fbpx
Wikipedia

Early modern human

January 31, 2023

This article is about the first Homo sapiens, specifically during the Old Stone Age. For a broader perspective on the human species, see Human.

Early modern human (EMH) or anatomically modern human (AMH)[1] are terms used to distinguish Homo sapiens (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 archaeological site in south-western Ethiopia, dating to about 233,000[2] to 196,000 years ago,[3] the Florisbad site in South Africa, dating to about 259,000 years ago, and the Jebel Irhoud site in Morocco, dated about 315,000 years ago.

100 to 80 thousand year old Skhul V from Israel

Extinct species of the genus Homo include Homo erectus (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 around 300,000 years ago, with the earliest genetic splits among modern people, according to some evidence, dating to around the same time.[4][5][note 1][8] 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.[9]

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.[10] 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]

Estimates for the split between the Homo sapiens line and combined Neanderthal/Denisovan line range from between 503,000 and 565,000 years ago;[15] between 550,000 and 765,000 years ago;[16] and (based on rates of dental evolution) possibly more than 800,000 years ago.[17]

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).[21][22] 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.[23]

All humans are considered to be a part of the subspecies H. sapiens sapiens,[24] 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.[24]

Age and speciation process

Further information: Human evolution, Homo, Multiregional origin of modern humans, Timeline of human evolution, and Early human migrations

Derivation from H. erectus

Further information: Homo ergaster, Homo antecessor, Homo heidelbergensis, Homo rhodesiensis, and Acheulean
 
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 Homo at a given time and place; late survival of robust australopithecines alongside Homo is indicated in purple). Based on Springer (2012), Homo heidelbergensis[7] 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.
 
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 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).[8][5] But the oldest split among modern human populations (such as the Khoisan split from other groups) has been recently dated to between 350,000 and 260,000 years ago,[25][26] and the earliest known examples of H. sapiens fossils also date to about that period, including the Jebel Irhoud remains from Morocco (ca. 300,000 or 350–280,000 years ago),[27] 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).[28][29][30][31][2]

An mtDNA study in 2019 proposed an origin of modern humans in Botswana (and a Khoisan split) of around 200,000 years.[32] However, this proposal has been widely criticized by scholars,[33][34][35] 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.[35]

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.[36][4]

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).[37]

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 selection throughout the Pleistocene.[38]

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.[39][40]

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.[41][42][43][44][45] H. s. idaltu, dated to 160,000 years ago, has been postulated as an extinct subspecies of H. sapiens in 2003.[46][24] H. neanderthalensis, which became extinct about 40,000 years ago, was also at one point considered to be a subspecies, H. s. neanderthalensis.[24]

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.[47][48]

Early Homo sapiens

Further information: Human subspecies, Middle Paleolithic, Mousterian, Interbreeding between archaic and modern humans, Homo sapiens idaltu, and Skhul and Qafzeh hominins

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, Omo, Herto, Florisbad, Skhul, and Peștera cu Oase exhibit a mix of archaic and modern traits.[49][50][27] 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.[51][52]

The "gracile" or lightly built skeleton of anatomically modern humans has been connected to a change in behavior, including increased cooperation and "resource transport".[53][54]

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."[55] 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.[56][57]

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 5] 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).[59][60][61]

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 6]

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 around 315,000 years ago).[27][30] There is indirect evidence for H. sapiens presence in West Asia around 270,000 years ago.[63]

The Florisbad Skull from Florisbad, South Africa, dated to about 259,000 years ago, has also been classified as representing early H. sapiens.[28][29][31][4]

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.[36][4]

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 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.[6]

H. s. idaltu, found at Middle Awash in Ethiopia, lived about 160,000 years ago,[64] and H. sapiens lived at Omo Kibish in Ethiopia about 233,000-195,000 years ago.[65][2] Two fossils from Guomde, Kenya, dated to at least (and likely more than) 180,000 years ago[28] and (more precisely) to 300–270,000 years ago,[4] have been tentatively assigned to H. sapiens and similarities have been noted between them and the Omo Kibbish remains.[28] Fossil evidence for modern human presence in West Asia is ascertained for 177,000 years ago,[66] and disputed fossil evidence suggests expansion as far as East Asia by 120,000 years ago.[67][68]

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.[69][70][71]

A significant dispersal event, within Africa and to West Asia, is associated with the African megadroughts during MIS 5, beginning 130,000 years ago.[72] 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).[73]

 
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",[74] and 42,400–41,700 BP for "Ethelruda".[74]

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. 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.[75][76][77][78] 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 between 65,000 and 50,000 years ago,[79][80][81][82] while Europe, East and North Asia 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.[citation needed]

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 7][84] and has also been supported by Y DNA and autosomal DNA.[78] 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.[85][39] 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.[86][87][88] 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.[89] 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.[90] 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 addition to neanderthal admixture).[39][51] Cumulatively, about 20% of the Neanderthal genome is estimated to remain present spread in contemporary populations.[91]

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 while North-African fossils may represent a population which introgressed into Neandertals during the LMP.[36][4]

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.[74]

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.[92] 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").[93] 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 8]

In this more narrow definition of H. sapiens, the subspecies Homo sapiens idaltu, discovered in 2003, also falls under the umbrella of "anatomically modern".[95] 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.[96] However, biological anthropologist Chris Stringer does not consider idaltu distinct enough within H. sapiens to warrant its own subspecies designation.[97][28]

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.[98]

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.[99] 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.[100]

Modern humans commonly have a steep, even vertical forehead whereas their predecessors had foreheads that sloped strongly backwards.[101] According to Desmond Morris, the vertical forehead in humans plays an important role in human communication through eyebrow movements and forehead skin wrinkling.[102]

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.[103][note 9]

Jaw anatomy

Compared to archaic people, anatomically modern humans have smaller, differently shaped teeth.[106][107] 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.[108] 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.[109] The same adaptation is found in some modern people living in the polar regions.[110]

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, which is largely identical to pre-industrial average heights for AMH.[note 10] 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 contemporary height distribution measured among Malay people, for one.[note 11]

Recent evolution

Main article: Recent human evolution
Further information: Human genetic variation, 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 had 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.[113]

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 12] A 2017 study found correlation of Neanderthal admixture in phenotypic traits in modern European populations.[115]

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 13]

Recent divergence of Eurasian lineages was sped up significantly during the Last Glacial Maximum (LGM), the Mesolithic and the Neolithic, due to increased selection pressures and due to founder effects associated with migration.[118] Alleles predictive of light skin have been found in Neanderthals,[119] 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.[118] 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 for adult males. Larger cranial volume is associated with climatic region, the largest averages being found in populations of Siberia and the Arctic.[note 14][121] 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.[120]

Examples for still later adaptations related to agriculture and animal domestication including East Asian types of ADH1B associated with rice domestication,[122] or lactase persistence,[123][124] 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.[125][126]

Behavioral modernity

 
Lithic Industries of early Homo sapiens at Blombos Cave (M3 phase, MIS 5), Southern Cape, South Africa (c. 105,000 – 90,000 years old)

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).[127]

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,[128] 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,[129] while others use the term for the rapid developments occurring around 50,000 years ago.[130][131][132] It has been proposed that the emergence of behavioral modernity was a gradual process.[133][134][135][136][137]

Examples of behavioural modernity

 
Claimed "oldest known drawing by human hands", discovered in Blombos Cave in South Africa. Estimated to be a 73,000-year-old work of a Homo sapiens.[138]

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 at 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.[139] 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."[140] 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 were early and enduring; a small sample of excavated sites in Africa is the best explanation for any perceived 'flickering' pattern."[141] 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.[142] 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.[143]

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.[144][145] Ostrich egg shell containers engraved with geometric designs dating to 60,000 years ago were found at Diepkloof, South Africa.[146] 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,[147] and shell beads dating to about 75,000 years ago have been found at Blombos Cave, South Africa.[148][149][150] 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[151][152][153][154][155] some of which may have been tipped with poisons,[156] and bone harpoons at the Central African site of Katanda dating ca. 90,000 years ago.[157] 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.[158][141]

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.[159][160][161] 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.[162] 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, and date to around 279,000 years ago.[163]

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.[164] 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.[147]

Humans in North Africa (Nazlet Sabaha, Egypt) are known to have dabbled in chert mining, as early as ≈100,000 years ago, for the construction of stone tools.[165][166]

Evidence was found in 2018, dating to about 320,000 years ago at the site of 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.[167][168][169]

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.[170]

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.[171]

Notes

  1. ^ a b Based on Schlebusch et al., "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago",[6] Fig. 3 (H. sapiens divergence times) and Stringer (2012),[7] (archaic admixture).
  2. ^ This is a matter of convention (rather than a factual dispute), and there is no universal consensus on terminology. Some scholars include humans of up to 600,000 years ago under the same species. See Bryant (2003), p. 811.[11] See also Tattersall (2012), Page 82 (cf. Unfortunately this consensus in principle hardly clarifies matters much in practice. For there is no agreement on what the 'qualities of a man' actually are," [...]).[12]
  3. ^ Werdelin[13] citing Lieberman et al.[14]
  4. ^ The history of claimed or proposed subspecies of H. sapiens is complicated and fraught with controversy. The only widely recognized archaic subspecies[citation needed] is H. sapiens idaltu (2003). The name H. s. sapiens is due to Linnaeus (1758), and refers by definition the subspecies of which Linnaeus himself is the type specimen. However, Linnaeus postulated four other extant subspecies, viz. H. s. afer, H. s. americanus, H. s. asiaticus and H. s. ferus for Africans, Americans, Asians and Malay. This classification remained in common usage until the mid 20th century, sometimes alongside H. s. tasmanianus for Australians. See, for example, Bailey, 1946;[18] Hall, 1946.[19] The division of extant human populations into taxonomic subspecies was gradually given up in the 1970s (for example, Grzimek's Animal Life Encyclopedia[20]).
  5. ^ (95% confidence interval 237–581 kya)[58]
  6. ^ "Although none of the Qesem teeth shows a suite of Neanderthal characters, a few traits may suggest some affinities with members of the Neanderthal evolutionary lineage. However, the balance of the evidence suggests a closer similarity with the Skhul/Qafzeh dental material, although many of these resemblances likely represent plesiomorphous features."[62]
  7. ^ "Currently available genetic and archaeological evidence is generally interpreted as supportive of a recent single origin of modern humans in East Africa."[83]
  8. ^ It is important to note that this is a question of conventional terminology, not one of a factual disagreement. Pääbo (2014) frames this as a debate that is unresolvable in principle, "since there is no definition of species perfectly describing the case."[94]
  9. ^ Contemporary human endocranial volume averages at 1,350 cm3 (82 cu in), with significant differences between populations, global group means range 1,085–1,580 cm3 (66.2–96.4 cu in).[104] Neanderthal average is close to 1,450 cm3 (88 cu in) (male average 1,600 cm3 (98 cu in), female average 1,300 cm3 (79 cu in)), with a range extending up to 1,736 cm3 (105.9 cu in) (Amud 1).[105]
  10. ^ "Based on 45 long bones from maximally 14 males and 7 females, Neanderthals' height averages between 164 and 168 (males) resp. 152 to 156 cm (females). This height is indeed 12–14 cm lower than the height of post-WWII Europeans, but compared to Europeans some 20,000 or 100 years ago, it is practically identical or even slightly higher."[111]
  11. ^ Malay, 20–24 (N= m:749 f:893, Median= m:166 cm (5 ft 5+1⁄2 in) f:155 cm (5 ft 1 in), SD= m:6.46 cm (2+1⁄2 in) f:6.04 cm (2+1⁄2 in))[112]
  12. ^ "Specifically, genes in the LCP [lipid catabolic process] term had the greatest excess of NLS in populations of European descent, with an average NLS frequency of 20.8±2.6% versus 5.9±0.08% genome wide (two-sided t-test, P<0.0001, n=379 Europeans and n=246 Africans). Further, among examined out-of-Africa human populations, the excess of NLS [Neanderthal-like genomic sites] in LCP genes was only observed in individuals of European descent: the average NLS frequency in Asians is 6.7±0.7% in LCP genes versus 6.2±0.06% genome wide."[114]
  13. ^ Traits affected by the mutation are sweat glands, teeth, hair thickness and breast tissue.[116][117]
  14. ^ "We offer an alternative hypothesis that suggests that hominid expansion into regions of cold climate produced change in head shape. Such change in shape contributed to the increased cranial volume. Bioclimatic effects directly upon body size (and indirectly upon brain size) in combination with cranial globularity appear to be a fairly powerful explanation of ethnic group differences." (figure in Beals, p304)[120]

