
the Bilzingsleben site, Thuringia, Germany.
There is, however, indirect evidence of human
presence in Europe as early as 1.6 Mya, in the
form of stone tools discovered at Lézignan-la-
Cèbe, France, in 2008.[56] Other stone tools
found in France and thought to predate 1 Mya
are from Chilhac, Haute-Loire) and from
the Grotte du Vallonnet, near Menton. Human
presence in Great Britain close to 1 Mya is
established by stone tools and fossilized
footprints found near Happisburgh, Norfolk.[57][58]
Homo sapiens is the only
extant human species. The name is Latin for
"wise man" and was introduced in 1758 by Carl
Linnaeus (who is himself the lectotype for the
species).
Extinct species of the
genus Homo include Homo erectus, extant
during roughly 1.9 to 0.4 million years ago, and
a number of other species (by some authors
considered subspecies of either H.
sapiens or H. erectus). The age
of speciation of H. sapiens out of ancestral H.
erectus (or an intermediate species such
as Homo antecessor) is estimated to have
been roughly 350,000 years ago.[note
1]
Sustained archaic admixture is known to have
taken place both in Africa and (following
the recent Out-Of-Africa expansion) in Eurasia,
between about 100,000 and 30,000 years
ago.[4]
The term anatomically modern
humans[5] (AMH) is used to distinguish H.
sapiens having an anatomy consistent with
the range of phenotypesseen in contemporary
humans from varieties of extinct archaic
humans. This is useful especially for times and
regions where anatomically modern and
archaic humans co-existed, for example,
in Paleolithic Europe.
Name and taxonomy
Main article: Human taxonomy
Further information: Homo and Names for the
human species
The binomial name Homo sapiens was coined
by Linnaeus, 1758.[6] The Latin noun homō (gen
itive hominis) means "human being", while the
participle sapiēns means "discerning, wise,
sensible".
Age and speciation process
Further information: Human
The species was initially thought to have
evolution, Homo, Timeline of human evolution,
emerged from a predecessor within the
and Early human migrations
genus Homo around 300,000 to 200,000 years
ago.[note 2] A problem with the morphological
classification of "anatomically modern" was that Schematic representation of the emergence of H.
it would not have included certain extant
sapiens from earlier species of Homo. The
populations. For this reason, a lineage-based
(cladistic) definition of H. sapiens has been horizontal axis represents geographic location; the
suggested, in which H. sapiens would by vertical axis represents time in millions of years
definition refer to the modern human lineage
following the split from the Neanderthal lineage. ago (blue areas denote the presence of a certain
Such a cladistic definition would extend the age species of Homoat a given time and place; late
of H. sapiens to over 500,000 years.[note 3] survival of robust
Extant human populations have historically australopithecines alongside Homo is indicated in
been divided into subspecies, but since around
purple). Based on Springer (2012), Homo
the 1980s all extant groups have tended to be
subsumed into a single species, H. sapiens, heidelbergensis[3] is shown as diverging into
avoiding division into subspecies altogether.[note
4] Neanderthals, Denisovans and H. sapiens. With the
rapid expansion of H. sapiens after 60 kya,
Some sources show Neanderthals (H.
neanderthalensis) as a subspecies (H. sapiens Neanderthals, Denisovans and unspecified archaic
neanderthalensis).[14][15]Similarly, the discovered
African hominins are shown as again subsumed into
specimens of the H. rhodesiensis species have
been classified by some as a subspecies (H. the H. sapiens lineage.
sapiens rhodesiensis), although it remains
more common to treat these last two as Derivation from H. erectus
separate species within the genus Homo rather Further information: Homo antecessor, Homo
than as subspecies within H. sapiens.[16] heidelbergensis, Homo rhodesiensis,
and Acheulean
The subspecies name H. sapiens sapiens is
sometimes used informally instead of "modern
humans" or "anatomically modern humans". It A model of the phylogeny of H. sapiens during
has no formal authority associated with it.[note the Middle Paleolithic. The horizontal axis
5]
By the early 2000s, it had become common to
use H. s. sapiens for the ancestral population of represents geographic location; the vertical axis
all contemporary humans, and as such it is represents time in thousands of years ago.[note
equivalent to the binomial H. sapiens in the 1] Neanderthals, Denisovans and unspecified archaic
more restrictive sense (considering H.
neanderthalensis a separate species).[note 6] African hominins are shown as admixed into the H.
sapiens lineage. In addition, prehistoric Archaic
Human and Eurasian admixture events in modern
African populations are indicated.
