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Saturday, March 12, 2022

Genetic history of the Middle East

From Wikipedia, the free encyclopedia
 

The genetic history of the Middle East is the subject of research within the fields of human population genomics, archaeogenetics and Middle Eastern studies. Researchers use Y-DNA, mtDNA, and other autosomal DNAs to identify the genetic history of ancient and modern populations of Egypt, Persia, Mesopotamia, Anatolia, Arabia, the Levant, and other areas.

History

Developments in DNA sequencing in the 1970s and 1980s provided researchers with the tools needed to study human genetic variation and the genetics of human populations to discover founder populations of modern people groups and human migrations.

In 2005, National Geographic launched The Genographic Project, led by 12 prominent scientists and researchers, to study and map historical human migration patterns by collecting and analyzing DNA samples from hundreds of thousands of people from around the world.

Regions

Egypt

Contamination from handling and intrusion from microbes create obstacles to the recovery of Ancient DNA. Consequently, most DNA studies have been carried out on modern Egyptian populations with the intent of learning about the influences of historical migrations on the population of Egypt.

In general, various DNA studies have found that the genetic variant frequencies of North African populations are intermediate between those of the Near East, the Horn of Africa, southern Europe and Sub Saharan Africa, though Egypt's NRY frequency distributions appear to be much more similar to those of the Middle East than to any sub-Saharan African population, suggesting a much larger Eurasian genetic component in samples examined .

A recent genetic study published in the "European Journal of Human Genetics" (2019) showed that Northern Africans (including Egyptians) are closely related to Europeans and West Asians as well as to Southwest Asians. Northern Africans can clearly be distinguished from West Africans and other African populations dwelling south of the Sahara.

Blood groups

Blood typing and DNA sampling on ancient Egyptian mummies is scant; however, a 1982 study of blood typing of dynastic mummies found ABO frequencies to be most similar to modern Egyptians and some also to Northern Haratin populations. ABO blood group distribution shows that the Egyptians form a sister group to North African populations, including Berbers, Nubians and Canary Islanders.

Ancient Egyptians

In 2013, Nature announced the publication of the first genetic study utilizing next-generation sequencing to ascertain the ancestral lineage of an Ancient Egyptian individual. The research was led by Carsten Pusch of the University of Tübingen in Germany and Rabab Khairat, who released their findings in the Journal of Applied Genetics. DNA was extracted from the heads of five Egyptian mummies that were housed at the institution. All the specimens were dated between 806 BC and 124 AD, a timeframe corresponding with the late Dynastic period. The researchers observed that one of the mummified individuals likely belonged to the mtDNA haplogroup I2, a maternal clade that is believed to have originated in Western Asia.

In a 2017 study published in Nature, three Egyptian mummies were obtained spanning around 1,300 years of Egyptian history from the New Kingdom to the Roman period. Analyses revealed that ancient Egyptians shared more ancestry with Near Easterners than present-day Egyptians, who received additional sub-Saharan admixture in more recent times, around 750 years ago.

Iran

Iranian peoples descended from two closely related West-Eurasian groups, specifically from "Neolithic Iranian farmers" and from "Yamnaya Steppe pastoralists". Recent population genomic studies found that the genetic structure of Iranian peoples formed already about 5,000 years ago and show high continuity since then, suggesting that they were largely unaffected by migration events from outside groups. Genetically speaking, Iranian peoples generally cluster closely with European peoples and populations south to the Caucasus, however certain ethno-linguistic minority groups, such as Turkmens, are more distant and show evidence of recent admixture. Persians, Kurds, Azeris, Tajiks, Pashtuns, Balochis, and certain Uzbek samples, cluster tightly together, forming a single cluster, known as CIC (Central Iranian cluster). Compared with worldwide populations, Iranians (CIC) cluster in the center of the wider West-Eurasian cluster, close to Europeans, Middle Easterners, and South-Central Asians. Interestingly, Iranian Arabs and Azeris are nearly indistinguishable from other Iranian groups, suggesting that linguistic identity is less relevant within Iran. Parsi samples from India were surprisingly found to be part of the Iranian cluster (CIC). The genetic substructure of Iranians is surprisingly low and homogeneous, compared with other "1000G" populations. Europeans, and certain South Asians (Indians) showed the highest affinity with Iranians, while Sub-Saharan Africans and East Asians showed the highest differentiation with Iranians.

Genetic links to neolithic Anatolia

A 2017 study analyzed the autosomal DNA and genome of an Iron Age Iranian sample taken from Teppe Hasanlu (F38_Hasanlu, dated to 971-832 BCE) and revealed it had close affinities to a neolithic North-West Anatolian individual from Kumtepe even closer than Neolithic Iranians.