References

  1. ^ Nitecki, Matthew H; Nitecki, Doris V (1994). Origins of Anatomically Modern Humans. Springer. ISBN 1489915079.
  2. ^ a b c Vidal, Celine M.; Lane, Christine S.; Asfawrossen, Asrat; et al. (Jan 2022). "Age of the oldest known Homo sapiens from eastern Africa". Nature. 601 (7894): 579–583. Bibcode:2022Natur.601..579V. doi:10.1038/s41586-021-04275-8. PMC 8791829. PMID 35022610.
  3. ^ Hammond, Ashley S.; Royer, Danielle F.; Fleagle, John G. (Jul 2017). "The Omo-Kibish I pelvis". Journal of Human Evolution. 108: 199–219. doi:10.1016/j.jhevol.2017.04.004. ISSN 1095-8606. PMID 28552208.
  4. ^ a b c d e f Mounier, Aurélien; Lahr, Marta (2019). "Deciphering African late middle Pleistocene hominin diversity and the origin of our species". Nature Communications. 10 (1): 3406. Bibcode:2019NatCo..10.3406M. doi:10.1038/s41467-019-11213-w. PMC 6736881. PMID 31506422.
  5. ^ a b Scerri, Eleanor M. L.; Thomas, Mark G.; Manica, Andrea; Gunz, Philipp; Stock, Jay T.; Stringer, Chris; Grove, Matt; Groucutt, Huw S.; Timmermann, Axel; Rightmire, G. Philip; d'Errico, Francesco (2018-08-01). "Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter?". Trends in Ecology & Evolution. 33 (8): 582–594. doi:10.1016/j.tree.2018.05.005. ISSN 0169-5347. PMC 6092560. PMID 30007846.
  6. ^ a b Schlebusch; et al. (3 November 2017). "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago". Science. 358 (6363): 652–655. Bibcode:2017Sci...358..652S. doi:10.1126/science.aao6266. PMID 28971970.
  7. ^ a b Stringer, C (2012). "What makes a modern human". Nature. 485 (7396): 33–35. Bibcode:2012Natur.485...33S. doi:10.1038/485033a. PMID 22552077. S2CID 4420496.
  8. ^ a b Neubauer, Simon; Hublin, Jean-Jacques; Gunz, Philipp (2018-01-01). "The evolution of modern human brain shape". Science Advances. 4 (1): eaao5961. Bibcode:2018SciA....4.5961N. doi:10.1126/sciadv.aao5961. ISSN 2375-2548. PMC 5783678. PMID 29376123.
  9. ^ Harrod, James. "Harrod (2014) Suppl File Table 1 mtDNA language myth Database rev May 17 2019.doc". Mother Tongue.
  10. ^ Linné, Carl von (1758). Systema naturæ. Regnum animale (10th ed.). Sumptibus Guilielmi Engelmann. pp. 18, 20. Retrieved 2019-05-06.
  11. ^ Bryant, Clifton D (2003). Handbook of Death and Dying. SAGE. ISBN 0761925147.
  12. ^ Tattersall, Ian (2012). Masters of the Planet: The Search for Our Human Origins. St Martin's Press. ISBN 978-1137000385.
  13. ^ Werdelin, Lars; Sanders, William Joseph (2010). Cenozoic Mammals of Africa. Univ of California Press. p. 517. ISBN 978-0520257214.
  14. ^ Lieberman, DE; McBratney, BM; Krovitz, G (2002). "The evolution and development of cranial form in Homo sapiens". PNAS. 99 (3): 1134–1139. Bibcode:2002PNAS...99.1134L. doi:10.1073/pnas.022440799. PMC 122156. PMID 11805284.
  15. ^ Hajdinjak, Mateja; Fu, Qiaomei; Hübner, Alexander; Petr, Martin; et al. (2018-03-01). "Reconstructing the genetic history of late Neanderthals". Nature. 555 (7698): 652–656. Bibcode:2018Natur.555..652H. doi:10.1038/nature26151. ISSN 1476-4687. PMC 6485383. PMID 29562232.
  16. ^ Meyer, Matthias; Arsuaga, Juan-Luis; de Filippo, Cesare; Nagel, Sarah; et al. (2016-03-01). "Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins". Nature. 531 (7595): 504–507. Bibcode:2016Natur.531..504M. doi:10.1038/nature17405. ISSN 1476-4687. PMID 26976447. S2CID 4467094.
  17. ^ Gómez-Robles, Aida (2019-05-01). "Dental evolutionary rates and its implications for the Neanderthal–modern human divergence". Science Advances. 5 (5): 1268. Bibcode:2019SciA....5.1268G. doi:10.1126/sciadv.aaw1268. ISSN 2375-2548. PMC 6520022. PMID 31106274.
  18. ^ Bailey, John Wendell (1946). The Mammals of Virginia. p. 356.
  19. ^ Hall, E (1946). "Zoological Subspecies of Man at the Peace Table". Journal of Mammalogy. 27 (4): 358–364. doi:10.2307/1375342. JSTOR 1375342. PMID 20247535.
  20. ^ Grzimek, Bernhard (1970). Grzimek's Animal Life Encyclopedia. Vol. 11. p. 55. ISBN 978-0442784782.
  21. ^ Hublin, J. J. (2009). "The origin of Neandertals". Proceedings of the National Academy of Sciences. 106 (38): 16022–16027. Bibcode:2009PNAS..10616022H. doi:10.1073/pnas.0904119106. JSTOR 40485013. PMC 2752594. PMID 19805257.
  22. ^ Harvati, K.; Frost, S.R.; McNulty, K.P. (2004). "Neanderthal taxonomy reconsidered: implications of 3D primate models of intra- and interspecific differences". Proc. Natl. Acad. Sci. U.S.A. 101 (5): 1147–1152. Bibcode:2004PNAS..101.1147H. doi:10.1073/pnas.0308085100. PMC 337021. PMID 14745010.
  23. ^ "Homo neanderthalensis King, 1864". Wiley-Blackwell Encyclopedia of Human Evolution. Chichester, West Sussex: Wiley-Blackwell. 2013. pp. 328–331.
  24. ^ a b c d Rafferty, John P. "Homo sapiens sapiens". Encyclopedia Britannica. Retrieved 2020-08-11.
  25. ^ Schlebusch, Carina M.; Malmström, Helena; Günther, Torsten; Sjödin, Per; Coutinho, Alexandra; Edlund, Hanna; Munters, Arielle R.; Steyn, Maryna; Soodyall, Himla; Lombard, Marlize; Jakobsson, Mattias (5 June 2017). "Ancient genomes from southern Africa pushes modern human divergence beyond 260,000 years ago". bioRxiv 10.1101/145409.
  26. ^ Schlebusch, Carina M.; Malmström, Helena; Günther, Torsten; Sjödin, Per; Coutinho, Alexandra; Edlund, Hanna; Munters, Arielle R.; Vicente, Mário; Steyn, Maryna; Soodyall, Himla; Lombard, Marlize (2017-11-03). "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago". Science. 358 (6363): 652–655. Bibcode:2017Sci...358..652S. doi:10.1126/science.aao6266. ISSN 0036-8075. PMID 28971970.
  27. ^ a b c Callaway, Ewan (7 June 2017). "Oldest Homo sapiens fossil claim rewrites our species' history". Nature. doi:10.1038/nature.2017.22114. Retrieved 11 June 2017.
  28. ^ a b c d e Stringer, C. (2016). "The origin and evolution of Homo sapiens". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 371 (1698): 20150237. doi:10.1098/rstb.2015.0237. PMC 4920294. PMID 27298468.
  29. ^ a b Sample, Ian (7 June 2017). "Oldest Homo sapiens bones ever found shake foundations of the human story". The Guardian. Retrieved 7 June 2017.
  30. ^ a b Hublin, Jean-Jacques; Ben-Ncer, Abdelouahed; Bailey, Shara E.; Freidline, Sarah E.; Neubauer, Simon; Skinner, Matthew M.; Bergmann, Inga; Le Cabec, Adeline; Benazzi, Stefano; Harvati, Katerina; Gunz, Philipp (2017). "New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens" (PDF). Nature. 546 (7657): 289–292. Bibcode:2017Natur.546..289H. doi:10.1038/nature22336. PMID 28593953.
  31. ^ a b Scerri, M.L.; et al. (2018). "Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter?". Trends in Ecology & Evolution. 33 (8): 582–594. doi:10.1016/j.tree.2018.05.005. PMC 6092560. PMID 30007846.
  32. ^ Chan, Eva, K. F. (28 October 2019). "Human origins in a southern African palaeo-wetland and first migrations". Nature. 857 (7781): 185–189. Bibcode:2019Natur.575..185C. doi:10.1038/s41586-019-1714-1. PMID 31659339. S2CID 204946938.
  33. ^ Sample, Ian (28 October 2019). "Ancestral home of modern humans is in Botswana, study finds". The Guardian. ISSN 0261-3077. Retrieved 29 October 2019.
  34. ^ Woodward, Aylin (28 October 2019). "New Study Pinpoints The Ancestral Homeland of All Humans Alive Today". ScienceAlert.com. Retrieved 29 October 2019.
  35. ^ a b Yong, Ed (28 October 2019). "Has Humanity's Homeland Been Found?". The Atlantic. Retrieved 28 October 2019.
  36. ^ a b c Zimmer, Carl (10 September 2019). "Scientists Find the Skull of Humanity's Ancestor – on a Computer – By comparing fossils and CT scans, researchers say they have reconstructed the skull of the last common forebear of modern humans". The New York Times. Retrieved 10 September 2019.
  37. ^ Rogers, Alan R.; Bohlender, Ryan J.; Huff, Chad D. (12 September 2017). "Early history of Neanderthals and Denisovans". Proceedings of the National Academy of Sciences. 114 (37): 9859–9863. doi:10.1073/pnas.1706426114. PMC 5604018. PMID 28784789.
  38. ^ Wolpoff, M. H.; Spuhler, J. N.; Smith, F. H.; Radovcic, J.; Pope, G.; Frayer, D. W.; Eckhardt, R.; Clark, G. (1988). "Modern Human Origins". Science. 241 (4867): 772–774. Bibcode:1988Sci...241..772W. doi:10.1126/science.3136545. PMID 3136545. S2CID 5223638.
  39. ^ a b c Green RE, Krause J, Briggs AW, Maricic T, Stenzel U, Kircher M, Patterson N, Li H, Zhai W, Fritz MH, Hansen NF, Durand EY, Malaspinas A, Jensen JD, Marques-Bonet T, Alkan C, Prüfer K, Meyer M, Burbano HA, Good JM, Schultz R, Aximu-Petri A, Butthof A, Höber B, Höffner B, Siegemund M, Weihmann A, Nusbaum C, Lander ES, et al. (May 2010). "A draft sequence of the Neandertal genome". Science. 328 (5979): 710–722. Bibcode:2010Sci...328..710G. doi:10.1126/science.1188021. PMC 5100745. PMID 20448178.
  40. ^ Reich D, Patterson N, Kircher M, Delfin F, Nandineni MR, Pugach I, Ko AM, Ko Y, Jinam TA, Phipps ME, Saitou N, Wollstein A, Kayser M, Pääbo S, Stoneking M (2011). "Denisova admixture and the first modern human dispersals into southeast Asia and oceania". Am J Hum Genet. 89 (4): 516–528. doi:10.1016/j.ajhg.2011.09.005. PMC 3188841. PMID 21944045.
  41. ^ "New Clues Add 40,000 Years to Age of Human Species". www.nsf.gov. NSF – National Science Foundation.
  42. ^ "Age of ancient humans reassessed". BBC News. February 16, 2005. Retrieved April 10, 2010.
  43. ^ "The Oldest Homo Sapiens: Fossils Push Human Emergence Back To 195,000 Years Ago". ScienceDaily. February 28, 2005. Retrieved 2019-05-06.
  44. ^ Alemseged, Z.; Coppens, Y.; Geraads, D. (2002). "Hominid cranium from Homo: Description and taxonomy of Homo-323-1976-896" (PDF). Am J Phys Anthropol. 117 (2): 103–112. doi:10.1002/ajpa.10032. PMID 11815945.
  45. ^ Stoneking, Mark; Soodyall, Himla (1996). "Human evolution and the mitochondrial genome". Current Opinion in Genetics & Development. 6 (6): 731–736. doi:10.1016/S0959-437X(96)80028-1. PMID 8994844.
  46. ^ Human evolution: the fossil evidence in 3D, by Philip L. Walker and Edward H. Hagen, Dept. of Anthropology, University of California, Santa Barbara. Retrieved April 5, 2005.
  47. ^ Meyer, Matthias; Arsuaga, Juan-Luis; de Filippo, Cesare; Nagel, Sarah; Aximu-Petri, Ayinuer; Nickel, Birgit; Martínez, Ignacio; Gracia, Ana; de Castro, José María Bermúdez; Carbonell, Eudald; Viola, Bence; Kelso, Janet; Prüfer, Kay; Pääbo, Svante (14 March 2016). "Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins". Nature. 531 (7595): 504–507. Bibcode:2016Natur.531..504M. doi:10.1038/nature17405. PMID 26976447. S2CID 4467094.
  48. ^ Callaway, Ewen (14 March 2016). "Oldest ancient-human DNA details dawn of Neanderthals". Nature. 531 (7594): 296–286. Bibcode:2016Natur.531..296C. doi:10.1038/531286a. PMID 26983523. S2CID 4459329.
  49. ^ Oppenheimer, S. (2003). Out of Eden: The Peopling of the World. ISBN 978-1841196978.
  50. ^ Trinkaus, E.; Moldovan, O.; Milota, Ș.; Bîlgăr, A.; Sarcina, L.; Athreya, S.; Bailey, S. E.; Rodrigo, R.; et al. (2003). "An early modern human from Peștera cu Oase, Romania". PNAS. 100 (20): 11231–11236. Bibcode:2003PNAS..10011231T. doi:10.1073/pnas.2035108100. PMC 208740. PMID 14504393.
  51. ^ a b Reich, David; Green, Richard E.; Kircher, Martin; Krause, Johannes; Patterson, Nick; Durand, Eric Y.; Viola, Bence; Briggs, Adrian W.; et al. (2010). "Genetic history of an archaic hominin group from Denisova Cave in Siberia". Nature. 468 (7327): 1053–1060. Bibcode:2010Natur.468.1053R. doi:10.1038/nature09710. hdl:10230/25596. PMC 4306417. PMID 21179161.
  52. ^ Trinkaus, Erik (October 2005). "Early modern humans". Annual Review of Anthropology. 34 (1): 207–230. doi:10.1146/annurev.anthro.34.030905.154913. S2CID 9039428.
  53. ^ Meldrum, Jeff; Hilton, Charles E. (2004). From Biped to Strider: The Emergence of Modern Human Walking, Running, and Resource Transport. Springer Science & Business Media. ISBN 978-0306480003.
  54. ^ Vonk, Jennifer; Shackelford, Todd K. (2012). The Oxford Handbook of Comparative Evolutionary Psychology. Oxford University Press, US. pp. 429–. ISBN 978-0199738182.
  55. ^ Bozek, Katarzyna; Wei, Yuning; Yan, Zheng; Liu, Xiling; Xiong, Jieyi; Sugimoto, Masahiro; Tomita, Masaru; Pääbo, Svante; Pieszek, Raik; Sherwood, Chet C.; Hof, Patrick R.; Ely, John J.; Steinhauser, Dirk; Willmitzer, Lothar; Bangsbo, Jens; Hansson, Ola; Call, Josep; Giavalisco, Patrick; Khaitovich, Philipp (2014). "Exceptional Evolutionary Divergence of Human Muscle and Brain Metabolomes Parallels Human Cognitive and Physical Uniqueness". PLOS Biology. 12 (5): e1001871. doi:10.1371/journal.pbio.1001871. PMC 4035273. PMID 24866127.
  56. ^ Thieme, H (2007). "Der große Wurf von Schöningen: Das neue Bild zur Kultur des frühen Menschen". Die Schöninger Speere – Mensch und Jagd vor 400 000 Jahren. Konrad Theiss Verlag. pp. 224–328. ISBN 978-3896460400.
  57. ^ Haidle, M.N. (2006). "Menschenaffen? Affenmenschen? Mensch! Kognition und Sprache im Altpaläolithikum". In Conard, N.J. (ed.). Woher kommt der Mensch. Attempto Verlag. pp. 69–97. ISBN 3893083812.
  58. ^ Mendez, Fernando; Krahn, Thomas; Schrack, Bonnie; Krahn, Astrid-Maria; Veeramah, Krishna; Woerner, August; Fomine, Forka Leypey Mathew; Bradman, Neil; Thomas, Mark (7 March 2013). "An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree" (PDF). American Journal of Human Genetics. 92 (3): 454–459. doi:10.1016/j.ajhg.2013.02.002. PMC 3591855. PMID 23453668.
  59. ^ Krings M, Stone A, Schmitz RW, Krainitzki H, Stoneking M, Pääbo S (July 1997). "Neandertal DNA sequences and the origin of modern humans". Cell. 90 (1): 19–30. doi:10.1016/S0092-8674(00)80310-4. hdl:11858/00-001M-0000-0025-0960-8. PMID 9230299. S2CID 13581775.
