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Early modern human (EMH) or anatomically modern human (AMH) are terms used to distinguish ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture ...
'' (the only extant Hominina species) that are anatomically consistent with the range of phenotypes seen in contemporary humans from extinct archaic human species. This distinction is useful especially for times and regions where anatomically modern and archaic humans co-existed, for example, in Paleolithic Europe. Among the oldest known remains of ''Homo sapiens'' are those found at the
Omo-Kibish I The Omo remains are a collection of homininThis article quotes historic texts that use the terms 'hominid' and 'hominin' with meanings that may be different from their modern usages. This is because several revisions in classifying the great apes h ...
archaeological site in south-western Ethiopia, dating to about 233,000 to 196,000 years ago, the Florisbad site in South Africa, dating to about 259,000 years ago, and the
Jebel Irhoud Jebel Irhoud or Adrar n Ighoud ( zgh, ⴰⴷⵔⴰⵔ ⵏ ⵉⵖⵓⴷ, Adrar n Iɣud; ar, جبل إيغود, žbəl iġud), is an archaeological site located just north of the locality known as Tlet Ighoud, approximately south-east of the cit ...
site in Morocco, dated about 315,000 years ago.
Extinct species This page features lists of extinct species, organisms that have become Extinction, extinct, either in the wild or completely disappeared from Earth. In actual theoretical practice, a species not definitely located in the wild in the last fifty ...
of the genus ''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus '' Australopithecus'' that encompasses the extant species ''Homo sapiens'' ( modern humans), plus several extinct species classified as either ancestral to or closely rela ...
'' include ''
Homo erectus ''Homo erectus'' (; meaning "upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as '' H. heidelbergensis'' and '' H. antecessor ...
'' (extant from roughly 2 to 0.1 million years ago) and a number of other species (by some authors considered subspecies of either ''H. sapiens'' or ''H. erectus''). The divergence of the lineage leading to ''H. sapiens'' out of ancestral ''H. erectus'' (or an intermediate species such as '' Homo antecessor'') is estimated to have occurred in Africa roughly 500,000 years ago. The earliest fossil evidence of early modern humans appears in
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
around 300,000 years ago, with the earliest genetic splits among modern people, according to some evidence, dating to around the same time. Sustained archaic human admixture with modern humans 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.


Name and taxonomy

The binomial name ''Homo sapiens'' was coined by Linnaeus, 1758. The
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
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. 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. Estimates for the split between the Homo sapiens line and combined
Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an Extinction, extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ag ...
/ Denisovan line range from between 503,000 and 565,000 years ago; between 550,000 and 765,000 years ago; and (based on rates of dental evolution) possibly more than 800,000 years ago. 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. Some sources show Neanderthals (''H. neanderthalensis'') as a subspecies (''H. sapiens neanderthalensis''). 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''. All humans are considered to be a part of the subspecies '' H. sapiens sapiens'', a designation which has been a matter of debate since a species is usually not given a subspecies category unless there is evidence of multiple distinct subspecies.


Age and speciation process


Derivation from ''H. erectus''