Gilaks and Mazandaranis

A 2006 genetic research was made by Nasidze et al. on the North Iranian populations on the Gilaks and Mazandaranis, spanning the southwestern coast of the Caspian Sea, up to the border with neighbouring Azerbaijan. The Gilaks and Mazandaranis comprise 7% of the Iranian population. The study suggested that their ancestors came from the Caucasus region, perhaps displacing an earlier group in the South Caspian. Linguistic evidence supports this scenario, in that the Gilaki and Mazandarani languages (but not other Iranian languages) share certain typological features with Caucasian languages, and specifically South Caucasian languages. There have been patterns analyzed of mtDNA and Y chromosome variation in the Gilaki and Mazandarani.

Based on mtDNA HV1 sequences tested by Nasidze et al., the Gilaks and Mazandarani most closely resemble their geographic and linguistic neighbors, namely other Iranian groups. However, their Y chromosome types most closely resemble those found in groups from the South Caucasus. A scenario that explains these differences is a south Caucasian origin for the ancestors of the Gilani and Mazandarani, followed by introgression of women (but not men) from local Iranian groups, possibly because of patrilocality. Given that both mtDNA and language are maternally transmitted, the incorporation of local Iranian women would have resulted in the concomitant replacement of the ancestral Caucasian language and mtDNA types of the Gilani and Mazandarani with their current Iranian language and mtDNA types. Concomitant replacement of language and mtDNA may be a more general phenomenon than previously recognized.

The Mazandarani and Gilani groups fall inside a major cluster consisting of populations from the Caucasus and West Asia and are particularly close to the South Caucasus groups—Georgians, Armenians, and Azerbaijanis. Iranians from Tehran and Isfahan are situated more distantly from these groups.

Iranian Azeris

The 2013 comparative study on the complete mitochondrial DNA diversity in Iranians has indicated that Iranian Azerbaijanis are more related to the people of Georgia, than they are to other Iranians (Like Persians), while the Persians, Armenians and Qashqai on the other hand were more related to each other. It furthermore showed that overall, the complete mtDNA sequence analysis revealed an extremely high level of genetic diversity in the Iranian populations studied which is comparable to the other groups from the South Caucasus, Anatolia and Europe. The same 2013 research further noted that "the results of AMOVA and MDS analyses did not associate any regional and/or linguistic group of populations in the Anatolia, Caucasus and Iran region pointing to strong genetic affinity of Indo-European speaking Persians and Turkic-speaking Qashqais, thus suggesting their origin from a common maternal ancestral gene pool. The pronounced influence of the South Caucasus populations on the maternal diversity of Iranian Azeris is also evident from the MDS analysis results." The study also notes that "It is worth pointing out the position of Azeris from the Caucasus region, who despite their supposed common origin with Iranian Azeris, cluster quite separately and occupy an intermediate position between the Azeris/Georgians and Turks/Iranians grouping". The MtDNA results from the samples overall on average closely resemble those found in the neighbouring regions of the Caucasus, Anatolia, and to a lesser extent (Northern) Mesopotamia.

Among the most common MtDNA lineages in the nation, namely U3b3, appears to be restricted to populations of Iran and the Caucasus, while the sub-cluster U3b1a is common in the whole Near East region.

Iraq

Ancient genetic links to South Asia

A 2013 study based on DNA extracted from the dental remains of four individuals from different time eras (200–300 CE, 2650-2450 BCE, 2200–1900 BCE) unearthed at Tell Ashara (ancient Terqa, in modern Syria) and Tell Masaikh (ancient Kar-Assurnasirpal) suggested a possible genetic link between the people of Bronze Age Mesopotamia and Northern India. According to the study, "We anticipate that the analysed remains from [northern] Mesopotamia belonged to people with genetic affinity to the Indian subcontinent since the distribution of identified ancient haplotypes indicates solid link with populations from the region of South Asia-Tibet (Trans-Himalaya). They may have been descendants of migrants from much earlier times, spreading the clades of the macrohaplogroup M throughout Eurasia and founding regional Mesopotamian groups like that of Terqa or just merchants moving along trade routes passing near or through the region." A 2014 study expanding on the 2013 study and based on analysis of 15751 DNA samples arrives at the conclusion, that "M65a, M49 and/or M61 haplogroups carrying ancient Mesopotamians might have been the merchants from India".

Assyrians

In the 1995 book The History and Geography of Human Genes the authors wrote that: "The Assyrians are a fairly homogeneous group of people, believed to originate from the land of old Assyria in northern Iraq [..] they are Christians and are bona fide descendants of their ancient namesakes." In a 2006 study of the Y chromosome DNA of six regional populations, including, for comparison, Assyrians and Syrians, researchers found that, "the two Semitic populations (Assyrians and Syrians) are very distinct from each other according to both [comparative] axes. This difference supported also by other methods of comparison points out the weak genetic affinity between the two populations with different historical destinies."

A 2008 study on the genetics of "old ethnic groups in Mesopotamia," including 340 subjects from seven ethnic communities ("These populations included Assyrians, Jews, Zoroastrians, Armenians, Arabs and Turkmen (representing ethnic groups from Iran, restricted by rules of their religion), and the Iraqi and Kuwaiti populations from Iraq and Kuwait.") found that Assyrians were homogeneous with respect to all other ethnic groups sampled in the study, regardless of religious affiliation.