  60. ^ Hill, Deborah (16 March 2004). "No Neandertals in the Gene Pool". Science. Retrieved 2019-05-06.
  61. ^ Serre, D; Langaney, A; Chech, M; Teschler-Nicola, M; Paunovic, M; Mennecier, P; Hofreiter, M; Possnert, G; Pääbo, S (2004). "No evidence of Neandertal mtDNA contribution to early modern humans". PLOS Biology. 2 (3): 313–317. doi:10.1371/journal.pbio.0020057. PMC 368159. PMID 15024415.
  62. ^ Hershkovitz, I; Smith, P; Sarig, R; Quam, R; Rodríguez, L; García, R; Arsuaga, JL; Barkai, R; Gopher, A (2011). "Middle pleistocene dental remains from Qesem Cave (Israel)". American Journal of Physical Anthropology. 144 (4): 575–592. doi:10.1002/ajpa.21446. PMID 21404234. S2CID 3106938.
  63. ^ Posth, Cosimo; et al. (4 July 2017). "Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals". Nature Communications. 8: 16046. Bibcode:2017NatCo...816046P. doi:10.1038/ncomms16046. PMC 5500885. PMID 28675384.
  64. ^ White, Tim D.; Asfaw, Berhane; Degusta, David; Gilbert, Henry; Richards, Gary D.; Suwa, Gen; Howell, Clark F. (June 2003). "Pleistocene Homo sapiens from Middle Awash, Ethiopia". Nature. 423 (6941): 742–747. Bibcode:2003Natur.423..742W. doi:10.1038/nature01669. PMID 12802332. S2CID 4432091.
  65. ^ "Fossil Reanalysis Pushes Back Origin of Homo sapiens". Scientific American. 2005-02-17. Retrieved 2019-05-06.
  66. ^ Mehta, Ankita (26 January 2018). "A 177,000-year-old jawbone fossil discovered in Israel is oldest human remains found outside Africa". International Business Times. Retrieved 2019-05-06.
  67. ^ Bae, Christopher J.; Douka, Katerina; Petraglia, Michael D. (8 December 2017). "On the origin of modern humans: Asian perspectives". Science. 358 (6368): eaai9067. doi:10.1126/science.aai9067. PMID 29217544.
  68. ^ Kuo, Lily (10 December 2017). "Early humans migrated out of Africa much earlier than we thought". Quartz. Retrieved 2019-05-06.
  69. ^ Zimmer, Carl (10 July 2019). "A Skull Bone Discovered in Greece May Alter the Story of Human Prehistory – The bone, found in a cave, is the oldest modern human fossil ever discovered in Europe. It hints that humans began leaving Africa far earlier than once thought". The New York Times. Retrieved 11 July 2019.
  70. ^ Staff (10 July 2019). "'Oldest remains' outside Africa reset human migration clock". Phys.org. Retrieved 10 July 2019.
  71. ^ Harvati, Katerina; et al. (10 July 2019). "Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia". Nature. 571 (7766): 500–504. doi:10.1038/s41586-019-1376-z. PMID 31292546. S2CID 195873640.
  72. ^ Rito, T; Richards, MB; Fernandes, V; Alshamali, F; Cerny, V; Pereira, L; Soares, P (2013). "The first modern human dispersals across Africa". PLOS ONE. 8 (11): e80031. Bibcode:2013PLoSO...880031R. doi:10.1371/journal.pone.0080031. PMC 3827445. PMID 24236171.
  73. ^ Henn, Brenna; Gignoux, Christopher R.; Jobin, Matthew (2011). "Hunter-gatherer genomic diversity suggests a southern African origin for modern humans". Proceedings of the National Academy of Sciences of the United States of America. 108 (13): 5154–5162. Bibcode:2011PNAS..108.5154H. doi:10.1073/pnas.1017511108. PMC 3069156. PMID 21383195.
  74. ^ a b c Higham, Thomas F. G.; Wesselingh, Frank P.; Hedges, Robert E. M.; Bergman, Christopher A.; Douka, Katerina (2013-09-11). "Chronology of Ksar Akil (Lebanon) and Implications for the Colonization of Europe by Anatomically Modern Humans". PLOS ONE. 8 (9): e72931. Bibcode:2013PLoSO...872931D. doi:10.1371/journal.pone.0072931. ISSN 1932-6203. PMC 3770606. PMID 24039825.
  75. ^ Posth C, Renaud G, Mittnik M, Drucker DG, Rougier H, Cupillard C, Valentin F, Thevenet C, Furtwängler A, Wißing C, Francken M, Malina M, Bolus M, Lari M, Gigli E, Capecchi G, Crevecoeur I, Beauval C, Flas D, Germonpré M, van der Plicht J, Cottiaux R, Gély B, Ronchitelli A, Wehrberger K, Grigorescu D, Svoboda J, Semal P, Caramelli D, Bocherens H, Harvati K, Conard NJ, Haak W, Powell A, Krause J (2016). "Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe". Current Biology. 26 (6): 827–833. doi:10.1016/j.cub.2016.01.037. hdl:2440/114930. PMID 26853362. S2CID 140098861.
  76. ^ Kamin M, Saag L, Vincente M, et al. (April 2015). "A recent bottleneck of Y chromosome diversity coincides with a global change in culture". Genome Research. 25 (4): 459–466. doi:10.1101/gr.186684.114. PMC 4381518. PMID 25770088.
  77. ^ Vai S, Sarno S, Lari M, Luiselli D, Manzi G, Gallinaro M, Mataich S, Hübner A, Modi A, Pilli E, Tafuri MA, Caramelli D, di Lernia S (March 2019). "Ancestral mitochondrial N lineage from the Neolithic 'green' Sahara". Sci Rep. 9 (1): 3530. Bibcode:2019NatSR...9.3530V. doi:10.1038/s41598-019-39802-1. PMC 6401177. PMID 30837540.
  78. ^ a b Haber M, Jones AL, Connel BA, Asan, Arciero E, Huanming Y, Thomas MG, Xue Y, Tyler-Smith C (June 2019). "A Rare Deep-Rooting D0 African Y-chromosomal Haplogroup and its Implications for the Expansion of Modern Humans Out of Africa". Genetics. 212 (4): 1421–1428. doi:10.1534/genetics.119.302368. PMC 6707464. PMID 31196864.
  79. ^ Clarkson, Chris; Jacobs, Zenobia; Pardoe, Colin (2017). "Human occupation of northern Australia by 65,000 years ago" (PDF). Nature. 547 (7663): 306–310. Bibcode:2017Natur.547..306C. doi:10.1038/nature22968. hdl:2440/107043. PMID 28726833. S2CID 205257212.
  80. ^ St. Fleu, Nicholas (July 19, 2017). "Humans First Arrived in Australia 65,000 Years Ago, Study Suggests". The New York Times.
  81. ^ Wood R (2017-09-02). "Comments on the chronology of Madjedbebe". Australian Archaeology. 83 (3): 172–174. doi:10.1080/03122417.2017.1408545. ISSN 0312-2417. S2CID 148777016.
  82. ^ O'Connell JF, Allen J, Williams MA, Williams AN, Turney CS, Spooner NA, et al. (August 2018). "Homo sapiens first reach Southeast Asia and Sahul?". Proceedings of the National Academy of Sciences of the United States of America. 115 (34): 8482–8490. doi:10.1073/pnas.1808385115. PMC 6112744. PMID 30082377.
  83. ^ Liu, Hua; et al. (2006). "A Geographically Explicit Genetic Model of Worldwide Human-Settlement History". The American Journal of Human Genetics. 79 (2): 230–237. doi:10.1086/505436. PMC 1559480. PMID 16826514.
  84. ^ "Out of Africa Revisited". Science. 308 (5724): 921g. 2005-05-13. doi:10.1126/science.308.5724.921g. S2CID 220100436.
  85. ^ Sankararaman, Sriram; Mallick, Swapan; Patterson, Nick; Reich, David (2016). "The Combined Landscape of Denisovan and Neanderthal Ancestry in Present-Day Humans". Current Biology. 26 (9): 1241–1247. doi:10.1016/j.cub.2016.03.037. ISSN 0960-9822. PMC 4864120. PMID 27032491.
  86. ^ Sánchez-Quinto, F; Botigué, LR; Civit, S; Arenas, C; Avila-Arcos, MC; Bustamante, CD; Comas, D; Lalueza-Fox, C (October 17, 2012). "North African Populations Carry the Signature of Admixture with Neandertals". PLOS ONE. 7 (10): e47765. Bibcode:2012PLoSO...747765S. doi:10.1371/journal.pone.0047765. PMC 3474783. PMID 23082212.
  87. ^ Fu, Q; Li, H; Moorjani, P; Jay, F; Slepchenko, SM; Bondarev, AA; Johnson, PL; Aximu-Petri, A; Prüfer, K; de Filippo, C; Meyer, M; Zwyns, N; Salazar-García, DC; Kuzmin, YV; Keates, SG; Kosintsev, PA; Razhev, DI; Richards, MP; Peristov, NV; Lachmann, M; Douka, K; Higham, TF; Slatkin, M; Hublin, JJ; Reich, D; Kelso, J; Viola, TB; Pääbo, S (October 23, 2014). "Genome sequence of a 45,000-year-old modern human from western Siberia". Nature. 514 (7523): 445–449. Bibcode:2014Natur.514..445F. doi:10.1038/nature13810. PMC 4753769. PMID 25341783.
  88. ^ Brahic, Catherine (February 3, 2014). "Humanity's forgotten return to Africa revealed in DNA". The New Scientist. Retrieved 2019-05-06.
  89. ^ Kuhlwilm, Martin (17 February 2016). "Ancient gene flow from early modern humans into Eastern Neanderthals". Nature. 530 (7591): 429–433. Bibcode:2016Natur.530..429K. doi:10.1038/nature16544. PMC 4933530. PMID 26886800.
  90. ^ Ding, Q.; Hu, Y.; Xu, S.; Wang, J.; Jin, L. (2014) [Online 2013]. "Neanderthal Introgression at Chromosome 3p21.31 was Under Positive Natural Selection in East Asians". Molecular Biology and Evolution. 31 (3): 683–695. doi:10.1093/molbev/mst260. PMID 24336922.
  91. ^ Vernot, B.; Akey, J. M. (2014). "Resurrecting Surviving Neandertal Lineages from Modern Human Genomes". Science. 343 (6174): 1017–1021. Bibcode:2014Sci...343.1017V. doi:10.1126/science.1245938. PMID 24476670. S2CID 23003860.
  92. ^ Ayala, Francisco José; Conde, Camilo José Cela (2017). Processes in Human Evolution: The Journey from Early Hominins to Neanderthals and Modern Humans. ISBN 978-0198739906.
  93. ^ Schopf, J. William (1992). Major Events in the History of Life. Jones & Bartlett Learning. pp. 168–. ISBN 978-0867202687.
  94. ^ Pääbo, Svante (2014). Neanderthal Man: In Search of Lost Genomes. New York: Basic Books. p. 237.
  95. ^ Sanders, Robert (11 June 2003). "160,000-year-old fossilized skulls uncovered in Ethiopia are oldest anatomically modern humans". UC Berkeley News. Retrieved 2019-05-07.
  96. ^ White, Tim D.; Asfaw, B.; DeGusta, D.; Gilbert, H.; Richards, G. D.; Suwa, G.; Howell, F. C. (2003). "Pleistocene Homo sapiens from Middle Awash, Ethiopia". Nature. 423 (6491): 742–747. Bibcode:2003Natur.423..742W. doi:10.1038/nature01669. PMID 12802332. S2CID 4432091.
  97. ^ Stringer, Chris (June 12, 2003). "Human evolution: Out of Ethiopia". Nature. 423 (6941): 693–695. Bibcode:2003Natur.423..692S. doi:10.1038/423692a. PMID 12802315. S2CID 26693109.
  98. ^ Hawks, J.; Wang, E. T.; Cochran, G. M.; Harpending, H. C.; Moyzis, R. K. (2007). "Recent acceleration of human adaptive evolution". Proceedings of the National Academy of Sciences. 104 (52): 20753–20758. Bibcode:2007PNAS..10420753H. doi:10.1073/pnas.0707650104. PMC 2410101. PMID 18087044.
  99. ^ Bhupendra, P. (April 2019). "Forehead Anatomy". Medscape references. Retrieved 2019-05-06.
  100. ^ "How to ID a modern human?". News, 2012. Natural History Museum, London. Retrieved 11 December 2013.
  101. ^ . Encarta. Archived from the original on 29 October 2009.
  102. ^ Morris, Desmond (2007). "The Brow". The Naked Woman: A Study of the Female Body. ISBN 978-0312338534.
  103. ^ Pearce, Eiluned; Stringer, Chris; Dunbar, R. I. M. (2013-05-07). "New insights into differences in brain organization between Neanderthals and anatomically modern humans". Proceedings of the Royal Society of London B: Biological Sciences. 280 (1758): 20130168. doi:10.1098/rspb.2013.0168. ISSN 0962-8452. PMC 3619466. PMID 23486442.
  104. ^ Smith, C. L.; Beals, K. L. (1990). "Cultural correlates with cranial capacity". American Anthropologist. 92: 193–200. doi:10.1525/aa.1990.92.1.02a00150. S2CID 162406199.
  105. ^ Stringer, C (1984). "Human evolution and biological adaptation in the Pleistocene". In Foley, R (ed.). Hominid evolution and community ecology. New York: Academic Press. ISBN 978-0122619205.
  106. ^ Townsend G, Richards L, Hughes T (May 2003). "Molar intercuspal dimensions: genetic input to phenotypic variation". Journal of Dental Research. 82 (5): 350–355. doi:10.1177/154405910308200505. PMID 12709500. S2CID 26123427.
  107. ^ Keith A (1913). "Problems relating to the Teeth of the Earlier Forms of Prehistoric Man". Proceedings of the Royal Society of Medicine. 6 (Odontol Sect): 103–124. doi:10.1177/003591571300601018. PMC 2005996. PMID 19977113.
  108. ^ Tattersall, Jeffrey H; Schwartz, Ian (2003). The human fossil record Craniodental Morphology of Genus Homo (Africa and Asia) (vol 2). Wiley-Liss. pp. 327–328. ISBN 978-0471319283.
  109. ^ Steegmann, A. Theodore; Cerny, Frank J.; Holliday, Trenton W. (2002). "Neandertal cold adaptation: Physiological and energetic factors". American Journal of Human Biology. 14 (5): 566–583. doi:10.1002/ajhb.10070. PMID 12203812. S2CID 2437566.
  110. ^ Stock, J.T. (October 2006). "Hunter-gatherer postcranial robusticity relative to patterns of mobility, climatic adaptation, and selection for tissue economy". American Journal of Physical Anthropology. 131 (2): 194–204. doi:10.1002/ajpa.20398. PMID 16596600.
  111. ^ Helmuth H (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift für Morphologie und Anthropologie. 82 (1): 1–12. doi:10.1127/zma/82/1998/1. PMID 9850627.
  112. ^ Lim TO, Ding LM, Zaki M, et al. (March 2000). "Distribution of Body Weight, Height and Body Mass Index in a National Sample of Malaysian Adults" (PDF). Med. J. Malaysia. 55 (1): 108–128. PMID 11072496.
  113. ^ Wade, N (2006-03-07). "Still Evolving, Human Genes Tell New Story". The New York Times. Retrieved 2008-07-10.
  114. ^ Khrameeva, E; Bozek, K; He, L; Yan, Z; Jiang, X; Wei, Y; Tang, K; Gelfand, MS; Prüfer, K; Kelso, J; Pääbo, S; Giavalisco, P; Lachmann, M; Khaitovich, P (2014). "Neanderthal ancestry drives evolution of lipid catabolism in contemporary Europeans". Nature Communications. 5 (3584): 3584. Bibcode:2014NatCo...5.3584K. doi:10.1038/ncomms4584. PMC 3988804. PMID 24690587.
  115. ^ Michael Dannemann 1 and Janet Kelso, "The Contribution of Neanderthals to Phenotypic Variation in Modern Humans", The American Journal of Human Genetics 101, 578–589, October 5, 2017.
  116. ^ Kamberov, Yana G (14 February 2013). "Modeling Recent Human Evolution in Mice by Expression of a Selected EDAR Variant". Cell. 152 (4): 691–702. doi:10.1016/j.cell.2013.01.016. PMC 3575602. PMID 23415220.
  117. ^ Wade, Nicholas (14 February 2013). "East Asian Physical Traits Linked to 35,000-Year-Old Mutation". The New York Times. Retrieved 2019-05-06.
  118. ^ a b Beleza, Sandra; Santos, A. M.; McEvoy, B.; Alves, I.; Martinho, C.; Cameron, E.; Shriver, M. D.; Parra, E. J.; Rocha, J. (2012). "The timing of pigmentation lightening in Europeans". Molecular Biology and Evolution. 30 (1): 24–35. doi:10.1093/molbev/mss207. PMC 3525146. PMID 22923467.