The divergence of the lineage that would lead to ''H. sapiens'' out of archaic human varieties derived from ''H. erectus'', is estimated as having taken place over 500,000 years ago (marking the split of the ''H. sapiens'' lineage from ancestors shared with other known archaic hominins). But the oldest split among modern human populations (such as the
Khoisan Khoisan , or (), according to the contemporary Khoekhoegowab orthography, is a catch-all term for those indigenous peoples of Southern Africa who do not speak one of the Bantu languages, combining the (formerly "Khoikhoi") and the or ( in ...
split from other groups) has been recently dated to between 350,000 and 260,000 years ago, and the earliest known examples of ''H. sapiens'' fossils also date to about that period, including the
Jebel Irhoud Jebel Irhoud or Adrar n Ighoud ( zgh, ⴰⴷⵔⴰⵔ ⵏ ⵉⵖⵓⴷ, Adrar n Iɣud; ar, جبل إيغود, žbəl iġud), is an archaeological site located just north of the locality known as Tlet Ighoud, approximately south-east of the cit ...
remains from Morocco (ca. 300,000 or 350–280,000 years ago), the Florisbad Skull from South Africa (ca. 259,000 years ago), and the Omo remains from Ethiopia (ca. 195,000, or, as more recently dated, ca. 233,000 years ago). An mtDNA study in 2019 proposed an origin of modern humans in Botswana (and a Khoisan split) of around 200,000 years. However, this proposal has been widely criticized by scholars, with the recent evidence overall (genetic, fossil, and archaeological) supporting an origin for ''H. sapiens'' approximately 100,000 years earlier and in a broader region of Africa than the study proposes. In September 2019, scientists proposed that the earliest ''H. sapiens'' (and last common human ancestor to modern humans) arose between 350,000 and 260,000 years ago through a merging of populations in East and
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
. An alternative suggestion defines ''H. sapiens'' cladistically as including the lineage of modern humans since the split from the lineage of
Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an Extinction, extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ag ...
s, 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). 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 The multiregional hypothesis, multiregional evolution (MRE), or polycentric hypothesis is a scientific model that provides an alternative explanation to the more widely accepted "Out of Africa" model of monogenesis for the pattern of human evoluti ...
" model postulated the survival of regional forms of archaic humans, gradually converging into the modern human varieties by the mechanism of
clinal variation Cline may refer to: Science * Cline (biology), a measurable gradient in a single trait in a species across its geographical range * Cline (hydrology), a fluid layer with a property that varies * Cline (mathematics) or generalised circle, a circ ...
, via
genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
,
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
and selection throughout the Pleistocene. Since the 2000s, the availability of data from archaeogenetics and
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and po ...
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. Since the 1970s, the Omo remains, originally 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. ''H. s. idaltu'', dated to 160,000 years ago, has been postulated as an extinct subspecies of ''H. sapiens'' in 2003. ''H. neanderthalensis'', which became extinct about 40,000 years ago, was also at one point considered to be 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''.


Early ''Homo sapiens''

The term Middle Paleolithic is intended to cover the time between the first emergence of ''H. sapiens'' (roughly 300,000 years ago) and the period held by some to mark the emergence of full behavioral modernity (roughly by 50,000 years ago, corresponding to the start of the Upper Paleolithic). Many of the early modern human finds, like those of
Jebel Irhoud Jebel Irhoud or Adrar n Ighoud ( zgh, ⴰⴷⵔⴰⵔ ⵏ ⵉⵖⵓⴷ, Adrar n Iɣud; ar, جبل إيغود, žbəl iġud), is an archaeological site located just north of the locality known as Tlet Ighoud, approximately south-east of the cit ...
,
Omo Omo or OMO may refer to: Geography Ethiopia * Omo River (Ethiopia), in southern Ethiopia is the largest Ethiopian river outside the Nile Basin and namesake for all the topics below * Omo Nada, one of the woredas in the Oromia Region of Ethiopia * ...
, Herto, Florisbad, Skhul, and Peștera cu Oase exhibit a mix of archaic and modern traits. 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 Mixed race people are people of more than one race or ethnicity. A variety of terms have been used both historically and presently for mixed race people in a variety of contexts, including ''multiethnic'', ''polyethnic'', occasionally ''bi-ethn ...
or retention of older traits. The "gracile" or lightly built skeleton of anatomically modern humans has been connected to a change in behavior, including increased cooperation and "resource transport". 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." 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. 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. 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). 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.