Marsh Arabs

A study published in 2011 looking at the relationship between Iraq's Marsh Arabs and ancient Sumerians concluded "the modern Marsh Arabs of Iraq harbour mtDNAs and Y chromosomes that are predominantly of Middle Eastern origin. Therefore, certain cultural features of the area such as water buffalo breeding and rice farming, which were most likely introduced from the Indian sub-continent, only marginally affected the gene pool of the autochthonous people of the region. Moreover, a Middle Eastern ancestral origin of the modern population of the marshes of southern Iraq implies that, if the Marsh Arabs are descendants of the ancient Sumerians, also Sumerians were not of Indian or Southern Asian ancestry." The same 2011 study, when focusing on the genetics of the Maʻdān people of Iraq, identified Y chromosome haplotypes shared by Marsh Arabs, Arabic speaking Iraqis, Assyrians and Mandeans "supporting a common local background."

Levant

Chalcolithic and Bronze Age periods

From a 2020 study published in Cell: "Understanding the nature of this movement was the primary motivation behind this study. Here, we present a large-scale analysis of genome-wide data from key sites of prehistoric Anatolia, the Northern Levant, and the Southern Caucasian lowlands ... In the Northern Levant, we identified a major genetic shift between the Chalcolithic and Bronze Age periods. During this transition, Northern Levantine populations experienced gene flow from new groups harboring ancestries related to both Zagros/Caucasus and the Southern Levant. This suggests a shift in social orientation, perhaps in response to the rise of urban centers in Mesopotamia, which to date remain genetically unsampled." They further add: "This expansion is recorded in the region of the Northern Levant ca. 2800 BCE and could be associated with the movement/ migration of people from Eastern Anatolia and the Southern Caucasian highlands. However, our results do not support this scenario for a number of reasons". "There are extensive textual references from the end of the EBA through the LBA referring to groups of people arriving into the area of the Amuq Valley. Although these groups were named, likely based on designations (e.g., Amorites, Hurrians), the formative context of their (cultural) identity and their geographic origins remain debated. One recent hypothesis (Weiss, 2014, 2017; Akar and Kara, 2020) associates the arrival of these groups with climate-forced population movement during the ‘‘4.2k BP event,’’ a Mega Drought that led to the abandonment of the entire Khabur river valley in Northern Mesopotamia and the search of nearby habitable areas."

The study also suggested a substantial genetic continuity from the Levantine Bronze Age both in modern-day Arabic-speaking Levantine populations (such as Syrians, Druze, Lebanese, and Palestinians) and Jewish groups (such as Moroccan, Ashkenazi, and Mizrahi Jews), who are all suggested to derive a majority (about half or more) of their ancestry from Canaanite-related or Bronze Age Levantine populations (with differering variables for different communities, and with Ashkenazi Jews deriving just over half of their ancestry from Bronze-Age Levantines/Canaanite-related peoples and the rest from Europeans, and Arabic-speaking Levantines, Moroccan Jews, and Mizrahi Jews deriving a larger majority of their ancestry from Bronze Age Canaanite-related peoples). The study concludes that this does not mean that any of these present-day groups bear direct ancestry from people who lived in the Middle-to-Late Bronze Age Levant or in Chalco-lithic Zagros; rather, it indicates that they have ancestries from populations whose ancient proxy can be related to the Middle East.

Canaanites and Phoenicians

Zalloua and Wells (2004), under the auspices of a grant from National Geographic Magazine, examined the origins of the Canaanite Phoenicians. The debate between Wells and Zalloua was whether haplogroup J2 (M172) should be identified as that of the Phoenicians or that of its "parent" haplogroup M89 on the YDNA phylogenetic tree. Initial consensus suggested that J2 be identified with the Canaanite-Phoenician (North Levantine) population, with avenues open for future research. As Wells commented, "The Phoenicians were the Canaanites" It was reported in the PBS description of the National Geographic TV Special on this study entitled "Quest for the Phoenicians" that ancient DNA was included in this study as extracted from the tooth of a 2500-year-old Phoenician mummy.

Wells identified the haplogroup of the Canaanites as haplogroup J2 which originated from Anatolia and the Caucasus. The National Geographic Genographic Project linked haplogroup J2 to the site of Jericho, Tel el-Sultan, ca. 8500 BCE and indicated that in modern populations, haplogroup J2 is found primarily in the Middle East, but also along the coasts of North Africa and Southern Europe, with especially high distribution among present-day Jewish populations (30%), Southern Italians (20%), and lower frequencies in Southern Spain (10%).

Cyprus

Cruciani in 2007 found E1b1b1a2 (E-V13) [one from Sub Clades of E1b1b1a1 (E-M78)] in high levels (>10% of the male population) in Cypriot and Druze lineages. Recent genetic clustering analyses of ethnic groups are consistent with the close ancestral relationship between the Druze and Cypriots, and also identified similarity to the general Syrian and Lebanese populations, as well as a variety of Jewish lineages (Ashkenazi, Sephardi, Iraqi Jewish, and Moroccan Jews).

