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Reconstruction of earlyHomo sapiensfromJebel Irhoud,Moroccoc. 315 000 years BP

Early modern human(EMH), oranatomically modern human(AMH),[1]are terms used to distinguishHomo sapiens(the only extantHomininaspecies) that areanatomicallyconsistent with therange of phenotypesseen in contemporary humans, from extinctarchaic humanspecies. This distinction is useful especially for times and regions where anatomically modern and archaic humans co-existed, for example, inPaleolithic Europe.Among the oldest known remains ofHomo sapiensare those found at theOmo-Kibish Iarchaeological site in south-westernEthiopia,dating to about 233,000[2]to 196,000 years ago,[3]theFlorisbadsite in South Africa, dating to about 259,000 years ago, and theJebel Irhoudsite in Morocco, dated about 315,000 years ago.

Extinct speciesof the genusHomoincludeHomo erectus(extant from roughly 2 to 0.1 million years ago) and a number of other species (by some authors consideredsubspeciesof eitherH. sapiensorH. erectus). The divergence of the lineage leading toH. sapiensout of ancestralH. erectus(or an intermediate species such asHomo antecessor) is estimated to have occurred in Africa roughly 500,000 years ago. The earliest fossil evidence of early modern humans appears inAfricaaround 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]Sustainedarchaic human admixture with modern humansis known to have taken place both in Africa and (following therecent Out-Of-Africa expansion) in Eurasia, between about 100,000 and 30,000 years ago.[9]

Name and taxonomy

Thebinomial nameHomo sapienswas coined byLinnaeus,1758.[10]TheLatinnounhomō(genitivehominis) means "human being", while the participlesapiēnsmeans "discerning, wise, sensible".

The species was initially thought to have emerged from a predecessor within the genusHomoaround 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 ofH. sapienshas been suggested, in whichH. sapienswould by definition refer to the modern human lineage following the split from the Neanderthal lineage. Such a cladistic definition would extend the age ofH. sapiensto over 500,000 years.[note 3]

Estimates for the split between the Homo sapiens line and combinedNeanderthal/Denisovanline 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 intosubspecies,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 theH. rhodesiensisspecies 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 genusHomorather than as subspecies withinH. sapiens.[23]

All humans are considered to be a part of the subspeciesH. 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

Derivation fromH. erectus

The divergence of the lineage that would lead toH. sapiensout ofarchaic humanvarieties derived fromH. erectus,is estimated as having taken place over 500,000 years ago (marking the split of theH. sapienslineage from ancestors shared with other known archaic hominins).[8][5]But the oldest split among modern human populations (such as theKhoisansplit from other groups) has been recently dated to between 350,000 and 260,000 years ago,[25][26]and the earliest known examples ofH. sapiensfossils also date to about that period, including theJebel Irhoudremains from Morocco (ca. 300,000 or 350–280,000 years ago),[27]theFlorisbad Skullfrom South Africa (ca. 259,000 years ago), and theOmo remainsfrom Ethiopia (ca. 195,000, or, as more recently dated, ca. 233,000 years ago).[28][2]

An mtDNA study in 2019 proposed an origin of modern humans in Botswana (and a Khoisan split) of around 200,000 years.[29]However, this proposal has been widely criticized by scholars,[30][31][32]with the recent evidence overall (genetic, fossil, and archaeological) supporting an origin forH. sapiensapproximately 100,000 years earlier and in a broader region of Africa than the study proposes.[32]

In September 2019, scientists proposed that the earliestH. sapiens(and last common human ancestor to modern humans) arose between 350,000 and 260,000 years ago through a merging of populations inEastandSouth Africa.[33][4]

An alternative suggestion definesH. sapienscladisticallyas including the lineage of modern humans since the split from the lineage ofNeanderthals,roughly 500,000 to 800,000 years ago.