  119. ^ Lalueza-Fox; Römpler, H; Caramelli, D; Stäubert, C; Catalano, G; Hughes, D; Rohland, N; Pilli, E; Longo, L; Condemi, S; de la Rasilla, M; Fortea, J; Rosas, A; Stoneking, M; Schöneberg, T; Bertranpetit, J; Hofreiter, M; et al. (2007). "A melanocortin-1 receptor allele suggests varying pigmentation among Neanderthals". Science. 318 (5855): 1453–1455. Bibcode:2007Sci...318.1453L. doi:10.1126/science.1147417. PMID 17962522. S2CID 10087710.
  120. ^ a b Beals, Kenneth L; Smith, Courtland L; Dodd, Stephen M (1984). "Brain Size, Cranial Morphology, Climate, and Time Machines". Current Anthropology. 25 (3): 301–330. doi:10.1086/203138. S2CID 86147507.
  121. ^ Nowaczewska, Wioletta; Dabrowski, Pawel; Kuźmiński, Lukasz (2011). . Collegium Antropologicum. 35 (3): 625–636. PMID 22053534. Archived from the original on 2014-10-11.
  122. ^ Peng, Y.; et al. (2010). "The ADH1B Arg47His polymorphism in East Asian populations and expansion of rice domestication in history". BMC Evolutionary Biology. 10: 15. doi:10.1186/1471-2148-10-15. PMC 2823730. PMID 20089146.
  123. ^ Ségurel, Laure; Bon, Céline (2017). "On the Evolution of Lactase Persistence in Humans". Annual Review of Genomics and Human Genetics. 18 (1): 297–319. doi:10.1146/annurev-genom-091416-035340. PMID 28426286.
  124. ^ Ingram, Catherine J. E.; Mulcare, Charlotte A.; Itan, Yuval; Thomas, Mark G.; Swallow, Dallas M. (2008-11-26). "Lactose digestion and the evolutionary genetics of lactase persistence". Human Genetics. 124 (6): 579–591. doi:10.1007/s00439-008-0593-6. ISSN 0340-6717. PMID 19034520. S2CID 3329285.
  125. ^ Ilardo, M. A.; Moltke, I.; Korneliussen, T. S.; Cheng, J.; Stern, A. J.; Racimo, F.; de Barros Damgaard, P.; Sikora, M.; Seguin-Orlando, A.; Rasmussen, S.; van den Munckhof, I. C. L.; ter Horst, R.; Joosten, L. A. B.; Netea, M. G.; Salingkat, S.; Nielsen, R.; Willerslev, E. (2018-04-18). "Physiological and Genetic Adaptations to Diving in Sea Nomads". Cell. 173 (3): 569–580.e15. doi:10.1016/j.cell.2018.03.054. PMID 29677510.
  126. ^ Gislén, A; Dacke, M; Kröger, RH; Abrahamsson, M; Nilsson, DE; Warrant, EJ (2003). "Superior Underwater Vision in a Human Population of Sea Gypsies". Current Biology. 13 (10): 833–836. doi:10.1016/S0960-9822(03)00290-2. PMID 12747831. S2CID 18731746.
  127. ^ Klein, Richard (1995). "Anatomy, behavior, and modern human origins". Journal of World Prehistory. 9 (2): 167–198. doi:10.1007/bf02221838. S2CID 10402296.
  128. ^ Feierman, Jay R. (2009). The Biology of Religious Behavior: The Evolutionary Origins of Faith and Religion. ABC-CLIO. p. 220. ISBN 978-0313364303.
  129. ^ Soressi M. (2005) Late Mousterian lithic technology. Its implications for the pace of the emergence of behavioural modernity and the relationship between behavioural modernity and biological modernity, pp. 389–417 in L. Backwell et F. d'Errico (eds.) From Tools to Symbols, Johannesburg: University of Witswatersand Press. ISBN 1868144178.
  130. ^ Companion encyclopedia of archaeology (1999). Routledge. ISBN 0415213304. Vol. 2. p. 763 (cf., ... "effectively limited to organic samples" [ed. organic compounds ] "or biogenic carbonates that date to less than 50 ka (50,000 years ago)."). See also: Later Stone Age and Upper Paleolithic.
  131. ^ Mellars, Paul (2006). "Why did modern human populations disperse from Africa ca. 60,000 years ago?". Proceedings of the National Academy of Sciences. 103 (25): 9381–9386. Bibcode:2006PNAS..103.9381M. doi:10.1073/pnas.0510792103. PMC 1480416. PMID 16772383.
  132. ^ Shea, John (2011). "Homo sapiens Is As Homo sapiens Was". Current Anthropology. 52 (1): 1–35. doi:10.1086/658067. S2CID 142517998.
  133. ^ McBrearty, Sally; Brooks, Allison (2000). "The revolution that wasn't: a new interpretation of the origin of modern human behavior". Journal of Human Evolution. 39 (5): 453–563. doi:10.1006/jhev.2000.0435. PMID 11102266. S2CID 42968840.
  134. ^ Henshilwood, Christopher; Marean, Curtis (2003). "The Origin of Modern Human Behavior: Critique of the Models and Their Test Implications". Current Anthropology. 44 (5): 627–651. doi:10.1086/377665. PMID 14971366. S2CID 11081605.
  135. ^ Marean, Curtis; et al. (2007). "Early human use of marine resources and pigment in South Africa during the Middle Pleistocene" (PDF). Nature. 449 (7164): 905–908. Bibcode:2007Natur.449..905M. doi:10.1038/nature06204. PMID 17943129. S2CID 4387442.
  136. ^ Powell, Adam; et al. (2009). "Late Pleistocene Demography and the Appearance of Modern Human Behavior" (PDF). Science. 324 (5932): 1298–1301. Bibcode:2009Sci...324.1298P. doi:10.1126/science.1170165. PMID 19498164. S2CID 206518315.
  137. ^ Premo, Luke; Kuhn, Steve (2010). "Modeling Effects of Local Extinctions on Culture Change and Diversity in the Paleolithic". PLOS ONE. 5 (12): e15582. Bibcode:2010PLoSO...515582P. doi:10.1371/journal.pone.0015582. PMC 3003693. PMID 21179418.
  138. ^ St. Fleur, Nicholas (12 September 2018). "Oldest Known Drawing by Human Hands Discovered in South African Cave". The New York Times. Retrieved 15 September 2018.
  139. ^ d'Errico, F.; Backwell, L.; Villa, P.; Degano, I.; Lucejko, J. J.; Bamford, M. K.; Higham, T. F. G.; Colombini, M. P.; Beaumont, P. B. (2012). "Early evidence of San material culture represented by organic artifacts from Border Cave, South Africa". Proceedings of the National Academy of Sciences. 109 (33): 13214–13219. Bibcode:2012PNAS..10913214D. doi:10.1073/pnas.1204213109. PMC 3421171. PMID 22847420.
  140. ^ Mourre, V.; Villa, P.; Henshilwood, C. S. (2010). "Early Use of Pressure Flaking on Lithic Artifacts at Blombos Cave, South Africa". Science. 330 (6004): 659–662. Bibcode:2010Sci...330..659M. doi:10.1126/science.1195550. PMID 21030655. S2CID 34833884.
  141. ^ a b Brown, Kyle S.; Marean, Curtis W.; Jacobs, Zenobia; Schoville, Benjamin J.; Oestmo, Simen; Fisher, Erich C.; Bernatchez, Jocelyn; Karkanas, Panagiotis; Matthews, Thalassa (2012). "An early and enduring advanced technology originating 71,000 years ago in South Africa". Nature. 491 (7425): 590–593. Bibcode:2012Natur.491..590B. doi:10.1038/nature11660. PMID 23135405. S2CID 4323569.
  142. ^ Scholz, C. A.; Johnson, T. C.; Cohen, A. S.; King, J. W.; Peck, J. A.; Overpeck, J. T.; Talbot, M. R.; Brown, E. T.; Kalindekafe, L.; Amoako, P. Y. O.; Lyons, R. P.; Shanahan, T. M.; Castaneda, I. S.; Heil, C. W.; Forman, S. L.; McHargue, L. R.; Beuning, K. R.; Gomez, J.; Pierson, J. (2007). "East African megadroughts between 135 and 75 thousand years ago and bearing on early-modern human origins". Proceedings of the National Academy of Sciences. 104 (42): 16416–16421. Bibcode:2007PNAS..10416416S. doi:10.1073/pnas.0703874104. PMC 1964544. PMID 17785420.
  143. ^ Wells, Spencer (2003). The Journey of Man: A Genetic Odyssey. Princeton, N.J. : Princeton University Press. ISBN 978-0691115320.
  144. ^ Henshilwood, Christopher; et al. (2002). "Emergence of Modern Human Behavior: Middle Stone Age Engravings from South Africa". Science. 295 (5558): 1278–1280. Bibcode:2002Sci...295.1278H. doi:10.1126/science.1067575. PMID 11786608. S2CID 31169551.
  145. ^ Henshilwood, Christopher S.; d'Errico, Francesco; Watts, Ian (2009). "Engraved ochres from the Middle Stone Age levels at Blombos Cave, South Africa". Journal of Human Evolution. 57 (1): 27–47. doi:10.1016/j.jhevol.2009.01.005. PMID 19487016.
  146. ^ Texier, PJ; Porraz, G; Parkington, J; Rigaud, JP; Poggenpoel, C; Miller, C; Tribolo, C; Cartwright, C; Coudenneau, A; Klein, R; Steele, T; Verna, C (2010). "A Howiesons Poort tradition of engraving ostrich eggshell containers dated to 60,000 years ago at Diepkloof Rock Shelter, South Africa". Proceedings of the National Academy of Sciences. 107 (14): 6180–6185. Bibcode:2010PNAS..107.6180T. doi:10.1073/pnas.0913047107. PMC 2851956. PMID 20194764.
  147. ^ a b McBrearty, Sally; Brooks, Allison (2000). "The revolution that wasn't: a new interpretation of the origin of modern human behavior". Journal of Human Evolution. 39 (5): 453–563. doi:10.1006/jhev.2000.0435. PMID 11102266. S2CID 42968840.
  148. ^ Henshilwood, Christopher S.; et al. (2004). "Middle Stone Age shell beads from South Africa". Science. 304 (5669): 404. doi:10.1126/science.1095905. PMID 15087540. S2CID 32356688.
  149. ^ d'Errico, Francesco; et al. (2005). "Nassarius kraussianus shell beads from Blombos Cave: evidence for symbolic behaviour in the Middle Stone Age". Journal of Human Evolution. 48 (1): 3–24. doi:10.1016/j.jhevol.2004.09.002. PMID 15656934.
  150. ^ Vanhaeren, Marian; et al. (2013). "Thinking strings: Additional evidence for personal ornament use in the Middle Stone Age at Blombos Cave, South Africa". Journal of Human Evolution. 64 (6): 500–517. doi:10.1016/j.jhevol.2013.02.001. PMID 23498114.
  151. ^ Backwell, L; d'Errico, F; Wadley, L (2008). "Middle Stone Age bone tools from the Howiesons Poort layers, Sibudu Cave, South Africa". Journal of Archaeological Science. 35 (6): 1566–1580. doi:10.1016/j.jas.2007.11.006.
  152. ^ Wadley, Lyn (2008). "The Howieson's Poort industry of Sibudu Cave". South African Archaeological Society Goodwin Series. 10.
  153. ^ Lombard M, Phillips L (2010). "Indications of bow and stone-tipped arrow use 64,000 years ago in KwaZulu-Natal, South Africa". Antiquity. 84 (325): 635–648. doi:10.1017/S0003598X00100134. S2CID 162438490.
  154. ^ Lombard M (2011). "Quartz-tipped arrows older than 60 ka: further use-trace evidence from Sibudu, Kwa-Zulu-Natal, South Africa". Journal of Archaeological Science. 38 (8): 1918–1930. doi:10.1016/j.jas.2011.04.001.
  155. ^ Backwell, L; Bradfield, J; Carlson, KJ; Jashashvili, T; Wadley, L; d'Errico, F (2018). "The antiquity of bow-and-arrow technology: evidence from Middle Stone Age layers at Sibudu Cave". Journal of Archaeological Science. 92 (362): 289–303. doi:10.15184/aqy.2018.11.
  156. ^ Lombard M (2020). "The tip cross-sectional areas of poisoned bone arrowheads from southern Africa". Journal of Archaeological Science: Reports. 33: 102477. doi:10.1016/j.jasrep.2020.102477. S2CID 224889105.
  157. ^ Yellen, JE; AS Brooks; E Cornelissen; MJ Mehlman; K Stewart (28 April 1995). "A middle stone age worked bone industry from Katanda, Upper Semliki Valley, Zaire". Science. 268 (5210): 553–556. Bibcode:1995Sci...268..553Y. doi:10.1126/science.7725100. PMID 7725100.
  158. ^ Brown, Kyle S.; Marean, Curtis W.; Herries, Andy I.R.; Jacobs, Zenobia; Tribolo, Chantal; Braun, David; Roberts, David L.; Meyer, Michael C.; Bernatchez, J. (14 August 2009), "Fire as an Engineering Tool of Early Modern Humans", Science, 325 (5942): 859–862, Bibcode:2009Sci...325..859B, doi:10.1126/science.1175028, hdl:11422/11102, PMID 19679810, S2CID 43916405
  159. ^ Amos, Jonathan (13 October 2011). "A Cultural Leap at the Dawn of Humanity – Ancient 'paint factory' unearthed". BBC News. Retrieved 13 October 2011.
  160. ^ Vastag, Brian (13 October 2011). "South African cave yields paint from dawn of humanity". The Washington Post. Retrieved 13 October 2011.
  161. ^ Henshilwood, Christopher S.; et al. (2011). "A 100,000-Year-Old Ochre-Processing Workshop at Blombos Cave, South Africa". Science. 334 (6053): 219–222. Bibcode:2011Sci...334..219H. doi:10.1126/science.1211535. PMID 21998386. S2CID 40455940.
  162. ^ Shipton C, d'Errico F, Petraglia M, et al. (2018). 78,000-year-old record of Middle and Later Stone Age innovation in an East African tropical forest. Nature Communications
  163. ^ Sahle, Y.; Hutchings, W. K.; Braun, D. R.; Sealy, J. C.; Morgan, L. E.; Negash, A.; Atnafu, B. (2013). Petraglia, Michael D (ed.). "Earliest Stone-Tipped Projectiles from the Ethiopian Rift Date to >279,000 Years Ago". PLOS ONE. 8 (11): e78092. Bibcode:2013PLoSO...878092S. doi:10.1371/journal.pone.0078092. PMC 3827237. PMID 24236011.
  164. ^ Marean, Curtis; et al. (2007). "Early human use of marine resources and pigment in South Africa during the Middle Pleistocene" (PDF). Nature. 449 (7164): 905–908. Bibcode:2007Natur.449..905M. doi:10.1038/nature06204. PMID 17943129. S2CID 4387442.
  165. ^ . Archived from the original on 2019-01-05. Retrieved 2019-09-11.
  166. ^ Guinness World Records (10 September 2015). Guinness World Records 2016. Guinness World Records. p. 27. ISBN 978-1910561034.
  167. ^ Chatterjee, Rhitu (15 March 2018). "Scientists Are Amazed By Stone Age Tools They Dug Up In Kenya". NPR. Retrieved 15 March 2018.
  168. ^ Yong, Ed (15 March 2018). "A Cultural Leap at the Dawn of Humanity – New finds from Kenya suggest that humans used long-distance trade networks, sophisticated tools, and symbolic pigments right from the dawn of our species". The Atlantic. Retrieved 15 March 2018.
  169. ^ Brooks AS, Yellen JE, Potts R, Behrensmeyer AK, Deino AL, Leslie DE, Ambrose SH, Ferguson JR, d'Errico F, Zipkin AM, Whittaker S, Post J, Veatch EG, Foecke K, Clark JB (2018). "Long-distance stone transport and pigment use in the earliest Middle Stone Age". Science. 360 (6384): 90–94. Bibcode:2018Sci...360...90B. doi:10.1126/science.aao2646. PMID 29545508.
  170. ^ Sahle Y, Brooks AS (2018). "Assessment of complex projectiles in the early Late Pleistocene at Aduma, Ethiopia". PLOS ONE. 14 (5): e0216716. Bibcode:2019PLoSO..1416716S. doi:10.1371/journal.pone.0216716. PMC 6508696. PMID 31071181.
  171. ^ Douglas, Kate (March 24, 2012). . New Scientist. Archived from the original on April 13, 2016. Retrieved May 4, 2022.{{cite magazine}}: CS1 maint: unfit URL (link)