A 2016 study on 600 Cypriot males asserts that "genome-wide studies indicate that the genetic affinity of Cyprus is nearest to current populations of the Levant". Analyses of Cypriot haplogroup data are consistent with two stages of prehistoric settlement. E-V13 and E-M34 are widespread, and PCA suggests sourcing them to the Balkans and Levant/Anatolia, respectively. Contrasting haplogroups in the PCA were used as surrogates of parental populations. Admixture analyses suggested that the majority of G2a-P15 and R1b-M269 components were contributed by Anatolia and Levant sources, respectively, while Greece/Balkans supplied the majority of E-V13 and J2a-M67. Haplotype-based expansion times were at historical levels suggestive of recent demography. On the other hand, more recent Principal Component Analyses based on autosomal DNA, have placed Cypriots clearly separate from Levantine and Middle Eastern groups, either at the easternmost flank of the south European cluster, or in an intermediate position between southern Europeans and northern Levantines. In a study by Harvard geneticist Iosif Lazarides and colleagues investigating the genetic origins of the Minoans and Mycenaeans, Cypriots were found to be the second least differentiated population from Bronze Age Mycenaeans based on FST index and also genetically differentiated from Levantines.

Palestine and Israel

A study published by the National Academy of Sciences found that "the paternal gene pools of Jewish communities from Europe, North Africa, and the Middle East descended from a common Middle Eastern ancestral population", and suggested that "most Jewish communities have remained relatively isolated from neighbouring non-Jewish communities during and after the Diaspora". Researchers expressed surprise at the remarkable genetic uniformity they found among modern Jews, no matter where the diaspora has become dispersed around the world. Skorecki and colleague wrote that "the extremely close affinity of Jewish and non-Jewish Middle Eastern populations observed ... supports the hypothesis of a common Middle Eastern origin".

This research has suggested that, in addition to Israelite male, significant female founder ancestry might also derive from the Middle East-with 40% of Ashkenazim descended from four women who lived about 2000–3000 years ago in the Middle East. In addition, Behar (2006) suggested that the rest of Ashkenazi mtDNA is originated from about 150 women; most of those were probably of Middle Eastern origin. A 2013 genetic study suggested that the four founding maternal lineages of Ashkenazi Jews originate in Europe and that only ~8% of Ashkenazi mtDNA can confidently be assigned a Near Eastern origin, while >80% of Ashkenazi maternal lineages have a likely European origin (with most Ashkenazi paternal lineages having a Middle Eastern origin), while a 2014 study carried out by Spanish geneticists suggested an ancient Near Eastern origin of the four founding maternal lineages of Ashkenazi Jews.

In 2004, a team of geneticists from Stanford University, the Hebrew University of Jerusalem, Tartu University (Estonia), Barzilai Medical Center (Ashkelon, Israel), and the Assaf Harofeh Medical Center (Zerifin, Israel), studied the modern Samaritan ethnic community living in Israel in comparison with modern Israeli populations to explore the ancient genetic history of these people groups. The Samaritans or Shomronim (singular: Shomroni; Hebrew: שומרוני) trace their origins to the Assyrian province of Shomron (Samaria) in ancient Israel in the period after the Assyrian conquest circa 722 BCE. Shomron was the capital of the Northern Kingdom of Israel when it was conquered by the Assyrians and gave the name to the ancient province of Samaria and the Samaritan people group. Jewish tradition holds that the Samaritans were a mixed group of Israelites who were not exiled or were sent back or returned from exile and non-Israelites relocated to the region by the Assyrians. The modern-day Samaritans are believed to be the direct descendants of the ancient Samaritans.

Their findings reported on four family lineages among the Samaritans: the Tsdaka family (tradition: tribe of Menasseh), the Joshua-Marhiv and Danfi families (tradition: tribe of Ephraim), and the Cohen family (tradition: tribe of Levi). All Samaritan families were found in haplogroups J1 and J2, except the Cohen family which was found in haplogroup E3b1a-M78. This article predated the E3b1a subclades based on the research of Cruciani, et al.

A 2018 study conducted by scholars from Tel-Aviv University, the Israel Antiquities Authority and Harvard University had discovered that 22 out of the 600 people who were buried in Peki'in cave from the Chalcolithic Period were of both local Levantine and Persian and Zagros area ancestries, or as phrased in the paper itself: "Ancient DNA from Chalcolithic Israel reveals the role of population mixture in cultural transformation," the scientists concluded that the homogeneous community found in the cave could source ~57% of its ancestry from groups related to those of the local Levant Neolithic, ~26% from groups related to those of the Anatolian Neolithic, and ~17% from groups related to those of the Iran Chalcolithic. The scholars noted that the Zagros genetic material held "Certain characteristics, such as genetic mutations contributing to blue eye color, were not seen in the DNA test results of earlier Levantine human remains"...The blue-eyed, fair-skinned community didn’t continue, but at least now researchers have an idea why. "These findings suggest that the rise and fall of the Chalcolithic culture are probably due to demographic changes in the region".