The time of divergence between archaicH. sapiensand ancestors of Neanderthals and Denisovans caused by agenetic bottleneckof the latter was dated at 744,000 years ago, combined with repeated early admixture events andDenisovansdiverging from Neanderthals 300 generations after their split fromH. sapiens,as calculated by Rogers et al. (2017).[34]

The derivation of a comparatively homogeneous single species ofH. sapiensfrom more diverse varieties ofarchaic humans(all of which were descended from theearly dispersalofH. erectussome 1.8 million years ago) was debated in terms of two competing models during the 1980s: "recent African origin"postulated the emergence ofH. sapiensfrom a single source population in Africa, which expanded and led to the extinction of all other human varieties, while the "multiregional evolution"model postulated the survival of regional forms of archaic humans, gradually converging into themodern human varietiesby the mechanism ofclinal variation,viagenetic drift,gene flowandselectionthroughout the Pleistocene.[35]

Since the 2000s, the availability of data fromarchaeogeneticsandpopulation geneticshas led to the emergence of a much more detailed picture, intermediate between the two competing scenarios outlined above: Therecent Out-of-Africaexpansion accounts for the predominant part of modern human ancestry, while there were also significantadmixture eventswith regional archaic humans.[36][37]

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 ofH. sapiensin journalistic publications.[38][39][40][41][42]H. s. idaltu,dated to 160,000 years ago, has been postulated as an extinct subspecies ofH. sapiensin 2003.[43][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 theSima de los Huesosfossils published in 2016 seems to suggest thatH. heidelbergensisin 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 ofH. heidelbergensis,to close to 800,000 years ago, the approximate time of disappearance ofH. antecessor.[44][45]

EarlyHomo sapiens

100 to 80 thousand year oldSkhul VfromIsrael

The termMiddle Paleolithicis intended to cover the time between the first emergence ofH. sapiens(roughly 300,000 years ago) and the period held by some to mark the emergence of fullbehavioral modernity(roughly by 50,000 years ago, corresponding to the start of theUpper Paleolithic).

Many of the early modern human finds, like those ofJebel Irhoud,Omo,Herto,Florisbad,Skhul,andPeștera cu Oaseexhibit a mix of archaic and modern traits.[46][47][27]Skhul V, for example, has prominent brow ridges and a projecting face. However, thebrain caseis 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 reflectsmixed ancestryor retention of older traits.[48][49]

The "gracile" or lightly built skeleton of anatomically modern humans has been connected to a change in behavior, including increased cooperation and "resource transport".[50][51]

There is evidence that the characteristic human brain development, especially the prefrontal cortex, was due to "an exceptional acceleration ofmetabolomeevolution... 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. "[52]TheSchöningen spearsand 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. heidelbergensisalready had intellectual and cognitive skills like anticipatory planning, thinking and acting that so far have only been attributed to modern man.[53][54]

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 EveandY-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).[56][57][58]

Fossil teeth found atQesem 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

Dispersal of earlyH. sapiensbegins soon after its emergence, as evidenced by the North AfricanJebel Irhoudfinds (dated to around 315,000 years ago).[27][60]There is indirect evidence forH. sapienspresence in West Asia around 270,000 years ago.[61]

TheFlorisbad Skullfrom Florisbad, South Africa, dated to about 259,000 years ago, has also been classified as representing earlyH. sapiens.[62][63]Scerri (2018),pp. 582–594[4]

In September 2019, scientists proposed that the earliestH. sapiens(and last common human ancestor to modern humans) arose between 350,000 and 260,000 years ago through a merging of populations inEastandSouth Africa.[33][4]

Among extant populations, theKhoi-San(or "Capoid") hunters-gatherers of Southern Africa may represent the human population with the earliest possible divergence within the groupHomo 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 earlyH. 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 archaicH. sapiensin southern Africa represented by, for example, the Florisbad skull dating to 259 (± 35) thousand years ago.[6]

H. s. idaltu,found atMiddle Awashin Ethiopia, lived about 160,000 years ago,[64]andH. sapienslived 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[62]and (more precisely) to 300–270,000 years ago,[4]have been tentatively assigned toH. sapiensand similarities have been noted between them and the Omo Kibbish remains.[62]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 aH. sapiensand 170,000 year old remains of aH. neanderthalensisinApidima Cave,Peloponnese,Greece,more than 150,000 years older than previousH. sapiensfinds in Europe.[69][70][71]

A significant dispersal event, within Africa and to West Asia, is associated with the AfricanmegadroughtsduringMIS 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 ofNamibiaandAngola).[73]