Dispersal and archaic admixture

Dispersal of early ''H. sapiens'' begins soon after its emergence, as evidenced by the North African
Jebel Irhoud Jebel Irhoud or Adrar n Ighoud ( zgh, ⴰⴷⵔⴰⵔ ⵏ ⵉⵖⵓⴷ, Adrar n Iɣud; ar, جبل إيغود, žbəl iġud), is an archaeological site located just north of the locality known as Tlet Ighoud, approximately south-east of the cit ...
finds (dated to around 315,000 years ago). There is indirect evidence for ''H. sapiens'' presence in West Asia around 270,000 years ago. The Florisbad Skull from Florisbad, South Africa, dated to about 259,000 years ago, has also been classified as representing early ''H. sapiens''. In September 2019, scientists proposed that the earliest ''H. sapiens'' (and last common human ancestor to modern humans) arose between 350,000 and 260,000 years ago through a merging of populations in East and
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
. Among extant populations, the
Khoi-San Khoisan , or (), according to the contemporary Khoekhoegowab orthography, is a catch-all term for those indigenous peoples of Southern Africa who do not speak one of the Bantu languages, combining the (formerly "Khoikhoi") and the or ( in t ...
(or "
Capoid Capoid race is a grouping formerly used for the Khoikhoi and San peoples in the context of a now-outdated model of dividing humanity into different races. The term was introduced by Carleton S. Coon in 1962 and named for the Cape of Good Hope.'' ...
") 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 between 350 and 260,000 years ago, compatible with the estimated age of early ''H. sapiens''. The study states that the deep split-time estimation of 350 to 260 thousand years ago is consistent with the archaeological estimate for the onset of the Middle Stone Age across sub-Saharan Africa and coincides with archaic ''H. sapiens'' in southern Africa represented by, for example, the Florisbad skull dating to 259 (± 35) thousand years ago. ''H. s. idaltu'', found at Middle Awash in Ethiopia, lived about 160,000 years ago, and ''H. sapiens'' lived at Omo Kibish in Ethiopia about 233,000-195,000 years ago. Two fossils from Guomde, Kenya, dated to at least (and likely more than) 180,000 years ago and (more precisely) to 300–270,000 years ago, have been tentatively assigned to ''H. sapiens'' and similarities have been noted between them and the Omo Kibbish remains. Fossil evidence for modern human presence in West Asia is ascertained for 177,000 years ago, and disputed fossil evidence suggests expansion as far as East Asia by 120,000 years ago. 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 Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders wi ...
, more than 150,000 years older than previous ''H. sapiens'' finds in Europe. A significant dispersal event, within Africa and to West Asia, is associated with the African megadroughts during MIS 5, beginning 130,000 years ago. 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 Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
and
Angola , national_anthem = "Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordinat ...
). While early modern human expansion in
Sub-Saharan Africa Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the List of sov ...
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 There is evidence for interbreeding between archaic and modern humans during the Middle Paleolithic and early Upper Paleolithic. The interbreeding happened in several independent events that included Neanderthals and Denisovans, as well as sever ...
. Eurasia was re-populated by early modern humans in the so-called "recent out-of-Africa migration" post-dating MIS5, beginning around 70,000–50,000 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 In the context of the recent African origin of modern humans, the Southern Dispersal scenario (also the coastal migration or great coastal migration hypothesis) refers to the early migration along the southern coast of Asia, from the Arabian Pen ...
spread to South Asia, Maritime South Asia and Oceania between 65,000 and 50,000 years ago, while
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
, East and
North Asia North Asia or Northern Asia, also referred to as Siberia, is the northern region of Asia, which is defined in geographical terms and is coextensive with the Asian part of Russia, and consists of three Russian regions east of the Ural Mountains ...
were reached by about 45,000 years ago. Some evidence suggests that an early wave humans may have reached the Americas by about 40,000–25,000 years ago. Evidence for the overwhelming contribution of this "recent" ( L3-derived) expansion to all non-African populations was established based on
mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...
, combined with evidence based on physical anthropology of archaic specimens, during the 1990s and 2000s, and has also been supported by
Y DNA The Y chromosome is one of two sex chromosomes (allosomes) in therian mammals, including humans, and many other animals. The other is the X chromosome. Y is normally the sex-determining chromosome in many species, since it is the presence or ...
and
autosomal DNA An autosome is any chromosome that is not a sex chromosome. The members of an autosome pair in a diploid cell have the same morphology, unlike those in allosomal (sex chromosome) pairs, which may have different structures. The DNA in autosomes ...
. 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 There is evidence for interbreeding between archaic and modern humans during the Middle Paleolithic and early Upper Paleolithic. The interbreeding happened in several independent events that included Neanderthals and Denisovans, as well as seve ...
, 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. 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. 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. The extent of
Neanderthal admixture There is evidence for interbreeding between archaic and modern humans during the Middle Paleolithic and early Upper Paleolithic. The interbreeding happened in several independent events that included Neanderthals and Denisovans, as well as seve ...
(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. The extent of archaic admixture is of the order of about 1% to 4% in Europeans and East Asians, and highest among
Melanesians Melanesians are the predominant and indigenous inhabitants of Melanesia, in a wide area from Indonesia's New Guinea to as far East as the islands of Vanuatu and Fiji. Most speak either one of the many languages of the Austronesian language fam ...
(the last also having Denisova hominin admixture at 4% to 6% in addition to neanderthal admixture). Cumulatively, about 20% of the Neanderthal genome is estimated to remain present spread in contemporary populations. 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 350,000 and 260,000 years ago through a merging of populations in East and
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
while North-African fossils may represent a population which introgressed into Neandertals during the LMP.