In a 2005 study of ASPM gene variants, Mekel-Bobrov et al. found that the Israeli Druze people of the Carmel region have among the highest rate of the newly evolved ASPM haplogroup D, at 52.2% occurrence of the approximately 6,000-year-old allele. While it is not yet known exactly what selective advantage is provided by this gene variant, the haplogroup D allele is thought to be positively selected in populations and to confer some substantial advantage that has caused its frequency to rapidly increase. According to DNA testing, Druze are remarkable for the high frequency (35%) of males who carry the Y-chromosomal haplogroup L, which is otherwise uncommon in the Mideast. This haplogroup originates from prehistoric South Asia and has spread from Pakistan into southern Iran.

Lebanon

In a 2011 genetic study by Haber et al which analyzed the male-line Y-chromosome genetics of the different religious groups of Lebanon, revealed no noticeable or significant genetic differentiation between the Maronites, Greek Orthodox Christians, Greek Catholic Christians, Sunni Muslims, Shiite Muslims, and Druze of the region on the more frequent haplogroups. Major differences between Lebanese groups were found among the less frequent haplogroups. In 1965, Ruffié and Taleb found significant differences of blood markers between ethno-religious groups. A 2005 study by Makhoul et al on Beta Thalassemia Heterogeneity in Lebanon found out that the thalassemia mutations in some Lebanese Christians are similar to the ones observed in Macedonia which "may confirm the presumed Macedonian origin of certain Lebanese Christians".

A 2013 genetic study carried out by Haber at al found "all Jews (Sephardi and Ashkenazi) cluster in one branch; Druze from Mount Lebanon and Druze from Mount Carmel are depicted on a private branch; and Lebanese Christians form a private branch with the Christian populations of Armenia and Cyprus placing the Lebanese Muslims as an outer group. Lebanese Muslims cluster towards the predominant Muslim populations of Syrians, Palestinians, and Jordanians, which in turn cluster on branches with other Muslim populations as distant as Morocco and Yemen."

The authors explained that "In particular, conversion of the region's populations to Islam appears to have introduced major rearrangements in populations' relations through admixture with culturally similar but geographically remote populations, leading to genetic similarities between remarkably distant populations like Jordanians, Moroccans, and Yemenis. Conversely, Christians, Jews and Druze became genetically isolated in the new cultural environment." In conclusions, the authors reconstructed the genetic structure of ancient Levantines and found that a pre-Islamic expansion Levant was more genetically similar to Europeans than to Arabians.

A 2017 study published by the American Journal of Human Genetics, concluded that present-day Lebanese derive most of their ancestry from a Canaanite-related population (Canaanite being a pre-Phoenician name), which therefore implies substantial genetic continuity in the Levant since at least the Bronze Age. More specifically, according to Chris Tyler-Smith, a geneticist and his colleagues at the Sanger Institute in Britain, who compared "sampled ancient DNA from five Canaanite people who lived 3,750 and 3,650 years ago" to modern people. "The comparison revealed that 93 percent of the genetic ancestry of people in Lebanon came from the Canaanites and the other 7 percent was of a Eurasian steppe population"

A 2019 study, carried out by the Wellcome Sanger Institute, United Kingdom, after analyzing the "DNA evidence from the remains of nine Crusaders found at a burial site in Lebanon", concludes that contrary to the popular belief, the Crusaders did not leave "a lasting effect on the genetics of modern-day Lebanese. Instead, today’s Lebanese Christians in particular are more genetically similar to locals from the Roman period, which preceded the Crusades by more than four centuries."

Turkey

Turkish genomic variation, along with several other Western Asian populations, looks most similar to genomic variation of South European populations such as southern Italians. Data from ancient DNA – covering the Paleolithic, the Neolithic, and the Bronze Age periods – showed that Western Asian genomes, including Turkish ones, have been greatly influenced by early agricultural populations in the area; later population movements, such as those of Turkic speakers, also contributed. The first and only (as of 2017) whole genome sequencing study in Turkey was done in 2014. Moreover, the genetic variation of various populations in Central Asia "has been poorly characterized"; Western Asian populations may also be "closely related to populations in the east". An earlier 2011 review had suggested that "small-scale, irregular punctuated migration events" caused changes in language and culture "among Anatolia's diverse autochthonous inhabitants," which explains Anatolian populations' profile today.

Early modern human

From Wikipedia, the free encyclopedia

100 to 80 thousand year old Skhul V from Israel

Early modern human (EMH) or anatomically modern human (AMH) 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 to 196,000 years ago, the Florisbad site in South Africa, dating to about 259,000 years ago, and the Jebel Irhoud site in Morocco, dated about 300,000 years ago.

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. Sustained archaic human admixture with modern humans is known to have taken place both in Africa and (following the recent Out-Of-Africa expansion) in Eurasia, between about 100,000 and 30,000 years ago.

Name and taxonomy

The binomial name Homo sapiens was coined by Linnaeus, 1758. The Latin noun homō (genitive hominis) means "human being", while the participle sapiēns means "discerning, wise, sensible".