Further reading

External links

  1. ^ Diamond, Jared (April 20, 2018). "A Brand-New Version of Our Origin Story". The New York Times. Retrieved April 23, 2018.

January 31, 2023
early, modern, human, this, article, about, first, homo, sapiens, specifically, during, stone, broader, perspective, human, species, human, anatomically, modern, human, terms, used, distinguish, homo, sapiens, only, extant, hominina, species, that, anatomicall. This article is about the first Homo sapiens specifically during the Old Stone Age For a broader perspective on the human species see Human Early modern human EMH or anatomically modern human AMH 1 are terms used to distinguish Homo sapiens 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 archaeological site in south western Ethiopia dating to about 233 000 2 to 196 000 years ago 3 the Florisbad site in South Africa dating to about 259 000 years ago and the Jebel Irhoud site in Morocco dated about 315 000 years ago 100 to 80 thousand year old Skhul V from Israel Extinct species of the genus Homo include Homo erectus 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 around 300 000 years ago with the earliest genetic splits among modern people according to some evidence dating to around the same time 4 5 note 1 8 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 9 Contents 1 Name and taxonomy 2 Age and speciation process 2 1 Derivation from H erectus 2 2 Early Homo sapiens 3 Dispersal and archaic admixture 4 Anatomy 4 1 Anatomical modernity 4 2 Braincase anatomy 4 3 Jaw anatomy 4 4 Body skeleton structure 5 Recent evolution 6 Behavioral modernity 6 1 Examples of behavioural modernity 7 Pace of progress during Homo sapiens history 8 Notes 9 References 10 Further reading 11 External linksName and taxonomyMain article Human taxonomy Further information Homo and Names for the human species The binomial name Homo sapiens was coined by Linnaeus 1758 10 The Latin noun homō genitive hominis means human being while the participle sapiens 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 Estimates for the split between the Homo sapiens line and combined Neanderthal Denisovan line range from between 503 000 and 565 000 years ago 15 between 550 000 and 765 000 years ago 16 and based on rates of dental evolution possibly more than 800 000 years ago 17 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 21 22 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 23 All humans are considered to be a part of the subspecies H sapiens sapiens 24 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 24 Age and speciation processFurther information Human evolution Homo Multiregional origin of modern humans Timeline of human evolution and Early human migrations Derivation from H erectus Further information Homo ergaster Homo antecessor Homo heidelbergensis Homo rhodesiensis and Acheulean 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 Homo at a given time and place late survival of robust australopithecines alongside Homo is indicated in purple Based on Springer 2012 Homo heidelbergensis 7 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 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 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 8 5 But the oldest split among modern human populations such as the Khoisan split from other groups has been recently dated to between 350 000 and 260 000 years ago 25 26 and the earliest known examples of H sapiens fossils also date to about that period including the Jebel Irhoud remains from Morocco ca 300 000 or 350 280 000 years ago 27 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 28 29 30 31 2 An mtDNA study in 2019 proposed an origin of modern humans in Botswana and a Khoisan split of around 200 000 years 32 However this proposal has been widely criticized by scholars 33 34 35 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 35 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 36 4 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 37 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 selection throughout the Pleistocene 38 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 39 40 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 41 42 43 44 45 H s idaltu dated to 160 000 years ago has been postulated as an extinct subspecies of H sapiens in 2003 46 24 H neanderthalensis which became extinct about 40 000 years ago was also at one point considered to be a subspecies H s neanderthalensis 24 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 47 48 Early Homo sapiens Further information Human subspecies Middle Paleolithic Mousterian Interbreeding between archaic and modern humans Homo sapiens idaltu and Skhul and Qafzeh hominins 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 Omo Herto Florisbad Skhul and Peștera cu Oase exhibit a mix of archaic and modern traits 49 50 27 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 51 52 The gracile or lightly built skeleton of anatomically modern humans has been connected to a change in behavior including increased cooperation and resource transport 53 54 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 55 The Schoningen 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 56 57 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 5 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 59 60 61 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 6 Dispersal and archaic admixtureFurther 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 around 315 000 years ago 27 30 There is indirect evidence for H sapiens presence in West Asia around 270 000 years ago 63 The Florisbad Skull from Florisbad South Africa dated to about 259 000 years ago has also been classified as representing early H sapiens 28 29 31 4 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 36 4 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 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 6 H s idaltu found at Middle Awash in Ethiopia lived about 160 000 years ago 64 and H sapiens lived at Omo Kibish in Ethiopia about 233 000 195 000 years ago 65 2 Two fossils from Guomde Kenya dated to at least and likely more than 180 000 years ago 28 and more precisely to 300 270 000 years ago 4 have been tentatively assigned to H sapiens and similarities have been noted between them and the Omo Kibbish remains 28 Fossil evidence for modern human presence in West Asia is ascertained for 177 000 years ago 66 and disputed fossil evidence suggests expansion as far as East Asia by 120 000 years ago 67 68 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 69 70 71 A significant dispersal event within Africa and to West Asia is associated with the African megadroughts during MIS 5 beginning 130 000 years ago 72 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 73 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 74 and 42 400 41 700 BP for Ethelruda 74 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 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 75 76 77 78 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 between 65 000 and 50 000 years ago 79 80 81 82 while Europe East and North Asia 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 citation needed 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 7 84 and has also been supported by Y DNA and autosomal DNA 78 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 85 39 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 86 87 88 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 89 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 90 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 addition to neanderthal admixture 39 51 Cumulatively about 20 of the Neanderthal genome is estimated to remain present spread in contemporary populations 91 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 while North African fossils may represent a population which introgressed into Neandertals during the LMP 36 4 AnatomySee 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 74 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 92 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 93 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 8 In this more narrow definition of H sapiens the subspecies Homo sapiens idaltu discovered in 2003 also falls under the umbrella of anatomically modern 95 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 96 However biological anthropologist Chris Stringer does not consider idaltu distinct enough within H sapiens to warrant its own subspecies designation 97 28 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 98 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 99 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 100 Modern humans commonly have a steep even vertical forehead whereas their predecessors had foreheads that sloped strongly backwards 101 According to Desmond Morris the vertical forehead in humans plays an important role in human communication through eyebrow movements and forehead skin wrinkling 102 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 103 note 9 Jaw anatomy Compared to archaic people anatomically modern humans have smaller differently shaped teeth 106 107 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 108 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 109 The same adaptation is found in some modern people living in the polar regions 110 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 which is largely identical to pre industrial average heights for AMH note 10 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 contemporary height distribution measured among Malay people for one note 11 Recent evolutionMain article Recent human evolution Further information Human genetic variation 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 had 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 113 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 12 A 2017 study found correlation of Neanderthal admixture in phenotypic traits in modern European populations 115 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 13 Recent divergence of Eurasian lineages was sped up significantly during the Last Glacial Maximum LGM the Mesolithic and the Neolithic due to increased selection pressures and due to founder effects associated with migration 118 Alleles predictive of light skin have been found in Neanderthals 119 but the alleles for light skin in Europeans and East Asians associated with KITLG and ASIP are as of 2012 update thought to have not been acquired by archaic admixture but recent mutations since the LGM 118 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 for adult males Larger cranial volume is associated with climatic region the largest averages being found in populations of Siberia and the Arctic note 14 121 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 120 Examples for still later adaptations related to agriculture and animal domestication including East Asian types of ADH1B associated with rice domestication 122 or lactase persistence 123 124 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 125 126 Behavioral modernity Lithic Industries of early Homo sapiens at Blombos Cave M3 phase MIS 5 Southern Cape South Africa c 105 000 90 000 years old 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 127 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 128 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 129 while others use the term for the rapid developments occurring around 50 000 years ago 130 131 132 It has been proposed that the emergence of behavioral modernity was a gradual process 133 134 135 136 137 Examples of behavioural modernity Claimed oldest known drawing by human hands discovered in Blombos Cave in South Africa Estimated to be a 73 000 year old work of a Homo sapiens 138 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 at 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 139 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 140 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 were early and enduring a small sample of excavated sites in Africa is the best explanation for any perceived flickering pattern 141 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 142 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 143 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 144 145 Ostrich egg shell containers engraved with geometric designs dating to 60 000 years ago were found at Diepkloof South Africa 146 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 147 and shell beads dating to about 75 000 years ago have been found at Blombos Cave South Africa 148 149 150 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 151 152 153 154 155 some of which may have been tipped with poisons 156 and bone harpoons at the Central African site of Katanda dating ca 90 000 years ago 157 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 158 141 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 159 160 161 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 162 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 and date to around 279 000 years ago 163 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 164 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 147 Humans in North Africa Nazlet Sabaha Egypt are known to have dabbled in chert mining as early as 100 000 years ago for the construction of stone tools 165 166 Evidence was found in 2018 dating to about 320 000 years ago at the site of 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 167 168 169 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 170 Pace of progress during Homo sapiens historyHomo 