The speciation of H. sapiens out of archaic
human varieties derived from H. erectus is
estimated as having taken place over 350,000
years ago, as the Khoisan split from other
populations is dated between 260,000 and
350,000 years ago.[20]
An alternative suggestion defines H.
sapiens cladistically as including the lineage of
modern humans since the split from the lineage
of Neanderthals, roughly 500,000 to 800,000
years ago.
The time of divergence between archaic H.
sapiens and ancestors of Neanderthals and
Denisovans caused by a genetic bottleneck of
the latter was dated at 744,000 years ago,
combined with repeated early admixture events
and Denisovans diverging from Neanderthals
300 generations after their split from H.
sapiens, as calculated by Rogers et al.
(2017).[21]
The derivation of a comparatively
homogeneous single species of H.
sapiens from more diverse varieties of archaic
humans (all of which were descended from
the early dispersal of H. erectus some 1.8
million years ago) was debated in terms of two
competing models during the 1980s: "recent
African origin" postulated the emergence of H.
sapiensfrom a single source population in
Africa, which expanded and led to the
extinction of all other human varieties, while the
"multiregional evolution" model postulated the
survival of regional forms of archaic humans,
gradually converging into the modern human
varieties by the mechanism of clinal variation,
via genetic drift, gene
flow and selectionthroughout the Pleistocene.[22]
Since the 2000s, the availability of data
from archaeogenetics and population
genetics has led to the emergence of a much
more detailed picture, intermediate between the
two competing scenarios outlined above:
The recent Out-of-Africa expansion accounts
for the predominant part of modern human
ancestry, while there were also
significant admixture events with regional
archaic humans.[23][24]
Since the 1970s, the Omo remains, dated to
some 195,000 years ago, have often been
taken as the conventional cut-off point for the
emergence of "anatomically modern humans".
Since the 2000s, the discovery of older remains
with comparable characteristics, and the
discovery of ongoing hybridization between
"modern" and "archaic" populations after the
time of the Omo remains, have opened up a
renewed debate on the age of H. sapiens in
journalistic publications.[25][26][27][28][29] H. s. idaltu,
dated to 160,000 years ago, has been
postulated as an extinct subspecies of H.
sapiens in 2003.[30][better source needed] H.
neanderthalensis, which became extinct about
40,000 years ago, has also been classified as a
subspecies, H. s. neanderthalensis.
H. heidelbergensis, dated 600,000 to 300,000
years ago, has long been thought to be a likely
candidate for the last common ancestor of the
Neanderthal and modern human lineages.
However, genetic evidence from the Sima de
los Huesos fossils published in 2016 seems to
suggest that H. heidelbergensis in its entirety
should be included in the Neanderthal lineage,
as "pre-Neanderthal" or "early Neanderthal",
while the divergence time between the
Neanderthal and modern lineages has been
pushed back to before the emergence of H.
heidelbergensis, to close to 800,000 years ago,
the approximate time of disappearance of H.
antecessor.[31][32]
Early Homo sapiens
Further information: Human subspecies, Middle
Paleolithic, Mousterian, Archaic human
admixture with modern humans, Homo sapiens
idaltu, and Skhul and Qafzeh hominins
Skhul V (dated at about 80,000–120,000 years old)
exhibiting a mix of archaic and modern traits.
The term Middle Paleolithic is intended to cover
the time between the first emergence of H.
sapiens (roughly 300,000 years ago) and the
emergence of full behavioral modernity (roughly
50,000 years ago, corresponding to the start of
the Upper Paleolithic).