Anatomy

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 Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago. While ...
vs. anatomically modern humans.


Anatomical 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''. 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"). 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. In this more narrow definition of ''H. sapiens'', the subspecies '' Homo sapiens idaltu'', discovered in 2003, also falls under the umbrella of "anatomically modern". The recognition of ''H. sapiens idaltu'' as a valid subspecies of the anatomically modern human lineage would justify the description of contemporary humans with the subspecies name ''Homo sapiens sapiens''. However, biological anthropologist
Chris Stringer Christopher Brian Stringer (born 1947) is a British physical anthropologist noted for his work on human evolution. Biography Growing up in a working-class family in the East End of London, Stringer's interest in anthropology began in primar ...
does not consider ''idaltu'' distinct enough within ''H. sapiens'' to warrant its own subspecies designation. A further division of AMH into "early" or "robust" vs. "post-glacial" or "
gracile Gracility is slenderness, the condition of being gracile, which means slender. It derives from the Latin adjective ''gracilis'' ( masculine or feminine), or ''gracile'' ( neuter), which in either form means slender, and when transferred for exa ...
" 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.


Braincase anatomy

The cranium lacks a pronounced
occipital bun The occipital bone () is a cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cere ...
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. 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. Modern humans commonly have a steep, even vertical forehead whereas their predecessors had foreheads that sloped strongly backwards. According to
Desmond Morris Desmond John Morris FLS ''hon. caus.'' (born 24 January 1928) is an English zoologist, ethologist and surrealist painter, as well as a popular author in human sociobiology. He is known for his 1967 book ''The Naked Ape'', and for his telev ...
, the vertical forehead in humans plays an important role in human communication through
eyebrow An eyebrow is an area of short hairs above each eye that follows the shape of the lower margin of the brow ridges of some mammals. In humans, eyebrows serve two main functions: first, communication through facial expression, and second, prev ...
movements and forehead skin wrinkling. 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.


Jaw anatomy

Compared to archaic people, anatomically modern humans have smaller, differently shaped teeth. 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. 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 In classical geometry, a radius (plural, : radii) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', ...
/ ulna and
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it conn ...
/
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
) are nearly the same size or slightly shorter than the proximal bones (the humerus and
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates ...
). In ancient people, particularly Neanderthals, the distal bones were shorter, usually thought to be an adaptation to cold climate. The same adaptation is found in some modern people living in the polar regions.
Height Height is measure of vertical distance, either vertical extent (how "tall" something or someone is) or vertical position (how "high" a point is). For example, "The height of that building is 50 m" or "The height of an airplane in-flight is ab ...
ranges overlap between Neanderthals and AMH, with Neanderthal averages cited as and for males and females, respectively, which is largely identical to pre-industrial average heights for AMH. Contemporary national averages range between in males and in females. Neanderthal ranges approximate the contemporary height distribution measured among
Malay people Malays ( ms, Orang Melayu, Jawi: أورڠ ملايو) are an Austronesian ethnic group native to eastern Sumatra, the Malay Peninsula and coastal Borneo, as well as the smaller islands that lie between these locations — areas that are c ...
, for one.