The species was initially thought to have emerged from a predecessor within the genus Homo around 300,000 to 200,000 years ago. A problem with the morphological classification of "anatomically modern" was that it would not have included certain extant populations. For this reason, a lineage-based (cladistic) definition of H. sapiens has been suggested, in which H. sapiens would by definition refer to the modern human lineage following the split from the Neanderthal lineage. Such a cladistic definition would extend the age of H. sapiens to over 500,000 years.

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

Extant human populations have historically been divided into subspecies, but since around the 1980s all extant groups have tended to be subsumed into a single species, H. sapiens, avoiding division into subspecies altogether.

Some sources show Neanderthals (H. neanderthalensis) as a subspecies (H. sapiens neanderthalensis). Similarly, the discovered specimens of the H. rhodesiensis species have been classified by some as a subspecies (H. sapiens rhodesiensis), although it remains more common to treat these last two as separate species within the genus Homo rather than as subspecies within H. sapiens.

All humans are considered to be a part of the subspecies H. sapiens sapiens, a designation which has been a matter of debate since a species is usually not given a subspecies category unless there is evidence of multiple distinct subspecies.

Age and speciation process

Derivation from H. erectus

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 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. 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). 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, 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), the Florisbad Skull from South Africa (ca. 259,000 years ago), and the Omo remains from Ethiopia (ca. 195,000, or, as more recently dated, ca. 233,000 years ago).

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

In September 2019, scientists proposed that the earliest H. sapiens (and last common human ancestor to modern humans) arose between 350,000 and 260,000 years ago through a merging of populations in East and South Africa.

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).

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.

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.

Since the 1970s, the Omo remains, originally dated to some 195,000 years ago, have often been taken as the conventional cut-off point for the emergence of "anatomically modern humans". Since the 2000s, the discovery of older remains with comparable characteristics, and the discovery of ongoing hybridization between "modern" and "archaic" populations after the time of the Omo remains, have opened up a renewed debate on the age of H. sapiens in journalistic publications. H. s. idaltu, dated to 160,000 years ago, has been postulated as an extinct subspecies of H. sapiens in 2003. H. neanderthalensis, which became extinct about 40,000 years ago, was also at one point considered to be a subspecies, H. s. neanderthalensis.

H. heidelbergensis, dated 600,000 to 300,000 years ago, has long been thought to be a likely candidate for the last common ancestor of the Neanderthal and modern human lineages. However, genetic evidence from the Sima de los Huesos fossils published in 2016 seems to suggest that H. heidelbergensis in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "early Neanderthal", while the divergence time between the Neanderthal and modern lineages has been pushed back to before the emergence of H. heidelbergensis, to close to 800,000 years ago, the approximate time of disappearance of H. antecessor.

Early Homo sapiens

The term Middle Paleolithic is intended to cover the time between the first emergence of H. sapiens (roughly 300,000 years ago) and the period held by some to mark the emergence of full behavioral modernity (roughly by 50,000 years ago, corresponding to the start of the Upper Paleolithic).

Many of the early modern human finds, like those of Jebel Irhoud, Omo, Herto, Florisbad, Skhul, Red Deer Cave people, and Peștera cu Oase exhibit a mix of archaic and modern traits. Skhul V, for example, has prominent brow ridges and a projecting face. However, the brain case is quite rounded and distinct from that of the Neanderthals and is similar to the brain case of modern humans. It is uncertain whether the robust traits of some of the early modern humans like Skhul V reflects mixed ancestry or retention of older traits.

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

There is evidence that the characteristic human brain development, especially the prefrontal cortex, was due to "an exceptional acceleration of metabolome evolution ... paralleled by a drastic reduction in muscle strength. The observed rapid metabolic changes in brain and muscle, together with the unique human cognitive skills and low muscle performance, might reflect parallel mechanisms in human evolution." The Schöningen spears and their correlation of finds are evidence that complex technological skills already existed 300,000 years ago, and are the first obvious proof of an active (big game) hunt. H. heidelbergensis already had intellectual and cognitive skills like anticipatory planning, thinking and acting that so far have only been attributed to modern man.

The ongoing admixture events within anatomically modern human populations make it difficult to estimate the age of the matrilinear and patrilinear most recent common ancestors of modern populations (Mitochondrial Eve and Y-chromosomal Adam). Estimates of the age of Y-chromosomal Adam have been pushed back significantly with the discovery of an ancient Y-chromosomal lineage in 2013, to likely beyond 300,000 years ago. There have, however, been no reports of the survival of Y-chromosomal or mitochondrial DNA clearly deriving from archaic humans (which would push back the age of the most recent patrilinear or matrilinear ancestor beyond 500,000 years).

Fossil teeth found at Qesem Cave (Israel) and dated to between 400,000 and 200,000 years ago have been compared to the dental material from the younger (120,000–80,000 years ago) Skhul and Qafzeh hominins.

Dispersal and archaic admixture

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). There is indirect evidence for H. sapiens presence in West Asia around 270,000 years ago.