Many of the early modern human finds, like
those of Omo, Herto, Skhul, Jebel
Irhoud and Peștera cu Oase exhibit a mix of
archaic and modern traits.[33][34][35] Skhul V, for
example, has prominent brow ridges and a
projecting face. However, the brain case is
quite rounded and distinct from that of the
Neanderthals and is similar to the brain case of
modern humans. It is uncertain whether the
robust traits of some of the early modern
humans like Skhul V reflects mixed ancestry or
retention of older traits.[36][37]
The "gracile" or lightly built skeleton of
anatomically modern humans has been
connected to a change in behavior, including
increased cooperation and "resource
transport".[38][39]
There is evidence that the characteristic human
brain development, especially the prefrontal
cortex, was due to "an exceptional acceleration
of metabolome evolution ... paralleled by a
drastic reduction in muscle strength. The
observed rapid metabolic changes in brain and
muscle, together with the unique human
cognitive skills and low muscle performance,
might reflect parallel mechanisms in human
evolution."[40] The Schöningen spears and their
correlation of finds are evidence that complex
technological skills already existed 300,000
years ago, and are the first obvious proof of an
active (big game) hunt. H.
heidelbergensis already had intellectual and
cognitive skills like anticipatory planning,
thinking and acting that so far have only been
attributed to modern man.[41][42]
The ongoing admixture events within
anatomically modern human populations make
it difficult to estimate the age of the matrilinear
and patrilinear most recent common ancestors
of modern populations (Mitochondrial
Eveand Y-chromosomal Adam). Estimates of
the age of Y-chromosomal Adam have been
pushed back significantly with the discovery of
an ancient Y-chromosomal lineage in 2013, to
likely beyond 300,000 years ago.[note 7]There
have, however, been no reports of the survival
of Y-chromosomal or mitochondrial DNA clearly
deriving from archaic humans (which would
push back the age of the most recent patrilinear
or matrilinear ancestor beyond 500,000
years).[44][45][46]
Fossil teeth found at Qesem Cave (Israel) and
dated to between 400,000 and 200,000 years
ago have been compared to the dental material
from the younger (120,000–80,000 years
ago) Skhul and Qafzeh hominins.[note 8]
Dispersal and archaic
admixture
Further information: Recent African origin of
modern humans, Southern Dispersal, Early
human migrations, and List of first human
settlements
Further information: Interbreeding between
archaic and modern humans
Overview map of the peopling of the world by
anatomically modern humans (numbers indicate
dates in thousands of years ago [ka])
Dispersal of early H. sapiensbegins soon after
its emergence, as evidenced by the North
African Jebel Irhoud finds (dated to between
280,000 and 350,000 years ago).[35] There is
indirect evidence for modern human presence
in West Asia around 270,000 years ago
and Dali Manfrom China is dated at 260,000
years ago.[48]
Among extant populations, the Khoi-San (or
"Capoid") hunters-gatherers of Southern Africa
may represent the human population with the
earliest possible divergence within the
group Homo sapiens sapiens. Their separation
time has been estimated in a 2017 study to be
as long as between 260,000 and 350,000 years
ago, compatible with the estimated age of H.
sapiens.[2] H. s. idaltu, found at Middle Awash in
Ethiopia, lived about 160,000 years
ago,[49] and H. sapiens lived at Omo Kibish in
Ethiopia about 195,000 years ago.[50] Fossil
evidence for modern human presence in West
Asia is ascertained for 177,000 years
ago,[51] and disputed fossil evidence suggests
expansion as far as East Asia by 120,000 years
ago.[52][53]
In July 2019, anthropologists reported the
discovery of 210,000 year old remains of a H.
sapiens and 170,000 year old remains of a H.
neanderthalensis in Apidima
Cave, Peloponnese, Greece, more than
150,000 years older than previous H.