Recent evolution

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'' had been essentially
isolated Isolation is the near or complete lack of social contact by an individual. Isolation or isolated may also refer to: Sociology and psychology *Isolation (health care), various measures taken to prevent contagious diseases from being spread **Is ...
for tens of thousands of years prior to the early modern
Age of Discovery The Age of Discovery (or the Age of Exploration), also known as the early modern period, was a period largely overlapping with the Age of Sail, approximately from the 15th century to the 17th century in European history, during which seafa ...
. Combined with
archaic admixture There is evidence for interbreeding between archaic and modern humans during the Middle Paleolithic and early Upper Paleolithic. The interbreeding happened in several independent events that included Neanderthals and Denisovans, as well as sever ...
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. 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 There is evidence for interbreeding between archaic and modern humans during the Middle Paleolithic and early Upper Paleolithic. The interbreeding happened in several independent events that included Neanderthals and Denisovans, as well as seve ...
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. A 2017 study found correlation of
Neanderthal admixture There is evidence for interbreeding between archaic and modern humans during the Middle Paleolithic and early Upper Paleolithic. The interbreeding happened in several independent events that included Neanderthals and Denisovans, as well as seve ...
in phenotypic traits in modern European populations. Physiological or phenotypical changes have been traced to Upper Paleolithic mutations, such as the East Asian variant of the
EDAR Ectodysplasin A receptor (EDAR) is a protein that in humans is encoded by the EDAR gene. EDAR is a cell surface receptor for ectodysplasin A which plays an important role in the development of ectodermal tissues such as the skin. It is structura ...
gene, dated to c. 35,000 years ago. Recent divergence of Eurasian lineages was sped up significantly during the
Last Glacial Maximum The Last Glacial Maximum (LGM), also referred to as the Late Glacial Maximum, was the most recent time during the Last Glacial Period that ice sheets were at their greatest extent. Ice sheets covered much of Northern North America, Northern Eu ...
(LGM), the Mesolithic and the
Neolithic The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several pa ...
, due to increased selection pressures and due to founder effects associated with migration. Alleles predictive of light skin have been found in
Neanderthals Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago. While ...
, but the alleles for light skin in Europeans and East Asians, associated with
KITLG Stem cell factor (also known as SCF, KIT-ligand, KL, or steel factor) is a cytokine that binds to the c-KIT receptor (CD117). SCF can exist both as a transmembrane protein and a soluble protein. This cytokine plays an important role in hematopoie ...
and ASIP, are () thought to have not been acquired by archaic admixture but recent mutations since the LGM. Phenotypes associated with the "
white White is the lightest color and is achromatic (having no hue). It is the color of objects such as snow, chalk, and milk, and is the opposite of black. White objects fully reflect and scatter all the visible wavelengths of light. 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 for adult males. Larger cranial volume is associated with climatic region, the largest averages being found in populations of
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part ...
and the
Arctic The Arctic ( or ) is a polar regions of Earth, polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenla ...
. Both
Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an Extinction, extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ag ...
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. Examples for still later adaptations related to
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people ...
and
animal domestication The domestication of animals is the mutual relationship between non-human animals and the humans who have influence on their care and reproduction. Charles Darwin recognized a small number of traits that made domesticated species different fro ...
including East Asian types of ADH1B associated with rice domestication, or lactase persistence, 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 Freediving, free-diving, free diving, breath-hold diving, or skin diving is a form of underwater diving that relies on breath-holding until resurfacing rather than the use of breathing apparatus such as scuba gear. Besides the limits of breath- ...
over the past thousand years or so.