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 171 Notes a b Based on Schlebusch et al Southern African ancient genomes estimate modern human divergence to 350 000 to 260 000 years ago 6 Fig 3 H sapiens divergence times and Stringer 2012 7 archaic admixture This is a matter of convention rather than a factual dispute and there is no universal consensus on terminology Some scholars include humans of up to 600 000 years ago under the same species See Bryant 2003 p 811 11 See also Tattersall 2012 Page 82 cf Unfortunately this consensus in principle hardly clarifies matters much in practice For there is no agreement on what the qualities of a man actually are 12 Werdelin 13 citing Lieberman et al 14 The history of claimed or proposed subspecies of H sapiens is complicated and fraught with controversy The only widely recognized archaic subspecies citation needed is H sapiens idaltu 2003 The name H s sapiens is due to Linnaeus 1758 and refers by definition the subspecies of which Linnaeus himself is the type specimen However Linnaeus postulated four other extant subspecies viz H s afer H s americanus H s asiaticus and H s ferus for Africans Americans Asians and Malay This classification remained in common usage until the mid 20th century sometimes alongside H s tasmanianus for Australians See for example Bailey 1946 18 Hall 1946 19 The division of extant human populations into taxonomic subspecies was gradually given up in the 1970s for example Grzimek s Animal Life Encyclopedia 20 95 confidence interval 237 581 kya 58 Although none of the Qesem teeth shows a suite of Neanderthal characters a few traits may suggest some affinities with members of the Neanderthal evolutionary lineage However the balance of the evidence suggests a closer similarity with the Skhul Qafzeh dental material although many of these resemblances likely represent plesiomorphous features 62 Currently available genetic and archaeological evidence is generally interpreted as supportive of a recent single origin of modern humans in East Africa 83 It is important to note that this is a question of conventional terminology not one of a factual disagreement Paabo 2014 frames this as a debate that is unresolvable in principle since there is no definition of species perfectly describing the case 94 Contemporary human endocranial volume averages at 1 350 cm3 82 cu in with significant differences between populations global group means range 1 085 1 580 cm3 66 2 96 4 cu in 104 Neanderthal average is close to 1 450 cm3 88 cu in male average 1 600 cm3 98 cu in female average 1 300 cm3 79 cu in with a range extending up to 1 736 cm3 105 9 cu in Amud 1 105 Based on 45 long bones from maximally 14 males and 7 females Neanderthals height averages between 164 and 168 males resp 152 to 156 cm females This height is indeed 12 14 cm lower than the height of post WWII Europeans but compared to Europeans some 20 000 or 100 years ago it is practically identical or even slightly higher 111 Malay 20 24 N m 749 f 893 Median m 166 cm 5 ft 5 1 2 in f 155 cm 5 ft 1 in SD m 6 46 cm 2 1 2 in f 6 04 cm 2 1 2 in 112 Specifically genes in the LCP lipid catabolic process term had the greatest excess of NLS in populations of European descent with an average NLS frequency of 20 8 2 6 versus 5 9 0 08 genome wide two sided t test P lt 0 0001 n 379 Europeans and n 246 Africans Further among examined out of Africa human populations the excess of NLS Neanderthal like genomic sites in LCP genes was only observed in individuals of European descent the average NLS frequency in Asians is 6 7 0 7 in LCP genes versus 6 2 0 06 genome wide 114 Traits affected by the mutation are sweat glands teeth hair thickness and breast tissue 116 117 We offer an alternative hypothesis that suggests that hominid expansion into regions of cold climate produced change in head shape Such change in shape contributed to the increased cranial volume Bioclimatic effects directly upon body size and indirectly upon brain size in combination with cranial globularity appear to be a fairly powerful explanation of ethnic group differences figure in Beals p304 120 References Nitecki Matthew H Nitecki Doris V 1994 Origins of Anatomically Modern Humans Springer ISBN 1489915079 a b c Vidal Celine M Lane Christine S Asfawrossen Asrat et al Jan 2022 Age of the oldest known Homo sapiens from eastern Africa Nature 601 7894 579 583 Bibcode 2022Natur 601 579V doi 10 1038 s41586 021 04275 8 PMC 8791829 PMID 35022610 Hammond Ashley S Royer Danielle F Fleagle John G Jul 2017 The Omo Kibish I pelvis Journal of Human Evolution 108 199 219 doi 10 1016 j jhevol 2017 04 004 ISSN 1095 8606 PMID 28552208 a b c d e f Mounier Aurelien Lahr Marta 2019 Deciphering African late middle Pleistocene hominin diversity and the origin of our species Nature Communications 10 1 3406 Bibcode 2019NatCo 10 3406M doi 10 1038 s41467 019 11213 w PMC 6736881 PMID 31506422 a b Scerri Eleanor M L Thomas Mark G Manica Andrea Gunz Philipp Stock Jay T Stringer Chris Grove Matt Groucutt Huw S Timmermann Axel Rightmire G Philip d Errico Francesco 2018 08 01 Did Our Species Evolve in Subdivided Populations across Africa and Why Does It Matter Trends in Ecology amp Evolution 33 8 582 594 doi 10 1016 j tree 2018 05 005 ISSN 0169 5347 PMC 6092560 PMID 30007846 a b Schlebusch et al 3 November 2017 Southern African ancient genomes estimate modern human divergence to 350 000 to 260 000 years ago Science 358 6363 652 655 Bibcode 2017Sci 358 652S doi 10 1126 science aao6266 PMID 28971970 a b Stringer C 2012 What makes a modern human Nature 485 7396 33 35 Bibcode 2012Natur 485 33S doi 10 1038 485033a PMID 22552077 S2CID 4420496 a b Neubauer Simon Hublin Jean Jacques Gunz Philipp 2018 01 01 The evolution of modern human brain shape Science Advances 4 1 eaao5961 Bibcode 2018SciA 4 5961N doi 10 1126 sciadv aao5961 ISSN 2375 2548 PMC 5783678 PMID 29376123 Harrod James Harrod 2014 Suppl File Table 1 mtDNA language myth Database rev May 17 2019 doc Mother Tongue Linne Carl von 1758 Systema naturae Regnum animale 10th ed Sumptibus Guilielmi Engelmann pp 18 20 Retrieved 2019 05 06 Bryant Clifton D 2003 Handbook of Death and Dying SAGE ISBN 0761925147 Tattersall Ian 2012 Masters of the Planet The Search for Our Human Origins St Martin s Press ISBN 978 1137000385 Werdelin Lars Sanders William Joseph 2010 Cenozoic Mammals of Africa Univ of California Press p 517 ISBN 978 0520257214 Lieberman DE McBratney BM Krovitz G 2002 The evolution and development of cranial form in Homo sapiens PNAS 99 3 1134 1139 Bibcode 2002PNAS 99 1134L doi 10 1073 pnas 022440799 PMC 122156 PMID 11805284 Hajdinjak Mateja Fu Qiaomei Hubner Alexander Petr Martin et al 2018 03 01 Reconstructing the genetic history of late Neanderthals Nature 555 7698 652 656 Bibcode 2018Natur 555 652H doi 10 1038 nature26151 ISSN 1476 4687 PMC 6485383 PMID 29562232 Meyer Matthias Arsuaga Juan Luis de Filippo Cesare Nagel Sarah et al 2016 03 01 Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins Nature 531 7595 504 507 Bibcode 2016Natur 531 504M doi 10 1038 nature17405 ISSN 1476 4687 PMID 26976447 S2CID 4467094 Gomez Robles Aida 2019 05 01 Dental evolutionary rates and its implications for the Neanderthal modern human divergence Science Advances 5 5 1268 Bibcode 2019SciA 5 1268G doi 10 1126 sciadv aaw1268 ISSN 2375 2548 PMC 6520022 PMID 31106274 Bailey John Wendell 1946 The Mammals of Virginia p 356 Hall E 1946 Zoological Subspecies of Man at the Peace Table Journal of Mammalogy 27 4 358 364 doi 10 2307 1375342 JSTOR 1375342 PMID 20247535 Grzimek Bernhard 1970 Grzimek s Animal Life Encyclopedia Vol 11 p 55 ISBN 978 0442784782 Hublin J J 2009 The origin of Neandertals Proceedings of the National Academy of Sciences 106 38 16022 16027 Bibcode 2009PNAS 10616022H doi 10 1073 pnas 0904119106 JSTOR 40485013 PMC 2752594 PMID 19805257 Harvati K Frost S R McNulty K P 2004 Neanderthal taxonomy reconsidered implications of 3D primate models of intra and interspecific differences Proc Natl Acad Sci U S A 101 5 1147 1152 Bibcode 2004PNAS 101 1147H doi 10 1073 pnas 0308085100 PMC 337021 PMID 14745010 Homo neanderthalensis King 1864 Wiley Blackwell Encyclopedia of Human Evolution Chichester West Sussex Wiley Blackwell 2013 pp 328 331 a b c d Rafferty John P Homo sapiens sapiens Encyclopedia Britannica Retrieved 2020 08 11 Schlebusch Carina M Malmstrom Helena Gunther Torsten Sjodin Per Coutinho Alexandra Edlund Hanna Munters Arielle R Steyn Maryna Soodyall Himla Lombard Marlize Jakobsson Mattias 5 June 2017 Ancient genomes from southern Africa pushes modern human divergence beyond 260 000 years ago bioRxiv 10 1101 145409 Schlebusch Carina M Malmstrom Helena Gunther Torsten Sjodin Per Coutinho Alexandra Edlund Hanna Munters Arielle R Vicente Mario Steyn Maryna Soodyall Himla Lombard Marlize 2017 11 03 Southern African ancient genomes estimate modern human divergence to 350 000 to 260 000 years ago Science 358 6363 652 655 Bibcode 2017Sci 358 652S doi 10 1126 science aao6266 ISSN 0036 8075 PMID 28971970 a b c Callaway Ewan 7 June 2017 Oldest Homo sapiens fossil claim rewrites our species history Nature doi 10 1038 nature 2017 22114 Retrieved 11 June 2017 a b c d e Stringer C 2016 The origin and evolution of Homo sapiens Philosophical Transactions of the Royal Society of London Series B Biological Sciences 371 1698 20150237 doi 10 1098 rstb 2015 0237 PMC 4920294 PMID 27298468 a b Sample Ian 7 June 2017 Oldest Homo sapiens bones ever found shake foundations of the human story The Guardian Retrieved 7 June 2017 a b Hublin Jean Jacques Ben Ncer Abdelouahed Bailey Shara E Freidline Sarah E Neubauer Simon Skinner Matthew M Bergmann Inga Le Cabec Adeline Benazzi Stefano Harvati Katerina Gunz Philipp 2017 New fossils from Jebel Irhoud Morocco and the pan African origin of Homo sapiens PDF Nature 546 7657 289 292 Bibcode 2017Natur 546 289H doi 10 1038 nature22336 PMID 28593953 a b Scerri M L et al 2018 Did Our Species Evolve in Subdivided Populations across Africa and Why Does It Matter Trends in Ecology amp Evolution 33 8 582 594 doi 10 1016 j tree 2018 05 005 PMC 6092560 PMID 30007846 Chan Eva K F 28 October 2019 Human origins in a southern African palaeo wetland and first migrations Nature 857 7781 185 189 Bibcode 2019Natur 575 185C doi 10 1038 s41586 019 1714 1 PMID 31659339 S2CID 204946938 Sample Ian 28 October 2019 Ancestral home of modern humans is in Botswana study finds The Guardian ISSN 0261 3077 Retrieved 29 October 2019 Woodward Aylin 28 October 2019 New Study Pinpoints The Ancestral Homeland of All Humans Alive Today ScienceAlert com Retrieved 29 October 2019 a b Yong Ed 28 October 2019 Has Humanity s Homeland Been Found The Atlantic Retrieved 28 October 2019 a b c Zimmer Carl 10 September 2019 Scientists Find the Skull of Humanity s Ancestor on a Computer By comparing fossils and CT scans researchers say they have reconstructed the skull of the last common forebear of modern humans The New York Times Retrieved 10 September 2019 Rogers Alan R Bohlender Ryan J Huff Chad D 12 September 2017 Early history of Neanderthals and Denisovans Proceedings of the National Academy of Sciences 114 37 9859 9863 doi 10 1073 pnas 1706426114 PMC 5604018 PMID 28784789 Wolpoff M H Spuhler J N Smith F H Radovcic J Pope G Frayer D W Eckhardt R Clark G 1988 Modern Human Origins Science 241 4867 772 774 Bibcode 1988Sci 241 772W doi 10 1126 science 3136545 PMID 3136545 S2CID 5223638 a b c Green RE Krause J Briggs AW Maricic T Stenzel U Kircher M Patterson N Li H Zhai W Fritz MH Hansen NF Durand EY Malaspinas A Jensen JD Marques Bonet T Alkan C Prufer K Meyer M Burbano HA Good JM Schultz R Aximu Petri A Butthof A Hober B Hoffner B Siegemund M Weihmann A Nusbaum C Lander ES et al May 2010 A draft sequence of the Neandertal genome Science 328 5979 710 722 Bibcode 2010Sci 328 710G doi 10 1126 science 1188021 PMC 5100745 PMID 20448178 Reich D Patterson N Kircher M Delfin F Nandineni MR Pugach I Ko AM Ko Y Jinam TA Phipps ME Saitou N Wollstein A Kayser M Paabo S Stoneking M 2011 Denisova admixture and the first modern human dispersals into southeast Asia and oceania Am J Hum Genet 89 4 516 528 doi 10 1016 j ajhg 2011 09 005 PMC 3188841 PMID 21944045 New Clues Add 40 000 Years to Age of Human Species www nsf gov NSF National Science Foundation Age of ancient humans reassessed BBC News February 16 2005 Retrieved April 10 2010 The Oldest Homo Sapiens Fossils Push Human Emergence Back To 195 000 Years Ago ScienceDaily February 28 2005 Retrieved 2019 05 06 Alemseged Z Coppens Y Geraads D 2002 Hominid cranium from Homo Description and taxonomy of Homo 323 1976 896 PDF Am J Phys Anthropol 117 2 103 112 doi 10 1002 ajpa 10032 PMID 11815945 Stoneking Mark Soodyall Himla 1996 Human evolution and the mitochondrial genome Current Opinion in Genetics amp Development 6 6 731 736 doi 10 1016 S0959 437X 96 80028 1 PMID 8994844 Human evolution the fossil evidence in 3D by Philip L Walker and Edward H Hagen Dept of Anthropology University of California Santa Barbara Retrieved April 5 2005 Meyer Matthias Arsuaga Juan Luis de Filippo Cesare Nagel Sarah Aximu Petri Ayinuer Nickel Birgit Martinez Ignacio Gracia Ana de Castro Jose Maria Bermudez Carbonell Eudald Viola Bence Kelso Janet Prufer Kay Paabo Svante 14 March 2016 Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins Nature 531 7595 504 507 Bibcode 2016Natur 531 504M doi 10 1038 nature17405 PMID 26976447 S2CID 4467094 Callaway Ewen 14 March 2016 Oldest ancient human DNA details dawn of Neanderthals Nature 531 7594 296 286 Bibcode 2016Natur 531 296C doi 10 1038 531286a PMID 26983523 S2CID 4459329 Oppenheimer S 2003 Out of Eden The Peopling of the World ISBN 978 1841196978 Trinkaus E Moldovan O Milota Ș Bilgăr