sapiens finds in Europe.[54][55][56]
A significant dispersal event, within Africa and
to West Asia, is associated with the
African megadroughts during MIS 5, beginning
130,000 years ago.[57] A 2011 study located the
origin of basal population of contemporary
human populations at 130,000 years ago, with
the Khoi-San representing an "ancestral
population cluster" located in southwestern
Africa (near the coastal border
of Namibia and Angola).[58]
Layer sequence at Ksar Akil in the Levantine
corridor, and discovery of two fossils of Homo
sapiens, dated to 40,800 to 39,200 years BP for
"Egbert",[59]and 42,400–41,700 BP for "Ethelruda".[59]
While early modern human expansion in Sub-
Saharan Africa before 130 kya persisted, early
expansion to North Africa and Asia appears to
have mostly disappeared by the end of MIS5
(75,000 years ago), and is known only from
fossil evidence and from archaic admixture.
Asia was re-populated by early modern
humans in the so-called "recent out-of-Africa
migration" post-dating MIS5, beginning around
70,000 years ago. In this expansion, bearers
of mt-DNA haplogroup L3 left East Africa, likely
reaching Arabia via the Bab-el-Mandeb, and in
the Great Coastal Migration spread to South
Asia, Maritime South Asia and Oceania by
65,000 years
ago,[60][61] while Europe, East and North Asia,
and possibly the Americas, were reached by
50,000 years ago.
Evidence for the overwhelming contribution of
this "recent" (L3-derived) expansion to all non-
African populations was established based
on mitochondrial DNA, combined with evidence
based on physical anthropology of
archaic specimens, during the 1990s and
2000s.[note 9][63] The assumption of complete
replacement has been revised in the 2010s
with the discovery of admixture
events (introgression) of populations of H.
sapiens with populations of archaic humans
over the period of between roughly 100,000
and 30,000 years ago, both in Eurasia and in
Sub-Saharan Africa. Neanderthal admixture, in
the range of 1-4%, is found in all modern
populations outside of Africa, including in
Europeans, Asians, Papua New Guineans,
Australian Aboriginals, Native Americans, and
other non-Africans.[64][23] This suggests that
interbreeding between Neanderthals and
anatomically modern humans took place after
the recent "out of Africa" migration, likely
between 60,000 and 40,000 years
ago.[65][66][67] Recent admixture analyses have
added to the complexity, finding that Eastern
Neanderthals derive up to 2% of their ancestry
from anatomically modern humans who left
Africa some 100 kya.[68] The extent
of Neanderthal admixture (and introgression of
genes acquired by admixture) varies
significantly between contemporary racial
groups, being absent in Africans, intermediate
in Europeans and highest in East Asians.
Certain genes related to UV-light adaptation
introgressed from Neanderthals have been
found to have been selected for in East Asians
specifically from 45,000 years ago until around
5,000 years ago.[69] The extent of archaic
admixture is of the order of about 1% to 4% in
Europeans and East Asians, and highest
among Melanesians (the last also
having Denisova hominin admixture at 4% to
6% in adition to neanderthal
admixture).[23][36] Cumulatively, about 20% of the
Neanderthal genome is estimated to remain
present spread in contemporary populations.[70]
Anatomy
See also: Human anatomy, Human physical
appearance, and Anthropometry
Known archaeological remains of Anatomically
Modern Humans in Europe and Africa, directly
dated, calibrated carbon dates as of 2013.[71]
Generally, modern humans are more lightly
built (or more "gracile") than the more
"robust" archaic humans. Nevertheless,
contemporary humans exhibit high variability in
many physiological traits, and may exhibit
remarkable "robustness". There are still a
number of physiological details which can be
taken as reliably differentiating the physiology
of Neanderthals vs. anatomically modern
humans.