Layer sequence atKsar Akilin theLevantine corridor,and discovery of two fossils ofHomo 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 inSub-Saharan Africabefore 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 fromarchaic 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]In this expansion, bearers ofmt-DNA haplogroup L3left East Africa, likely reaching Arabia via theBab-el-Mandeb,and in theGreat Coastal Migrationspread to South Asia, Maritime South Asia and Oceania between 65,000 and 50,000 years ago,[76][77][78][79]whileEurope,EastandNorth Asiawere reached by about 45,000 years ago. Some evidence suggests that an early wave of humans may have reachedthe Americasby 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 onmitochondrial DNA,combined with evidence based onphysical anthropologyof archaicspecimens,during the 1990s and 2000s,[note 7][81]and has also been supported byY DNAandautosomal DNA.[82]The assumption of complete replacement has been revised in the 2010s with the discovery ofadmixture events(introgression) of populations ofH. sapienswith 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.[83][36]This suggests that interbreeding between Neanderthals and anatomically modern humans took place after therecent "out of Africa" migration,likely between 60,000 and 40,000 years ago.[84][85][86]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 100kya.[87]The extent ofNeanderthal admixture(andintrogressionof 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.[88]The extent of archaic admixture is of the order of about 1% to 4% in Europeans and East Asians, and highest amongMelanesians(the last also havingDenisova homininadmixture at 4% to 6% in addition to neanderthal admixture).[36][48]Cumulatively, about 20% of the Neanderthal genome is estimated to remain present spread in contemporary populations.[89]

In September 2019, scientists reported the computerized determination, based on 260CT scans,of a virtualskull shapeof 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 inEastandSouth AfricawhileNorth-Africanfossils may represent a population which introgressed into Neandertals during the LMP.[33][4]

Anatomy

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 highvariability 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 ofNeanderthalsvs. anatomically modern humans.

Anatomical modernity

The term "anatomically modern humans" (AMH) is used with varying scope depending on context, to distinguish "anatomically modern"Homo sapiensfromarchaic humanssuch as Neanderthals and Middle andLower Paleolithichominins with transitional features intermediate betweenH. erectus,Neanderthals and early AMH calledarchaic Homo sapiens.[90]In a convention popular in the 1990s, Neanderthals were classified as asubspeciesofH. sapiens,asH. s. neanderthalensis,while AMH (orEuropean early modern humans,EEMH) was taken to refer to "Cro-Magnon"orH. s. sapiens.Under this nomenclature (Neanderthals consideredH. sapiens), the term "anatomically modernHomo sapiens"(AMHS) has also been used to refer to EEMH (" Cro-Magnons ").[91]It has since become more common to designate Neanderthals as a separate species,H. neanderthalensis,so that AMH in the European context refers toH. sapiens,but the question is by no means resolved.[note 8]

In this more narrow definition ofH. sapiens,the subspeciesHomo sapiens idaltu,discovered in 2003, also falls under the umbrella of "anatomically modern".[93]The recognition ofH. sapiens idaltuas avalid subspeciesof the anatomically modern human lineage would justify the description of contemporary humans with the subspecies nameHomo sapiens sapiens.[94]However, biological anthropologistChris Stringerdoes not consideridaltudistinct enough withinH. sapiensto warrant its own subspecies designation.[95][62]

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.[96]

Braincase anatomy

Anatomical comparison of skulls ofH. sapiens(left) andH. neanderthalensis(right)
(inCleveland Museum of Natural History)
Features compared are thebraincaseshape,forehead,browridge,nasal bone projection,cheek bone angulation,chinandoccipital contour

The cranium lacks a pronouncedoccipital bunin 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 reducedbrow 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 asupraorbital foramenor notch, forming a groove through the ridge above each eye.[97]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.[98]

Modern humans commonly have a steep, even verticalforeheadwhereas their predecessors had foreheads that sloped strongly backwards.[99]According toDesmond Morris,the vertical forehead in humans plays an important role in human communication througheyebrowmovements and forehead skin wrinkling.[100]

Brain sizein both Neanderthals and AMH is significantly larger on average (but overlapping in range) than brain size inH. 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.[101][note 9]

Jaw anatomy

Compared to archaic people, anatomically modern humans have smaller, differently shaped teeth.[104][105]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 themental trigon,not found in archaic humans.[106]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 (theradius/ulnaandtibia/fibula) are nearly the same size or slightly shorter than the proximal bones (thehumerusandfemur). In ancient people, particularly Neanderthals, the distal bones were shorter, usually thought to be an adaptation to cold climate.[107]The same adaptation is found in some modern people living in the polar regions.[108]

Heightranges 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 averagesrange 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 amongMalay people,for one.[note 11]

Recent evolution

Reconstruction of a modern man from southwestern Europec. 30 000 years BP,London Natural History Museum.