A Sarcina L Athreya S Bailey S E Rodrigo R et al 2003 An early modern human from Peștera cu Oase Romania PNAS 100 20 11231 11236 Bibcode 2003PNAS 10011231T doi 10 1073 pnas 2035108100 PMC 208740 PMID 14504393 a b Reich David Green Richard E Kircher Martin Krause Johannes Patterson Nick Durand Eric Y Viola Bence Briggs Adrian W et al 2010 Genetic history of an archaic hominin group from Denisova Cave in Siberia Nature 468 7327 1053 1060 Bibcode 2010Natur 468 1053R doi 10 1038 nature09710 hdl 10230 25596 PMC 4306417 PMID 21179161 Trinkaus Erik October 2005 Early modern humans Annual Review of Anthropology 34 1 207 230 doi 10 1146 annurev anthro 34 030905 154913 S2CID 9039428 Meldrum Jeff Hilton Charles E 2004 From Biped to Strider The Emergence of Modern Human Walking Running and Resource Transport Springer Science amp Business Media ISBN 978 0306480003 Vonk Jennifer Shackelford Todd K 2012 The Oxford Handbook of Comparative Evolutionary Psychology Oxford University Press US pp 429 ISBN 978 0199738182 Bozek Katarzyna Wei Yuning Yan Zheng Liu Xiling Xiong Jieyi Sugimoto Masahiro Tomita Masaru Paabo Svante Pieszek Raik Sherwood Chet C Hof Patrick R Ely John J Steinhauser Dirk Willmitzer Lothar Bangsbo Jens Hansson Ola Call Josep Giavalisco Patrick Khaitovich Philipp 2014 Exceptional Evolutionary Divergence of Human Muscle and Brain Metabolomes Parallels Human Cognitive and Physical Uniqueness PLOS Biology 12 5 e1001871 doi 10 1371 journal pbio 1001871 PMC 4035273 PMID 24866127 Thieme H 2007 Der grosse Wurf von Schoningen Das neue Bild zur Kultur des fruhen Menschen Die Schoninger Speere Mensch und Jagd vor 400 000 Jahren Konrad Theiss Verlag pp 224 328 ISBN 978 3896460400 Haidle M N 2006 Menschenaffen Affenmenschen Mensch Kognition und Sprache im Altpalaolithikum In Conard N J ed Woher kommt der Mensch Attempto Verlag pp 69 97 ISBN 3893083812 Mendez Fernando Krahn Thomas Schrack Bonnie Krahn Astrid Maria Veeramah Krishna Woerner August Fomine Forka Leypey Mathew Bradman Neil Thomas Mark 7 March 2013 An African American paternal lineage adds an extremely ancient root to the human Y chromosome phylogenetic tree PDF American Journal of Human Genetics 92 3 454 459 doi 10 1016 j ajhg 2013 02 002 PMC 3591855 PMID 23453668 Krings M Stone A Schmitz RW Krainitzki H Stoneking M Paabo S July 1997 Neandertal DNA sequences and the origin of modern humans Cell 90 1 19 30 doi 10 1016 S0092 8674 00 80310 4 hdl 11858 00 001M 0000 0025 0960 8 PMID 9230299 S2CID 13581775 Hill Deborah 16 March 2004 No Neandertals in the Gene Pool Science Retrieved 2019 05 06 Serre D Langaney A Chech M Teschler Nicola M Paunovic M Mennecier P Hofreiter M Possnert G Paabo S 2004 No evidence of Neandertal mtDNA contribution to early modern humans PLOS Biology 2 3 313 317 doi 10 1371 journal pbio 0020057 PMC 368159 PMID 15024415 Hershkovitz I Smith P Sarig R Quam R Rodriguez L Garcia R Arsuaga JL Barkai R Gopher A 2011 Middle pleistocene dental remains from Qesem Cave Israel American Journal of Physical Anthropology 144 4 575 592 doi 10 1002 ajpa 21446 PMID 21404234 S2CID 3106938 Posth Cosimo et al 4 July 2017 Deeply divergent archaic mitochondrial genome provides lower time boundary for African gene flow into Neanderthals Nature Communications 8 16046 Bibcode 2017NatCo 816046P doi 10 1038 ncomms16046 PMC 5500885 PMID 28675384 White Tim D Asfaw Berhane Degusta David Gilbert Henry Richards Gary D Suwa Gen Howell Clark F June 2003 Pleistocene Homo sapiens from Middle Awash Ethiopia Nature 423 6941 742 747 Bibcode 2003Natur 423 742W doi 10 1038 nature01669 PMID 12802332 S2CID 4432091 Fossil Reanalysis Pushes Back Origin of Homo sapiens Scientific American 2005 02 17 Retrieved 2019 05 06 Mehta Ankita 26 January 2018 A 177 000 year old jawbone fossil discovered in Israel is oldest human remains found outside Africa International Business Times Retrieved 2019 05 06 Bae Christopher J Douka Katerina Petraglia Michael D 8 December 2017 On the origin of modern humans Asian perspectives Science 358 6368 eaai9067 doi 10 1126 science aai9067 PMID 29217544 Kuo Lily 10 December 2017 Early humans migrated out of Africa much earlier than we thought Quartz Retrieved 2019 05 06 Zimmer Carl 10 July 2019 A Skull Bone Discovered in Greece May Alter the Story of Human Prehistory The bone found in a cave is the oldest modern human fossil ever discovered in Europe It hints that humans began leaving Africa far earlier than once thought The New York Times Retrieved 11 July 2019 Staff 10 July 2019 Oldest remains outside Africa reset human migration clock Phys org Retrieved 10 July 2019 Harvati Katerina et al 10 July 2019 Apidima Cave fossils provide earliest evidence of Homo sapiens in Eurasia Nature 571 7766 500 504 doi 10 1038 s41586 019 1376 z PMID 31292546 S2CID 195873640 Rito T Richards MB Fernandes V Alshamali F Cerny V Pereira L Soares P 2013 The first modern human dispersals across Africa PLOS ONE 8 11 e80031 Bibcode 2013PLoSO 880031R doi 10 1371 journal pone 0080031 PMC 3827445 PMID 24236171 Henn Brenna Gignoux Christopher R Jobin Matthew 2011 Hunter gatherer genomic diversity suggests a southern African origin for modern humans Proceedings of the National Academy of Sciences of the United States of America 108 13 5154 5162 Bibcode 2011PNAS 108 5154H doi 10 1073 pnas 1017511108 PMC 3069156 PMID 21383195 a b c Higham Thomas F G Wesselingh Frank P Hedges Robert E M Bergman Christopher A Douka Katerina 2013 09 11 Chronology of Ksar Akil Lebanon and Implications for the Colonization of Europe by Anatomically Modern Humans PLOS ONE 8 9 e72931 Bibcode 2013PLoSO 872931D doi 10 1371 journal pone 0072931 ISSN 1932 6203 PMC 3770606 PMID 24039825 Posth C Renaud G Mittnik M Drucker DG Rougier H Cupillard C Valentin F Thevenet C Furtwangler A Wissing C Francken M Malina M Bolus M Lari M Gigli E Capecchi G Crevecoeur I Beauval C Flas D Germonpre M van der Plicht J Cottiaux R Gely B Ronchitelli A Wehrberger K Grigorescu D Svoboda J Semal P Caramelli D Bocherens H Harvati K Conard NJ Haak W Powell A Krause J 2016 Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non Africans and a Late Glacial Population Turnover in Europe Current Biology 26 6 827 833 doi 10 1016 j cub 2016 01 037 hdl 2440 114930 PMID 26853362 S2CID 140098861 Kamin M Saag L Vincente M et al April 2015 A recent bottleneck of Y chromosome diversity coincides with a global change in culture Genome Research 25 4 459 466 doi 10 1101 gr 186684 114 PMC 4381518 PMID 25770088 Vai S Sarno S Lari M Luiselli D Manzi G Gallinaro M Mataich S Hubner A Modi A Pilli E Tafuri MA Caramelli D di Lernia S March 2019 Ancestral mitochondrial N lineage from the Neolithic green Sahara Sci Rep 9 1 3530 Bibcode 2019NatSR 9 3530V doi 10 1038 s41598 019 39802 1 PMC 6401177 PMID 30837540 a b Haber M Jones AL Connel BA Asan Arciero E Huanming Y Thomas MG Xue Y Tyler Smith C June 2019 A Rare Deep Rooting D0 African Y chromosomal Haplogroup and its Implications for the Expansion of Modern Humans Out of Africa Genetics 212 4 1421 1428 doi 10 1534 genetics 119 302368 PMC 6707464 PMID 31196864 Clarkson Chris Jacobs Zenobia Pardoe Colin 2017 Human occupation of northern Australia by 65 000 years ago PDF Nature 547 7663 306 310 Bibcode 2017Natur 547 306C doi 10 1038 nature22968 hdl 2440 107043 PMID 28726833 S2CID 205257212 St Fleu Nicholas July 19 2017 Humans First Arrived in Australia 65 000 Years Ago Study Suggests The New York Times Wood R 2017 09 02 Comments on the chronology of Madjedbebe Australian Archaeology 83 3 172 174 doi 10 1080 03122417 2017 1408545 ISSN 0312 2417 S2CID 148777016 O Connell JF Allen J Williams MA Williams AN Turney CS Spooner NA et al August 2018 Homo sapiens first reach Southeast Asia and Sahul Proceedings of the National Academy of Sciences of the United States of America 115 34 8482 8490 doi 10 1073 pnas 1808385115 PMC 6112744 PMID 30082377 Liu Hua et al 2006 A Geographically Explicit Genetic Model of Worldwide Human Settlement History The American Journal of Human Genetics 79 2 230 237 doi 10 1086 505436 PMC 1559480 PMID 16826514 Out of Africa Revisited Science 308 5724 921g 2005 05 13 doi 10 1126 science 308 5724 921g S2CID 220100436 Sankararaman Sriram Mallick Swapan Patterson Nick Reich David 2016 The Combined Landscape of Denisovan and Neanderthal Ancestry in Present Day Humans Current Biology 26 9 1241 1247 doi 10 1016 j cub 2016 03 037 ISSN 0960 9822 PMC 4864120 PMID 27032491 Sanchez Quinto F Botigue LR Civit S Arenas C Avila Arcos MC Bustamante CD Comas D Lalueza Fox C October 17 2012 North African Populations Carry the Signature of Admixture with Neandertals PLOS ONE 7 10 e47765 Bibcode 2012PLoSO 747765S doi 10 1371 journal pone 0047765 PMC 3474783 PMID 23082212 Fu Q Li H Moorjani P Jay F Slepchenko SM Bondarev AA Johnson PL Aximu Petri A Prufer K de Filippo C Meyer M Zwyns N Salazar Garcia DC Kuzmin YV Keates SG Kosintsev PA Razhev DI Richards MP Peristov NV Lachmann M Douka K Higham TF Slatkin M Hublin JJ Reich D Kelso J Viola TB Paabo S October 23 2014 Genome sequence of a 45 000 year old modern human from western Siberia Nature 514 7523 445 449 Bibcode 2014Natur 514 445F doi 10 1038 nature13810 PMC 4753769 PMID 25341783 Brahic Catherine February 3 2014 Humanity s forgotten return to Africa revealed in DNA The New Scientist Retrieved 2019 05 06 Kuhlwilm Martin 17 February 2016 Ancient gene flow from early modern humans into Eastern Neanderthals Nature 530 7591 429 433 Bibcode 2016Natur 530 429K doi 10 1038 nature16544 PMC 4933530 PMID 26886800 Ding Q Hu Y Xu S Wang J Jin L 2014 Online 2013 Neanderthal Introgression at Chromosome 3p21 31 was Under Positive Natural Selection in East Asians Molecular Biology and Evolution 31 3 683 695 doi 10 1093 molbev mst260 PMID 24336922 Vernot B Akey J M 2014 Resurrecting Surviving Neandertal Lineages from Modern Human Genomes Science 343 6174 1017 1021 Bibcode 2014Sci 343 1017V doi 10 1126 science 1245938 PMID 24476670 S2CID 23003860 Ayala Francisco Jose Conde Camilo Jose Cela 2017 Processes in Human Evolution The Journey from Early Hominins to Neanderthals and Modern Humans ISBN 978 0198739906 Schopf J William 1992 Major Events in the History of Life Jones amp Bartlett Learning pp 168 ISBN 978 0867202687 Paabo Svante 2014 Neanderthal Man In Search of Lost Genomes New York Basic Books p 237 Sanders Robert 11 June 2003 160 000 year old fossilized skulls uncovered in Ethiopia are oldest anatomically modern humans UC Berkeley News Retrieved 2019 05 07 White Tim D Asfaw B DeGusta D Gilbert H Richards G D Suwa G Howell F C 2003 Pleistocene Homo sapiens from Middle Awash Ethiopia Nature 423 6491 742 747 Bibcode 2003Natur 423 742W doi 10 1038 nature01669 PMID 12802332 S2CID 4432091 Stringer Chris June 12 2003 Human evolution Out of Ethiopia Nature 423 6941 693 695 Bibcode 2003Natur 423 692S doi 10 1038 423692a PMID 12802315 S2CID 26693109 Hawks J Wang E T Cochran G M Harpending H C Moyzis R K 2007 Recent acceleration of human adaptive evolution Proceedings of the National Academy of Sciences 104 52 20753 20758 Bibcode 2007PNAS 10420753H doi 10 1073 pnas 0707650104 PMC 2410101 PMID 18087044 Bhupendra P April 2019 Forehead Anatomy Medscape references Retrieved 2019 05 06 How to ID a modern human News 2012 Natural History Museum London Retrieved 11 December 2013 Encarta Human Evolution Encarta Archived from the original on 29 October 2009 Morris Desmond 2007 The Brow The Naked Woman A Study of the Female Body ISBN 978 0312338534 Pearce Eiluned Stringer Chris Dunbar R I M 2013 05 07 New insights into differences in brain organization between Neanderthals and anatomically modern humans Proceedings of the Royal Society of London B Biological Sciences 280 1758 20130168 doi 10 1098 rspb 2013 0168 ISSN 0962 8452 PMC 3619466 PMID 23486442 Smith C L Beals K L 1990 Cultural correlates with cranial capacity American Anthropologist 92 193 200 doi 10 1525 aa 1990 92 1 02a00150 S2CID 162406199 Stringer C 1984 Human evolution and biological adaptation in the Pleistocene In Foley R ed Hominid evolution and community ecology New York Academic Press ISBN 978 0122619205 Townsend G Richards L Hughes T May 2003 Molar intercuspal dimensions genetic input to phenotypic variation Journal of Dental Research 82 5 350 355 doi 10 1177 154405910308200505 PMID 12709500 S2CID 26123427 Keith A 1913 Problems relating to the Teeth of the Earlier Forms of Prehistoric Man Proceedings of the Royal Society of Medicine 6 Odontol Sect 103 124 doi 10 1177 003591571300601018 PMC 2005996 PMID 19977113 Tattersall Jeffrey H Schwartz Ian 2003 The human fossil record Craniodental Morphology of Genus Homo Africa and Asia vol 2 Wiley Liss pp 327 328 ISBN 978 0471319283 Steegmann A Theodore Cerny Frank J Holliday Trenton W 2002 Neandertal cold adaptation Physiological and energetic factors American Journal of Human Biology 14 5 566 583 doi 10 1002 ajhb 10070 PMID 12203812 S2CID 2437566 Stock J T October 2006 Hunter gatherer postcranial robusticity relative to patterns of mobility climatic adaptation and selection for tissue economy American Journal of Physical Anthropology 131 2 194 204 doi 10 1002 ajpa 20398 PMID 16596600 Helmuth H 1998 Body height body mass and surface area of the Neanderthals Zeitschrift fur Morphologie und Anthropologie 82 1 1 12 doi 10 1127 zma 82 1998 1 PMID 9850627 Lim TO Ding LM Zaki M et al March 2000 Distribution of Body Weight Height and Body Mass Index in a National Sample of Malaysian Adults PDF Med J Malaysia 55 1 108 128 PMID 11072496 Wade N 2006 03 07 Still Evolving Human Genes Tell New Story The New York Times Retrieved 2008 07 10 Khrameeva E Bozek K He L Yan Z Jiang X Wei Y Tang K Gelfand MS Prufer K Kelso J Paabo S Giavalisco P Lachmann M Khaitovich P 2014 