Anatomical modernity
See also: Behavioral modernity
The term "anatomically modern humans"
(AMH) is used with varying scope depending
on context, to distinguish "anatomically
modern" Homo sapiens from archaic
humans such as Neanderthals and Middle
and Lower Paleolithic hominins with transitional
features intermediate between H. erectus,
Neanderthals and early AMH called archaic
Homo sapiens.[72][73] In a convention popular in
the 1990s, Neanderthals were classified as
a subspecies of H. sapiens, as H. s.
neanderthalensis, while AMH (or European
early modern humans, EEMH) was taken to
refer to "Cro-Magnon" or H. s. sapiens. Under
this nomenclature (Neanderthals considered H.
sapiens), the term "anatomically modern Homo
sapiens" (AMHS) has also been used to refer to
EEMH ("Cro-Magnons").[74] It has since become
more common to designate Neanderthals as a
separate species, H. neanderthalensis, so that
AMH in the European context refers to H.
sapiens (but the question is by no means
resolved[note 10]).
In this more narrow definition of H. sapiens, the
subspecies H. s. idaltu, discovered in 2003,
also falls under the umbrella of "anatomically
modern".[76] The recognition of H. s. idaltu as
a valid subspecies of the anatomically modern
human lineage would justify the description of
contemporary humans with the subspecies
name H. s. sapiens.
A further division of AMH into "early" or "robust"
vs. "post-glacial" or "gracile" subtypes has
since been used for convenience. The
emergence of "gracile AMH" is taken to reflect
a process towards a smaller and more fine-
boned skeleton beginning around 50,000–
30,000 years ago.[77]
Braincase anatomy
Further information: Brain size
Anatomical comparison of skulls of H. sapiens(left)
and H. neanderthalensis (right)
(in Cleveland Museum of Natural History)
Features compared are
the braincase shape, forehead, browridge, nasal
bone, projection, cheek bone
angulation, chin and occipital contour
The cranium lacks a pronounced occipital
bun in the neck, a bulge that anchored
considerable neck muscles in Neanderthals.
Modern humans, even the earlier ones,
generally have a larger fore-brain than the
archaic people, so that the brain sits above
rather than behind the eyes. This will usually
(though not always) give a higher forehead,
and reduced brow ridge. Early modern people
and some living people do however have quite
pronounced brow ridges, but they differ from
those of archaic forms by having both
a supraorbital foramen or notch, forming a
groove through the ridge above each
eye.[78] This splits the ridge into a central part
and two distal parts. In current humans, often
only the central section of the ridge is
preserved (if it is preserved at all). This
contrasts with archaic humans, where the brow
ridge is pronounced and unbroken.[79]
Modern humans commonly have a steep, even
vertical forehead whereas their predecessors
had foreheads that sloped strongly
backwards.[80] According to Desmond Morris,
the vertical forehead in humans plays an
important role in human communication
through eyebrow movements and forehead skin
wrinkling.[81]
Brain size in both Neanderthals and AMH is
significantly larger on average (but overlapping
in range) than brain size in H. erectus.
Neanderthal and AMH brain sizes are in the
same range, but there are differences in the
relative sizes of individual brain areas, with
significantly larger visual systems in
Neanderthals than in AMH.[82][note 11]
Jaw anatomy
Compared to archaic people, anatomically
modern humans have smaller, differently
shaped teeth.[85][86] This results in a smaller,
more receded dentary, making the rest of the
jaw-line stand out, giving an often quite
prominent chin. The central part of the
mandible forming the chin carries a triangularly
shaped area forming the apex of the chin called
the mental trigon, not found in archaic
humans.[87] Particularly in living populations, the
use of fire and tools requires fewer jaw
muscles, giving slender, more gracile jaws.
Compared to archaic people, modern humans
have smaller, lower faces.