Behavioral modernity

Behavioral modernity, involving the development of
language Language is a structured system of communication. The structure of a language is its grammar and the free components are its vocabulary. Languages are the primary means by which humans communicate, and may be conveyed through a variety of ...
, figurative art and early forms of
religion Religion is usually defined as a social- cultural system of designated behaviors and practices, morals, beliefs, worldviews, texts, sanctified places, prophecies, ethics, or organizations, that generally relates humanity to supernatur ...
(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). 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 Language is a structured system of communication. The structure of a language is its grammar and the free components are its vocabulary. Languages are the primary means by which humans communicate, and may be conveyed through a variety of ...
(requiring the capacity for abstract thought),
artistic expression Art is a diverse range of human activity, and resulting product, that involves creative or imaginative talent expressive of technical proficiency, beauty, emotional power, or conceptual ideas. There is no generally agreed definition of wh ...
, early forms of religious behavior, 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 The art of the Upper Paleolithic represents the oldest form of prehistoric art. Figurative art is present in Europe and Southeast Asia, beginning between about 40,000 to 35,000 years ago. Non-figurative cave paintings, consisting of hand ...
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
Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an Extinction, extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ag ...
s. 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, while others use the term for the rapid developments occurring around 50,000 years ago. It has been proposed that the emergence of behavioral modernity was a gradual process.