Neanderthal ancestry drives evolution of lipid catabolism in contemporary Europeans Nature Communications 5 3584 3584 Bibcode 2014NatCo 5 3584K doi 10 1038 ncomms4584 PMC 3988804 PMID 24690587 Michael Dannemann 1 and Janet Kelso The Contribution of Neanderthals to Phenotypic Variation in Modern Humans The American Journal of Human Genetics 101 578 589 October 5 2017 Kamberov Yana G 14 February 2013 Modeling Recent Human Evolution in Mice by Expression of a Selected EDAR Variant Cell 152 4 691 702 doi 10 1016 j cell 2013 01 016 PMC 3575602 PMID 23415220 Wade Nicholas 14 February 2013 East Asian Physical Traits Linked to 35 000 Year Old Mutation The New York Times Retrieved 2019 05 06 a b Beleza Sandra Santos A M McEvoy B Alves I Martinho C Cameron E Shriver M D Parra E J Rocha J 2012 The timing of pigmentation lightening in Europeans Molecular Biology and Evolution 30 1 24 35 doi 10 1093 molbev mss207 PMC 3525146 PMID 22923467 Lalueza Fox Rompler H Caramelli D Staubert C Catalano G Hughes D Rohland N Pilli E Longo L Condemi S de la Rasilla M Fortea J Rosas A Stoneking M Schoneberg T Bertranpetit J Hofreiter M et al 2007 A melanocortin 1 receptor allele suggests varying pigmentation among Neanderthals Science 318 5855 1453 1455 Bibcode 2007Sci 318 1453L doi 10 1126 science 1147417 PMID 17962522 S2CID 10087710 a b Beals Kenneth L Smith Courtland L Dodd Stephen M 1984 Brain Size Cranial Morphology Climate and Time Machines Current Anthropology 25 3 301 330 doi 10 1086 203138 S2CID 86147507 Nowaczewska Wioletta Dabrowski Pawel Kuzminski Lukasz 2011 Morphological Adaptation to Climate in Modern Homo sapiens Crania The Importance of Basicranial Breadth Collegium Antropologicum 35 3 625 636 PMID 22053534 Archived from the original on 2014 10 11 Peng Y et al 2010 The ADH1B Arg47His polymorphism in East Asian populations and expansion of rice domestication in history BMC Evolutionary Biology 10 15 doi 10 1186 1471 2148 10 15 PMC 2823730 PMID 20089146 Segurel Laure Bon Celine 2017 On the Evolution of Lactase Persistence in Humans Annual Review of Genomics and Human Genetics 18 1 297 319 doi 10 1146 annurev genom 091416 035340 PMID 28426286 Ingram Catherine J E Mulcare Charlotte A Itan Yuval Thomas Mark G Swallow Dallas M 2008 11 26 Lactose digestion and the evolutionary genetics of lactase persistence Human Genetics 124 6 579 591 doi 10 1007 s00439 008 0593 6 ISSN 0340 6717 PMID 19034520 S2CID 3329285 Ilardo M A Moltke I Korneliussen T S Cheng J Stern A J Racimo F de Barros Damgaard P Sikora M Seguin Orlando A Rasmussen S van den Munckhof I C L ter Horst R Joosten L A B Netea M G Salingkat S Nielsen R Willerslev E 2018 04 18 Physiological and Genetic Adaptations to Diving in Sea Nomads Cell 173 3 569 580 e15 doi 10 1016 j cell 2018 03 054 PMID 29677510 Gislen A Dacke M Kroger RH Abrahamsson M Nilsson DE Warrant EJ 2003 Superior Underwater Vision in a Human Population of Sea Gypsies Current Biology 13 10 833 836 doi 10 1016 S0960 9822 03 00290 2 PMID 12747831 S2CID 18731746 Klein Richard 1995 Anatomy behavior and modern human origins Journal of World Prehistory 9 2 167 198 doi 10 1007 bf02221838 S2CID 10402296 Feierman Jay R 2009 The Biology of Religious Behavior The Evolutionary Origins of Faith and Religion ABC CLIO p 220 ISBN 978 0313364303 Soressi M 2005 Late Mousterian lithic technology Its implications for the pace of the emergence of behavioural modernity and the relationship between behavioural modernity and biological modernity pp 389 417 in L Backwell et F d Errico eds From Tools to Symbols Johannesburg University of Witswatersand Press ISBN 1868144178 Companion encyclopedia of archaeology 1999 Routledge ISBN 0415213304 Vol 2 p 763 cf effectively limited to organic samples ed organic compounds or biogenic carbonates that date to less than 50 ka 50 000 years ago See also Later Stone Age and Upper Paleolithic Mellars Paul 2006 Why did modern human populations disperse from Africa ca 60 000 years ago Proceedings of the National Academy of Sciences 103 25 9381 9386 Bibcode 2006PNAS 103 9381M doi 10 1073 pnas 0510792103 PMC 1480416 PMID 16772383 Shea John 2011 Homo sapiens Is As Homo sapiens Was Current Anthropology 52 1 1 35 doi 10 1086 658067 S2CID 142517998 McBrearty Sally Brooks Allison 2000 The revolution that wasn t a new interpretation of the origin of modern human behavior Journal of Human Evolution 39 5 453 563 doi 10 1006 jhev 2000 0435 PMID 11102266 S2CID 42968840 Henshilwood Christopher Marean Curtis 2003 The Origin of Modern Human Behavior Critique of the Models and Their Test Implications Current Anthropology 44 5 627 651 doi 10 1086 377665 PMID 14971366 S2CID 11081605 Marean Curtis et al 2007 Early human use of marine resources and pigment in South Africa during the Middle Pleistocene PDF Nature 449 7164 905 908 Bibcode 2007Natur 449 905M doi 10 1038 nature06204 PMID 17943129 S2CID 4387442 Powell Adam et al 2009 Late Pleistocene Demography and the Appearance of Modern Human Behavior PDF Science 324 5932 1298 1301 Bibcode 2009Sci 324 1298P doi 10 1126 science 1170165 PMID 19498164 S2CID 206518315 Premo Luke Kuhn Steve 2010 Modeling Effects of Local Extinctions on Culture Change and Diversity in the Paleolithic PLOS ONE 5 12 e15582 Bibcode 2010PLoSO 515582P doi 10 1371 journal pone 0015582 PMC 3003693 PMID 21179418 St Fleur Nicholas 12 September 2018 Oldest Known Drawing by Human Hands Discovered in South African Cave The New York Times Retrieved 15 September 2018 d Errico F Backwell L Villa P Degano I Lucejko J J Bamford M K Higham T F G Colombini M P Beaumont P B 2012 Early evidence of San material culture represented by organic artifacts from Border Cave South Africa Proceedings of the National Academy of Sciences 109 33 13214 13219 Bibcode 2012PNAS 10913214D doi 10 1073 pnas 1204213109 PMC 3421171 PMID 22847420 Mourre V Villa P Henshilwood C S 2010 Early Use of Pressure Flaking on Lithic Artifacts at Blombos Cave South Africa Science 330 6004 659 662 Bibcode 2010Sci 330 659M doi 10 1126 science 1195550 PMID 21030655 S2CID 34833884 a b Brown Kyle S Marean Curtis W Jacobs Zenobia Schoville Benjamin J Oestmo Simen Fisher Erich C Bernatchez Jocelyn Karkanas Panagiotis Matthews Thalassa 2012 An early and enduring advanced technology originating 71 000 years ago in South Africa Nature 491 7425 590 593 Bibcode 2012Natur 491 590B doi 10 1038 nature11660 PMID 23135405 S2CID 4323569 Scholz C A Johnson T C Cohen A S King J W Peck J A Overpeck J T Talbot M R Brown E T Kalindekafe L Amoako P Y O Lyons R P Shanahan T M Castaneda I S Heil C W Forman S L McHargue L R Beuning K R Gomez J Pierson J 2007 East African megadroughts between 135 and 75 thousand years ago and bearing on early modern human origins Proceedings of the National Academy of Sciences 104 42 16416 16421 Bibcode 2007PNAS 10416416S doi 10 1073 pnas 0703874104 PMC 1964544 PMID 17785420 Wells Spencer 2003 The Journey of Man A Genetic Odyssey Princeton N J Princeton University Press ISBN 978 0691115320 Henshilwood Christopher et al 2002 Emergence of Modern Human Behavior Middle Stone Age Engravings from South Africa Science 295 5558 1278 1280 Bibcode 2002Sci 295 1278H doi 10 1126 science 1067575 PMID 11786608 S2CID 31169551 Henshilwood Christopher S d Errico Francesco Watts Ian 2009 Engraved ochres from the Middle Stone Age levels at Blombos Cave South Africa Journal of Human Evolution 57 1 27 47 doi 10 1016 j jhevol 2009 01 005 PMID 19487016 Texier PJ Porraz G Parkington J Rigaud JP Poggenpoel C Miller C Tribolo C Cartwright C Coudenneau A Klein R Steele T Verna C 2010 A Howiesons Poort tradition of engraving ostrich eggshell containers dated to 60 000 years ago at Diepkloof Rock Shelter South Africa Proceedings of the National Academy of Sciences 107 14 6180 6185 Bibcode 2010PNAS 107 6180T doi 10 1073 pnas 0913047107 PMC 2851956 PMID 20194764 a b McBrearty Sally Brooks Allison 2000 The revolution that wasn t a new interpretation of the origin of modern human behavior Journal of Human Evolution 39 5 453 563 doi 10 1006 jhev 2000 0435 PMID 11102266 S2CID 42968840 Henshilwood Christopher S et al 2004 Middle Stone Age shell beads from South Africa Science 304 5669 404 doi 10 1126 science 1095905 PMID 15087540 S2CID 32356688 d Errico Francesco et al 2005 Nassarius kraussianus shell beads from Blombos Cave evidence for symbolic behaviour in the Middle Stone Age Journal of Human Evolution 48 1 3 24 doi 10 1016 j jhevol 2004 09 002 PMID 15656934 Vanhaeren Marian et al 2013 Thinking strings Additional evidence for personal ornament use in the Middle Stone Age at Blombos Cave South Africa Journal of Human Evolution 64 6 500 517 doi 10 1016 j jhevol 2013 02 001 PMID 23498114 Backwell L d Errico F Wadley L 2008 Middle Stone Age bone tools from the Howiesons Poort layers Sibudu Cave South Africa Journal of Archaeological Science 35 6 1566 1580 doi 10 1016 j jas 2007 11 006 Wadley Lyn 2008 The Howieson s Poort industry of Sibudu Cave South African Archaeological Society Goodwin Series 10 Lombard M Phillips L 2010 Indications of bow and stone tipped arrow use 64 000 years ago in KwaZulu Natal South Africa Antiquity 84 325 635 648 doi 10 1017 S0003598X00100134 S2CID 162438490 Lombard M 2011 Quartz tipped arrows older than 60 ka further use trace evidence from Sibudu Kwa Zulu Natal South Africa Journal of Archaeological Science 38 8 1918 1930 doi 10 1016 j jas 2011 04 001 Backwell L Bradfield J Carlson KJ Jashashvili T Wadley L d Errico F 2018 The antiquity of bow and arrow technology evidence from Middle Stone Age layers at Sibudu Cave Journal of Archaeological Science 92 362 289 303 doi 10 15184 aqy 2018 11 Lombard M 2020 The tip cross sectional areas of poisoned bone arrowheads from southern Africa Journal of Archaeological Science Reports 33 102477 doi 10 1016 j jasrep 2020 102477 S2CID 224889105 Yellen JE AS Brooks E Cornelissen MJ Mehlman K Stewart 28 April 1995 A middle stone age worked bone industry from Katanda Upper Semliki Valley Zaire Science 268 5210 553 556 Bibcode 1995Sci 268 553Y doi 10 1126 science 7725100 PMID 7725100 Brown Kyle S Marean Curtis W Herries Andy I R Jacobs Zenobia Tribolo Chantal Braun David Roberts David L Meyer Michael C Bernatchez J 14 August 2009 Fire as an Engineering Tool of Early Modern Humans Science 325 5942 859 862 Bibcode 2009Sci 325 859B doi 10 1126 science 1175028 hdl 11422 11102 PMID 19679810 S2CID 43916405 Amos Jonathan 13 October 2011 A Cultural Leap at the Dawn of Humanity Ancient paint factory unearthed BBC News Retrieved 13 October 2011 Vastag Brian 13 October 2011 South African cave yields paint from dawn of humanity The Washington Post Retrieved 13 October 2011 Henshilwood Christopher S et al 2011 A 100 000 Year Old Ochre Processing Workshop at Blombos Cave South Africa Science 334 6053 219 222 Bibcode 2011Sci 334 219H doi 10 1126 science 1211535 PMID 21998386 S2CID 40455940 Shipton C d Errico F Petraglia M et al 2018 78 000 year old record of Middle and Later Stone Age innovation in an East African tropical forest Nature Communications Sahle Y Hutchings W K Braun D R Sealy J C Morgan L E Negash A Atnafu B 2013 Petraglia Michael D ed Earliest Stone Tipped Projectiles from the Ethiopian Rift Date to gt 279 000 Years Ago PLOS ONE 8 11 e78092 Bibcode 2013PLoSO 878092S doi 10 1371 journal pone 0078092 PMC 3827237 PMID 24236011 Marean Curtis et al 2007 Early human use of marine resources and pigment in South Africa during the Middle Pleistocene PDF Nature 449 7164 905 908 Bibcode 2007Natur 449 905M doi 10 1038 nature06204 PMID 17943129 S2CID 4387442 5 Oldest Mines in the World A Casual Survey Archived from the original on 2019 01 05 Retrieved 2019 09 11 Guinness World Records 10 September 2015 Guinness World Records 2016 Guinness World Records p 27 ISBN 978 1910561034 Chatterjee Rhitu 15 March 2018 Scientists Are Amazed By Stone Age Tools They Dug Up In Kenya NPR Retrieved 15 March 2018 Yong Ed 15 March 2018 A Cultural Leap at the Dawn of Humanity New finds from Kenya suggest that humans used long distance trade networks sophisticated tools and symbolic pigments right from the dawn of our species The Atlantic Retrieved 15 March 2018 Brooks AS Yellen JE Potts R Behrensmeyer AK Deino AL Leslie DE Ambrose SH Ferguson JR d Errico F Zipkin AM Whittaker S Post J Veatch EG Foecke K Clark JB 2018 Long distance stone transport and pigment use in the earliest Middle Stone Age Science 360 6384 90 94 Bibcode 2018Sci 360 90B doi 10 1126 science aao2646 PMID 29545508 Sahle Y Brooks AS 2018 Assessment of complex projectiles in the early Late Pleistocene at Aduma Ethiopia PLOS ONE 14 5 e0216716 Bibcode 2019PLoSO 1416716S doi 10 1371 journal pone 0216716 PMC 6508696 PMID 31071181 Douglas Kate March 24 2012 Puzzles of Evolution Why was technological development so slow New Scientist Archived from the original on April 13 2016 Retrieved May 4 2022 a href Template Cite magazine html title Template Cite magazine cite magazine a CS1 maint unfit URL link Further readingReich David 2018 Who We Are And How We Got Here Ancient DNA and the New Science of the Human Past Pantheon Books ISBN 978 1101870327 1 External linksHuman Timeline Interactive Smithsonian National Museum of Natural History August 2016 Media related to Early modern human at Wikimedia Commons Data related to Early modern human at Wikispecies Portal Evolutionary biology Diamond Jared April 20 2018 A Brand New Version of Our Origin Story The New York Times Retrieved April 23 2018 Retrieved from https en wikipedia org w index php title Early modern human amp oldid 1134871489, wikipedia, wiki, book, books, library,

article

, read, download, free, free download, mp3, video, mp4, 3gp, jpg, jpeg, gif, png, picture, music, song, movie, book, game, games.