Following thepeopling of Africasome 130,000 years ago, and therecent Out-of-Africaexpansion some 70,000 to 50,000 years ago, some sub-populations ofH. sapienshad been essentiallyisolatedfor tens of thousands of years prior to the early modernAge of Discovery.Combined witharchaic admixturethis has resulted in significantgenetic variation,which in some instances has been shown to be the result ofdirectional selectiontaking place over the past 15,000 years, i.e., significantly later than possible archaic admixture events.[111]

Some climatic adaptations, such ashigh-altitude adaptation in humans,are thought to have been acquired by archaic admixture.Introgressionof genetic variants acquired byNeanderthal admixturehave different distributions inEuropeanandEast 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 toimmuneandhaematopoietic pathways,while European populations showed clustering in functional groups related to thelipid catabolic process.[note 12]A 2017 study found correlation ofNeanderthal admixturein phenotypic traits in modern European populations.[113]

Physiological or phenotypical changes have been traced to Upper Paleolithic mutations, such as the East Asian variant of theEDARgene, dated to c. 35,000 years ago.[note 13]

Recent divergence of Eurasian lineages was sped up significantly during theLast Glacial Maximum(LGM), theMesolithicand theNeolithic,due to increased selection pressures and due tofounder effectsassociated withmigration.[116]Alleles predictive oflight skinhave been found inNeanderthals,[117]but the alleles for light skin in Europeans and East Asians, associated withKITLGandASIP,are (as of 2012) thought to have not been acquired by archaic admixture but recent mutations since the LGM.[116]Phenotypes associated with the "white"or"Caucasian"populations of Western Eurasian stock emerge during the LGM, from about 19,000 years ago. Averagecranial capacityin modern human populations varies in the range of 1,200 to 1,450 cm3for adult males. Larger cranial volume is associated with climatic region, the largest averages being found in populations ofSiberiaand theArctic.[note 14][119]BothNeanderthalandEEMHhad 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.[118]

Examples for still later adaptations related toagricultureandanimal domesticationincludingEast Asiantypes ofADH1Bassociated withrice domestication,[120]orlactase persistence,[121][122]are due to recent selection pressures.

An even more recent adaptation has been proposed for the AustronesianSama-Bajau,developed under selection pressures associated with subsisting onfreedivingover the past thousand years or so.[123][124]

Behavioral modernity

Lithic Industries of earlyHomo sapiensatBlombos Cave(M3 phase, MIS 5), Southern Cape, South Africa (c. 105,000 – 90,000 years old)

Behavioral modernity,involving the development oflanguage,figurative artand early forms ofreligion(etc.) is taken to have arisen before 40,000 years ago, marking the beginning of theUpper Paleolithic(in African contexts also known as theLater Stone Age).[125]

There is considerable debate regarding whether the earliest anatomically modern humans behaved similarly to recent or existing humans.Behavioral modernityis taken to include fully developedlanguage(requiring the capacity forabstract thought),artistic expression,early forms ofreligious behavior,[126]increased cooperation and the formation of early settlements, and the production of articulated tools fromlithic cores,bone or antler. The termUpper Paleolithicis intended to cover the period since therapid expansionof modern humans throughout Eurasia, which coincides with the first appearance ofPaleolithic artsuch ascave paintingsand the development of technological innovation such as thespear-thrower.The Upper Paleolithic begins around 50,000 to 40,000 years ago, and also coincides with the disappearance of archaic humans such as theNeanderthals.