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

In September 2019, scientists proposed that the earliest H. sapiens (and last common human ancestor to modern humans) arose between 350,000 and 260,000 years ago through a merging of populations in East and South Africa.

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.

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

In July 2019, anthropologists reported the discovery of 210,000 year old remains of a H. sapiens and 170,000 year old remains of a H. neanderthalensis in Apidima Cave, Peloponnese, Greece, more than 150,000 years older than previous H. sapiens finds in Europe.

A significant dispersal event, within Africa and to West Asia, is associated with the African megadroughts during MIS 5, beginning 130,000 years ago. A 2011 study located the origin of basal population of contemporary human populations at 130,000 years ago, with the Khoi-San representing an "ancestral population cluster" located in southwestern Africa (near the coastal border of Namibia and Angola).

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

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. 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, 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 40–25,000 years ago.

Evidence for the overwhelming contribution of this "recent" (L3-derived) expansion to all non-African populations was established based on mitochondrial DNA, combined with evidence based on physical anthropology of archaic specimens, during the 1990s and 2000s, and has also been supported by Y DNA and autosomal DNA. 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. This suggests that interbreeding between Neanderthals and anatomically modern humans took place after the recent "out of Africa" migration, likely between 60,000 and 40,000 years ago. Recent admixture analyses have added to the complexity, finding that Eastern Neanderthals derive up to 2% of their ancestry from anatomically modern humans who left Africa some 100 kya. The extent of Neanderthal admixture (and introgression of genes acquired by admixture) varies significantly between contemporary racial groups, being absent in Africans, intermediate in Europeans and highest in East Asians. Certain genes related to UV-light adaptation introgressed from Neanderthals have been found to have been selected for in East Asians specifically from 45,000 years ago until around 5,000 years ago. The extent of archaic admixture is of the order of about 1% to 4% in Europeans and East Asians, and highest among Melanesians (the last also having Denisova hominin admixture at 4% to 6% in addition to neanderthal admixture). Cumulatively, about 20% of the Neanderthal genome is estimated to remain present spread in contemporary populations.

In September 2019, scientists reported the computerized determination, based on 260 CT scans, of a virtual skull shape of the last common human ancestor to modern humans/H. sapiens, representative of the earliest modern humans, and suggested that modern humans arose between 350,000 and 260,000 years ago through a merging of populations in East and South Africa while North-African fossils may represent a population which introgressed into Neandertals during the LMP.

Anatomy

Known archaeological remains of anatomically modern humans in Europe and Africa, directly dated, calibrated carbon dates as of 2013.

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

The term "anatomically modern humans" (AMH) is used with varying scope depending on context, to distinguish "anatomically modern" Homo sapiens from archaic humans such as Neanderthals and Middle and Lower Paleolithic hominins with transitional features intermediate between H. erectus, Neanderthals and early AMH called archaic Homo sapiens. In a convention popular in the 1990s, Neanderthals were classified as a subspecies of H. sapiens, as H. s. neanderthalensis, while AMH (or European early modern humans, EEMH) was taken to refer to "Cro-Magnon" or H. s. sapiens. Under this nomenclature (Neanderthals considered H. sapiens), the term "anatomically modern Homo sapiens" (AMHS) has also been used to refer to EEMH ("Cro-Magnons"). It has since become more common to designate Neanderthals as a separate species, H. neanderthalensis, so that AMH in the European context refers to H. sapiens, but the question is by no means resolved.

In this more narrow definition of H. sapiens, the subspecies Homo sapiens idaltu, discovered in 2003, also falls under the umbrella of "anatomically modern". The recognition of H. sapiens idaltu as a valid subspecies of the anatomically modern human lineage would justify the description of contemporary humans with the subspecies name Homo sapiens sapiens. However, biological anthropologist Chris Stringer does not consider idaltu distinct enough within H. sapiens to warrant its own subspecies designation.

A further division of AMH into "early" or "robust" vs. "post-glacial" or "gracile" subtypes has since been used for convenience. The emergence of "gracile AMH" is taken to reflect a process towards a smaller and more fine-boned skeleton beginning around 50,000–30,000 years ago.

Braincase anatomy

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. This splits the ridge into a central part and two distal parts. In current humans, often only the central section of the ridge is preserved (if it is preserved at all). This contrasts with archaic humans, where the brow ridge is pronounced and unbroken.

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

Brain size in both Neanderthals and AMH is significantly larger on average (but overlapping in range) than brain size in H. erectus. Neanderthal and AMH brain sizes are in the same range, but there are differences in the relative sizes of individual brain areas, with significantly larger visual systems in Neanderthals than in AMH.

Jaw anatomy

Compared to archaic people, anatomically modern humans have smaller, differently shaped teeth. This results in a smaller, more receded dentary, making the rest of the jaw-line stand out, giving an often quite prominent chin. The central part of the mandible forming the chin carries a triangularly shaped area forming the apex of the chin called the mental trigon, not found in archaic humans. Particularly in living populations, the use of fire and tools requires fewer jaw muscles, giving slender, more gracile jaws. Compared to archaic people, modern humans have smaller, lower faces.

