n taxonomy, 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 also the type specimen).
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.
The term anatomically modern humans[4] (AMH) is used to distinguish H. sapiens having
an anatomy consistent with the range of phenotypes seen 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.

Contents

 1Name and taxonomy

 2Age and speciation process
o 2.1Derivation from H. erectus
o 2.2Early Homo sapiens
 3Dispersal and archaic admixture
 4Anatomy
o 4.1Anatomical modernity
o 4.2Braincase anatomy
o 4.3Jaw anatomy
o 4.4Body skeleton structure
 5Recent evolution
 6Behavioral modernity
 7Notes
 8References
 9Further reading
 10External links

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.[5] The Latin noun homō (genitive hominis) means "human being", while the
participle sapiēns means "discerning, wise, sensible".
The species was initially thought to have emerged from a predecessor within the
genus Homo around 300,000 to 200,000 years ago.[note 2] A problem with the morphological
classification of "anatomically modern" was that it would not have included certain extant
populations. For this reason, a lineage-based (cladistic) definition of H. sapiens has been suggested,
in which H. sapiens would by definition refer to the modern human lineage following the split from
the Neanderthal lineage. Such a cladistic definition would extend the age of H. sapiens to over
500,000 years.[note 3]
Extant human populations have historically been divided into subspecies, but since around the
1980s all extant groups have tended to be subsumed into a single species, H. sapiens, avoiding
division into subspecies altogether.[note 4]
Some sources show Neanderthals (H. neanderthalensis) as a subspecies (H. sapiens
neanderthalensis).[13][14] Similarly, the discovered specimens of the H. rhodesiensis species have
been classified by some as a subspecies (H. sapiens rhodesiensis), although it remains more
common to treat these last two as separate species within the genus Homo rather than as
subspecies within H. sapiens.[15]
The subspecies name H. sapiens sapiens is sometimes used informally instead of "modern humans"
or "anatomically modern humans". It has no formal authority associated with it.[note 5] By the early
2000s, it had become common to use H. s. sapiens for the ancestral population of all contemporary
humans, and as such it is equivalent to the binomial H. sapiens in the more restrictive sense
(considering H. neanderthalensis a separate species).[note 6]

Age and speciation process

Further information: Human evolution, Homo, Timeline of human evolution, and Early human
migrations

Schematic representation of the emergence of H. sapiens from earlier species of Homo. The horizontal axis
represents geographic location; the vertical axis represents time in millions of years ago (blue areas denote the
presence of a certain species of Homoat a given time and place; late survival of robust
australopithecines alongside Homo is indicated in purple). Based on Springer (2012), Homo
heidelbergensis[3] is shown as diverging into Neanderthals, Denisovans and H. sapiens. With the rapid
expansion of H. sapiens after 60 kya, Neanderthals, Denisovans and unspecified archaic African hominins are
shown as again subsumed into the H. sapiens lineage.

Derivation from H. erectus

Further information: Homo antecessor, Homo heidelbergensis, Homo rhodesiensis, and Acheulean
A model of the phylogeny of H. sapiens during the Middle Paleolithic. The horizontal axis represents
geographic location; the vertical axis represents time in thousands of years ago.[note 1] Neanderthals, Denisovans
and unspecified archaic 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.[19]
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).[20]
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. sapiens from 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.[21]
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.[22][23]
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.[24][25][26][27][28] H. s.
idaltu, dated to 160,000 years ago, has been postulated as an extinct subspecies of H. sapiens in
2003.[29][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.[30][31]

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.[32][33][34] 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.[35][36]
The "gracile" or lightly built skeleton of anatomically modern humans has been connected to a
change in behavior, including increased cooperation and "resource transport".[37][38]
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."[39] 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.[40][41]
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 Eve and 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).[43][44][45]
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. sapiens begins soon after its emergence, as evidenced by the North
African Jebel Irhoud finds (dated to between 280,000 and 350,000 years ago).[34] There is indirect
evidence for modern human presence in West Asia around 270,000 years ago and Dali Man from
China is dated at 260,000 years ago.[47]
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 site Middle Awash in Ethiopia, lived about 160,000 years ago,[48] and H. Sapiens lived at
Omo Kibish in Ethiopia about 195,000 years ago.[49] Fossil evidence for modern human presence in
West Asia is ascertained for 177,000 years ago,[50] and disputed fossil evidence suggests expansion
as far as East Asia by 120,000 years ago.[51][52]
A significant dispersal event, within Africa and to West Asia, is associated with the
African megadroughts during MIS 5, beginning 130,000 years ago.[53] 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).[54]
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",[55]and 42,400–41,700 BP for "Ethelruda".[55]

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,[56][57] 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][59]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, Papuan New Guineans, Australian Aboriginals, and Native
Americans.[60][22] 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.[61][62][63] 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.[64] 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.[65] The extent of archaic admixture is
of the order of about 1% to 4% in Europeans and East Asians, and highest
among Melanesians (Denisova hominin admixture), at 4% to 6%.[22][35] Cumulatively, about 20% of
the Neanderthal genome is estimated to remain present spread in contemporary populations.[66]

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.[67]
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.[68][69] 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").[70] 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".[72] 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.[73]

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.[74] 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.[75]
Modern humans commonly have a steep, even vertical forehead whereas their predecessors had
foreheads that sloped strongly backwards.[76] According to Desmond Morris, the vertical forehead in
humans plays an important role in human communication through eyebrow movements and
forehead skin wrinkling.[77]
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.[78][note 11]

Jaw anatomy
Compared to archaic people, anatomically modern humans have smaller, differently shaped
teeth.[81][82] 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.[83] 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.[84] The same adaptation can be found in some
modern people living in the polar regions.[85]
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 approximating the height distribution
measured, e.g., among Malay people.[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 selection taking place over the past 15,000
years, i.e. significantly later than possible archaic admixture events.[88]
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.[90]
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.[93] Alleles predictive of light skin have been found in Neanderthals,[94] 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.[93] 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][96] 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.[95]
Examples for still later adaptations related to agriculture and animal domestication including East
Asian types of ADH1B associated with rice domestication,[97] or lactase persistence,[98][99] 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.[100][101]