Body skeleton structure
The body skeletons of even the earliest and
most robustly built modern humans were less
robust than those of Neanderthals (and from
what little we know from Denisovans), having
essentially modern proportions. Particularly
regarding the long bones of the limbs, the distal
bones (the radius/ulna and tibia/fibula) are
nearly the same size or slightly shorter than the
proximal bones (the humerus and femur). In
ancient people, particularly Neanderthals, the
distal bones were shorter, usually thought to be
an adaptation to cold climate.[88]The same
adaptation can be found in some modern
people living in the polar regions.[89]
Height ranges overlap between Neanderthals
and AMH, with Neanderthal averages cited as
164 to 168 cm (65 to 66 in) and 152 to 156 cm
(60 to 61 in) for males and females,
respectively.[note 12] By
comparison, contemporary national
averages range between 158 to 184 cm (62 to
72 in) in males and 147 to 172 cm (58 to 68 in)
in females. Neanderthal ranges approximate
the height distribution measured among Malay
people, for one.[note 13]
Recent evolution
Main article: Recent human evolution
Further information: Human genetic
variability, Race and genetics, and Sexual
selection in humans
Following the peopling of Africa some 130,000
years ago, and the recent Out-of-
Africa expansion some 70,000 to 50,000 years
ago, some sub-populations of H. sapiens have
been essentially isolated for tens of thousands
of years prior to the early modern Age of
Discovery. Combined with archaic
admixture this has resulted in
significant genetic variation, which in some
instances has been shown to be the result
of directional selectiontaking place over the
past 15,000 years, i.e. significantly later than
possible archaic admixture events.[92]
Some climatic adaptations, such as high-
altitude adaptation in humans, are thought to
have been acquired by archaic
admixture. Introgression of genetic variants
acquired by Neanderthal admixture have
different distributions in European and East
Asians, reflecting differences in recent selective
pressures. A 2014 study reported that
Neanderthal-derived variants found in East
Asian populations showed clustering in
functional groups related
to immune and haematopoietic pathways, while
European populations showed clustering in
functional groups related to the lipid catabolic
process.[note 14] A 2017 study found correlation
of Neanderthal admixture in phenotypic traits in
modern European populations.[94]
Physiological or phenotypical changes have
been traced to Upper Paleolithic mutations,
such as the East Asian variant of
the EDAR gene, dated to c. 35,000 years
ago.[note 15]
Recent divergence of Eurasian lineages was
sped up significantly during the Last Glacial
Maximum, the Mesolithic and the Neolithic, due
to increased selection pressures and due to
founder effects associated
with migration.[97] Alleles predictive of light
skin have been found in Neanderthals,[98] but
the alleles for light skin in Europeans and East
Asians, associated with KITLG and ASIP, are
(as of 2012) thought to have not been acquired
by archaic admixture but recent mutations
since the LGM.[97] Phenotypes associated with
the "white" or "Caucasian" populations of
Western Eurasian stock emerge during the
LGM, from about 19,000 years ago.
Average cranial capacity in modern human
populations varies in the range of 1,200 to
1,450 cm3 (adult male averages). Larger cranial
volume is associated with climatic region, the
largest averages being found in populations
of Siberia and the Arctic.[note
16][100]
Both Neanderthal and EEMH had
somewhat larger cranial volumes on average
than modern Europeans, suggesting the
relaxation of selection pressures for larger brain
volume after the end of the LGM.[99]
Examples for still later adaptations related
to agriculture and animal
domestication including East Asian types
of ADH1B associated with rice
domestication,[101] or lactase
persistence,[102][103] are due to recent selection
pressures.
An even more recent adaptation has been
proposed for the Austronesian Sama-Bajau,
developed under selection pressures
associated with subsisting on freediving over
the past thousand years or so.[104][105]
Behavioral modernity
Lithic Industries of early Homo sapiens at Blombos
Cave (M3 phase, MIS 5), Southern Cape, South
Africa (c. 105 – 90 Ka)
Behavioral modernity, involving the
development of language, figurative art and
early forms of religion(etc.) is taken to have
arisen before 40,000 years ago, marking the
beginning of the Upper Paleolithic (in African
contexts also known as the Later Stone
Age).[106]
There is considerable debate regarding
whether the earliest anatomically modern
humans behaved similarly to recent or existing
humans. Behavioral modernity is taken to
include fully developed language(requiring the
capacity for abstract thought), artistic
expression, early forms of religious
behavior,[107]increased cooperation and the
formation of early settlements, and the
production of articulated tools from lithic cores,
bone or antler. The term Upper Paleolithic is
intended to cover the period since the rapid
expansion of modern humans throughout
Eurasia, which coincides with the first
appearance of Paleolithic art such as cave
paintings and the development of technological
innovation such as the spear-thrower. The
Upper Paleolithic begins around 50,000 to
40,000 years ago, and also coincides with the
disappearance of archaic humans such as
the Neanderthals.