Examples of behavioural modernity

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 Southern Africa is the southernmost subregion of the African continent, south of the Congo and Tanzania. The physical location is the large part of Africa to the south of the extensive Congo River basin. Southern Africa is home to a number o ...
was also present by at least 40,000 years ago, including digging sticks of similar materials used today,
ostrich egg The egg of the ostrich (genus ''Struthio'') is the largest of any living bird. The shell has a long history of use by humans as a container and for decorative artwork. The eggs are not commonly eaten. Biology The female common ostrich lays her ...
shell beads, bone arrow heads with individual maker's marks etched and embedded with red
ochre Ochre ( ; , ), or ocher in American English, is a natural clay earth pigment, a mixture of ferric oxide and varying amounts of clay and sand. It ranges in colour from yellow to deep orange or brown. It is also the name of the colours produced ...
, and poison applicators. 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." 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 the South African 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 by 71 kya, 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 Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
were early and enduring; a small sample of excavated sites in Africa is the best explanation for any perceived 'flickering' pattern." Increases in behavioral complexity have been speculated to have been a linked to an earlier climatic change to much drier conditions between 135,000 and 75,000 years ago. 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
glacier A glacier (; ) is a persistent body of dense ice that is constantly moving under its own weight. A glacier forms where the accumulation of snow exceeds its ablation over many years, often centuries. It acquires distinguishing features, such a ...
s, 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 Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. With an area of , Australia is the largest country by ...
. In addition, a variety of other evidence of abstract imagery, widened subsistence strategies, and other "modern" behaviors has been discovered in Africa, especially South, North, and East Africa, predating 50,000 years ago (with some predating 100,000 years ago). The Blombos Cave site in South Africa, for example, is famous for rectangular slabs of ochre engraved with geometric designs. Using multiple dating techniques, the site was confirmed to be around 77,000 and 100,000–75,000 years old. Ostrich egg shell containers engraved with geometric designs dating to 60,000 years ago were found at Diepkloof, South Africa. Beads and other personal ornamentation have been found from Morocco which might be as much as 130,000 years old; as well, the Cave of Hearths in South Africa has yielded a number of beads dating from significantly prior to 50,000 years ago, and shell beads dating to about 75,000 years ago have been found at Blombos Cave, South Africa. Specialized projectile weapons as well have been found at various sites in Middle Stone Age Africa, including bone and stone arrowheads at South African sites such as Sibudu Cave (along with an early bone needle also found at Sibudu) dating approximately 72,000–60,000 years ago some of which may have been tipped with poisons, and bone harpoons at the Central African site of Katanda dating ca. 90,000 years ago. Evidence also exists for the systematic heat treating of silcrete stone to increase its flake-ability for the purpose of toolmaking, beginning approximately 164,000 years ago at the South African site of Pinnacle Point and becoming common there for the creation of microlithic tools at about 72,000 years ago. In 2008, an ochre processing workshop likely for the production of paints was uncovered dating to ca. 100,000 years ago at Blombos Cave, South Africa. Analysis shows that a liquefied pigment-rich mixture was produced and stored in the two abalone shells, and that ochre, bone, charcoal, grindstones and hammer-stones also formed a composite part of the toolkits. Evidence for the complexity of the task includes procuring and combining raw materials from various sources (implying they had a mental template of the process they would follow), possibly using pyrotechnology to facilitate fat extraction from bone, using a probable recipe to produce the compound, and the use of shell containers for mixing and storage for later use. Modern behaviors, such as the making of shell beads, bone tools and arrows, and the use of ochre pigment, are evident at a Kenyan site by 78,000-67,000 years ago. Evidence of early stone-tipped projectile weapons (a characteristic tool of ''Homo sapiens''), the stone tips of javelins or throwing spears, were discovered in 2013 at the Ethiopian site of
Gademotta The Gademotta Formation in the Main Ethiopian Rift Valley is known for its Middle Stone Age archaeological sites. It is located west of Lake Ziway. In addition to the type-site, which assumes the same name, the formation contains a cluster of s ...
, and date to around 279,000 years ago. Expanding subsistence strategies beyond big-game hunting and the consequential diversity in tool types have been noted as signs of behavioral modernity. A number of South African sites have shown an early reliance on aquatic resources from fish to shellfish. Pinnacle Point, in particular, shows exploitation of marine resources as early as 120,000 years ago, perhaps in response to more arid conditions inland. Establishing a reliance on predictable shellfish deposits, for example, could reduce mobility and facilitate complex social systems and symbolic behavior. Blombos Cave and Site 440 in Sudan both show evidence of fishing as well. Taphonomic change in fish skeletons from Blombos Cave have been interpreted as capture of live fish, clearly an intentional human behavior. Humans in North Africa ( Nazlet Sabaha,
Egypt Egypt ( ar, مصر , ), officially the Arab Republic of Egypt, is a List of transcontinental countries, transcontinental country spanning the North Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via a land bridg ...
) are known to have dabbled in
chert Chert () is a hard, fine-grained sedimentary rock composed of microcrystalline or cryptocrystalline quartz, the mineral form of silicon dioxide (SiO2). Chert is characteristically of biological origin, but may also occur inorganically as a ...
mining Mining is the extraction of valuable minerals or other geological materials from the Earth, usually from an ore body, lode, vein, seam, reef, or placer deposit. The exploitation of these deposits for raw material is based on the econom ...
, as early as ≈100,000 years ago, for the construction of
stone tool A stone tool is, in the most general sense, any tool made either partially or entirely out of stone. Although stone tool-dependent societies and cultures still exist today, most stone tools are associated with prehistoric (particularly Stone A ...
s. Evidence was found in 2018, dating to about 320,000 years ago at the site of
Olorgesailie Olorgesailie is a geological formation in East Africa, on the floor of the Eastern Rift Valley in southern Kenya, southwest of Nairobi along the road to Lake Magadi. It contains a group of Lower Paleolithic archaeological sites. Olorgesailie ...
in Kenya, of the early emergence of modern behaviors including: the trade and long-distance transportation of resources (such as obsidian), the use of pigments, and the possible making of projectile points. The authors of three 2018 studies on the site observe that the evidence of these behaviors is roughly contemporary with the earliest known ''Homo sapiens'' fossil remains from Africa (such as at Jebel Irhoud and Florisbad), and they suggest that complex and modern behaviors began in Africa around the time of the emergence of ''Homo sapiens''. In 2019, further evidence of Middle Stone Age complex projectile weapons in Africa was found at Aduma, Ethiopia, dated 100,000–80,000 years ago, in the form of points considered likely to belong to darts delivered by spear throwers.


Pace of progress during ''Homo sapiens'' history

''Homo sapiens'' technological and cultural progress appears to have been very much faster in recent millennia than in ''Homo sapiens'' early periods. The pace of development may indeed have accelerated, due to massively larger population (so more humans extant to think of innovations), more communication and sharing of ideas among human populations, and the accumulation of thinking tools. However it may also be that the pace of advancements always looks relatively faster to humans in the time they live, because previous advances are unrecognised.


Notes


References


Further reading

*


External links


Human Timeline (Interactive)
Smithsonian,
National Museum of Natural History The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7. ...
(August 2016). * * {{authority control Humans Mammals described in 1758 Taxa named by Carl Linnaeus Tool-using mammals