Bifacial silcrete point of earlyHomo sapiens,from M1 phase (71,000 BCE) layer ofBlombos 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 ofH. sapiensaround 200,000 years ago,[127]while others use the term for the rapid developments occurring around 50,000 years ago.[128][129][130]It has been proposed that the emergence of behavioral modernity was a gradual process.[131][132][133][134][135]

Examples of behavioural modernity

Claimed "oldest known drawing by human hands",discovered inBlombos CaveinSouth Africa.Estimated to be a 73,000-year-old work of aHomo sapiens.[136]

The equivalent of the Eurasian Upper Paleolithic in African archaeology is known as theLater 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 theVenus figurinesand other artefacts from theAurignacian,more recent archaeological research has shown that all essential elements of the kind of material culture typical of contemporarySanhunter-gatherers inSouthern Africawas also present by at least 40,000 years ago, including digging sticks of similar materials used today,ostrich eggshell beads, bonearrowheads with individual maker's marks etched and embedded with redochre,and poison applicators.[137]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 atBlombos Cave,South Africa. The technique was used during the final shaping of Still Bay bifacial points made on heat‐treated silcrete. "[138]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 AfricanPinnacle PointSite 5–6. These have been dated to approximately 71,000 years ago. The researchers suggest that their research "shows thatmicrolithic technologyoriginated 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 inAfricawere early and enduring; a small sample of excavated sites in Africa is the best explanation for any perceived 'flickering' pattern. "[139]Increases in behavioral complexity have been speculated to have been linked to an earlier climatic change to much drier conditions between 135,000 and 75,000 years ago.[140]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 inglaciers,such marshlands would have occurred all along the southern coasts of Eurasia. The use ofraftsand boats may well have facilitated exploration of offshore islands and travel along the coast, and eventually permitted expansion to New Guinea and then toAustralia.[141]

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 withgeometricdesigns. Using multiple dating techniques, the site was confirmed to be around 77,000 and 100,000–75,000 years old.[142][143]Ostrich egg shell containers engraved with geometric designs dating to 60,000 years ago were found atDiepkloof,South Africa.[144]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,[145]and shell beads dating to about 75,000 years ago have been found at Blombos Cave, South Africa.[146][147][148]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 asSibudu Cave(along with an early bone needle also found at Sibudu) dating approximately 72,000–60,000 years ago[149][150][151][152][153]some of which may have been tipped with poisons,[154]and bone harpoons at the Central African site of Katanda dating ca. 90,000 years ago.[155]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.[156][139]

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 mi xing and storage for later use.[157][158][159]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.[160]Evidence of early stone-tipped projectile weapons (a characteristic tool ofHomo sapiens), the stone tips of javelins or throwing spears, were discovered in 2013 at the Ethiopian site ofGademotta,and date to around 279,000 years ago.[161]

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.[162]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.[145]

Humans in North Africa (Nazlet Sabaha,Egypt) are known to have dabbled inchertmining,as early as ≈100,000 years ago, for the construction ofstone tools.[163][164]

Evidence was found in 2018, dating to about 320,000 years ago at the site ofOlorgesailiein 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 knownHomo sapiensfossil 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 ofHomo sapiens.[165][166][167]

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.[168]

Pace of progress duringHomo sapienshistory

Homo sapienstechnological and cultural progress appears to have been very much faster in recent millennia than inHomo sapiensearly 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.[169]

Notes

  1. ^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. sapiensdivergence 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 ofH. sapiensis complicated and fraught with controversy. The only widely recognized archaic subspecies[citation needed]isH. sapiens idaltu(2003). The nameH. s. sapiensis due toLinnaeus(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. asiaticusandH. s. ferusfor Africans, Americans, Asians andMalay.This classification remained in common usage until the mid 20th century, sometimes alongsideH. s. tasmanianusfor 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)[55]
  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."[59]
  7. ^"Currently available genetic and archaeological evidence is generally interpreted as supportive of a recent single origin of modern humans in East Africa."[80]
  8. ^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."[92]
  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).[102]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).[103]
  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."[109]
  11. ^Malay, 20–24 (N= m:749 f:893, Median= m:166 cm (5 ft5+12in) f:155 cm (5 ft 1 in), SD= m:6.46 cm (2+12in) f:6.04 cm (2+12in))[110]
  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."[112]
  13. ^Traits affected by the mutation are sweat glands, teeth, hair thickness and breast tissue.[114][115]
  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)[118]

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