Bifacial silcrete point of early Homo sapiens, from
M1 phase (71,000 BCE) layer of Blombos Cave,
South Africa
The term "behavioral modernity" is somewhat
disputed. It is most often used for the set of
characteristics marking the Upper Paleolithic,
but some scholars use "behavioral modernity"
for the emergence of H. sapiens around
200,000 years ago,[108] while others use the
term for the rapid developments occurring
around 50,000 years ago.[109][110][111] It has been
proposed that the emergence of behavioral
modernity was a gradual process.[112][113][114][115][116]
In January 2018, it was announced that modern
human finds at Misliya cave, Israel, in 2002,
had been dated to around 185,000 years ago,
the earliest evidence of their out of Africa
migration.[117][118][119][120]
The earliest H. sapiens (AMH) found
in Europe are the "Cro-Magnon" (named after
the site of first discovery in France), beginning
about 40,000 to 35,000 years ago. These are
also known as "European early modern
humans" in contrast to the
preceding Neanderthals.[121][122]
Claimed "Oldest known drawing by human hands",
discovered in Blombos Cave in South Africa.
Estimated to be a 73,000 years old work of a Homo
sapiens.[123]
The International Space Station, one of the latest
creations of Homo sapiens
The equivalent of the Eurasian Upper
Paleolithic in African archaeology is known as
the Later Stone Age, also beginning roughly
40,000 years ago. While most clear evidence
for behavioral modernity uncovered from the
later 19th century was from Europe, such as
the Venus figurines and other artefacts from
the Aurignacian, more recent archaeological
research has shown that all essential elements
of the kind of material culture typical of
contemporary San hunter-gatherers
in Southern Africa was also present by least
40,000 years ago, including digging sticks of
similar materials used today, ostrich egg shell
beads, bone arrow heads with individual
maker's marks etched and embedded with red
ochre, and poison applicators.[124] There is also
a suggestion that "pressure flaking best
explains the morphology of lithic artifacts
recovered from the c. 75-ka Middle Stone Age
levels at Blombos Cave, South Africa. The
technique was used during the final shaping of
Still Bay bifacial points made on heat‐treated
silcrete."[125] Both pressure flaking and heat
treatment of materials were previously thought
to have occurred much later in prehistory, and
both indicate a behaviourally modern
sophistication in the use of natural materials.
Further reports of research on cave sites along
the southern African coast indicate that "the
debate as to when cultural and cognitive
characteristics typical of modern humans first
appeared" may be coming to an end, as
"advanced technologies with elaborate chains
of production" which "often demand high-fidelity
transmission and thus language" have been
found at Pinnacle Point Site 5–6. These have
been dated to approximately 71,000 years ago.
The researchers suggest that their research
"shows that microlithic technology originated
early in South Africa, evolved over a vast time
span (c. 11,000 years), and was typically
coupled to complex heat treatment that
persisted for nearly 100,000 years. Advanced
technologies in Africa were early and enduring;
a small sample of excavated sites in Africa is
the best explanation for any perceived
'flickering' pattern."[126] These results suggest
that Late Stone Age foragers in Sub-Saharan
Africa had developed modern cognition and
behaviour by at least 50,000 years ago.[127] The
change in behavior has been speculated to
have been a consequence of an earlier climatic
change to much drier and colder conditions
between 135,000 and 75,000 years
ago.[128] This might have led to human groups
who were seeking refuge from the inland
droughts, expanded along the coastal marshes
rich in shellfish and other resources. Since sea
levels were low due to so much water tied up
in glaciers, such marshlands would have
occurred all along the southern coasts of
Eurasia. The use of rafts and boats may well
have facilitated exploration of offshore islands
and travel along the coast, and eventually
permitted expansion to New Guinea and then
to Australia.