Body skeleton structure

The body skeletons of even the earliest and most robustly built modern humans were less robust than those of Neanderthals (and from what little we know from Denisovans), having essentially modern proportions. Particularly regarding the long bones of the limbs, the distal bones (the radius/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. The same adaptation is found in some modern people living in the polar regions.

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. By comparison, contemporary national averages range between 158 to 184 cm (62 to 72 in) in males and 147 to 172 cm (58 to 68 in) in females. Neanderthal ranges approximate the height distribution measured among Malay people, for one.

Recent evolution

Following the peopling of Africa some 130,000 years ago, and the recent Out-of-Africa expansion some 70,000 to 50,000 years ago, some sub-populations of H. sapiens had been essentially isolated 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.

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. A 2017 study found correlation of Neanderthal admixture in phenotypic traits in modern European populations.

Physiological or phenotypical changes have been traced to Upper Paleolithic mutations, such as the East Asian variant of the EDAR gene, dated to c. 35,000 years ago.

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. Alleles predictive of light skin have been found in Neanderthals, 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. 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. 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.

Examples for still later adaptations related to agriculture and animal domestication including East Asian types of ADH1B associated with rice domestication, or lactase persistence, are due to recent selection pressures.

An even more recent adaptation has been proposed for the Austronesian Sama-Bajau, developed under selection pressures associated with subsisting on freediving over the past thousand years or so.

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).

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, 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, while others use the term for the rapid developments occurring around 50,000 years ago. It has been proposed that the emergence of behavioral modernity was a gradual process.

Examples of behavioural modernity

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.

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. There is also a suggestion that "pressure flaking best explains the morphology of lithic artifacts recovered from the c. 75-ka Middle Stone Age levels at Blombos Cave, South Africa. The technique was used during the final shaping of Still Bay bifacial points made on heat‐treated silcrete." Both pressure flaking and heat treatment of materials were previously thought to have occurred much later in prehistory, and both indicate a behaviourally modern sophistication in the use of natural materials. Further reports of research on cave sites along the southern African coast indicate that "the debate as to when cultural and cognitive characteristics typical of modern humans first appeared" may be coming to an end, as "advanced technologies with elaborate chains of production" which "often demand high-fidelity transmission and thus language" have been found at the South African Pinnacle Point Site 5–6. These have been dated to approximately 71,000 years ago. The researchers suggest that their research "shows that microlithic technology originated early in South Africa by 71 kya, evolved over a vast time span (c. 11,000 years), and was typically coupled to complex heat treatment that persisted for nearly 100,000 years. Advanced technologies in Africa were early and enduring; a small sample of excavated sites in Africa is the best explanation for any perceived 'flickering' pattern." These results suggest that Late Stone Age foragers in Sub-Saharan Africa had developed modern cognition and behaviour by at least 50,000 years ago. The change in behavior has been speculated to have been a consequence of an earlier climatic change to much drier and colder conditions between 135,000 and 75,000 years ago. 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.

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 100–75,000 years old. Ostrich egg shell containers engraved with geometric designs dating to 60,000 years ago were found at Diepkloof, South Africa. Beads and other personal ornamentation have been found from Morocco which might be as much as 130,000 years old; as well, the Cave of Hearths in South Africa has yielded a number of beads dating from significantly prior to 50,000 years ago, and shell beads dating to about 75,000 years ago have been found at Blombos Cave, South Africa. Specialized projectile weapons as well have been found at various sites in Middle Stone Age Africa, including bone and stone arrowheads at South African sites such as Sibudu Cave (along with an early bone needle also found at Sibudu) dating approximately 72,000–60,000 years ago some of which may have been tipped with poisons, and bone harpoons at the Central African site of Katanda dating ca. 90,000 years ago. Evidence also exists for the systematic heat treating of silcrete stone to increase its flake-ability for the purpose of toolmaking, beginning approximately 164,000 years ago at the South African site of Pinnacle Point and becoming common there for the creation of microlithic tools at about 72,000 years ago.

In 2008, an ochre processing workshop likely for the production of paints was uncovered dating to ca. 100,000 years ago at Blombos Cave, South Africa. Analysis shows that a liquefied pigment-rich mixture was produced and stored in the two abalone shells, and that ochre, bone, charcoal, grindstones and hammer-stones also formed a composite part of the toolkits. Evidence for the complexity of the task includes procuring and combining raw materials from various sources (implying they had a mental template of the process they would follow), possibly using pyrotechnology to facilitate fat extraction from bone, using a probable recipe to produce the compound, and the use of shell containers for mixing and storage for later use. Modern behaviors, such as the making of shell beads, bone tools and arrows, and the use of ochre pigment, are evident at a Kenyan site by 78,000-67,000 years ago. Evidence of early stone-tipped projectile weapons (a characteristic tool of Homo sapiens), the stone tips of javelins or throwing spears, were discovered in 2013 at the Ethiopian site of Gademotta, and date to around 279,000 years ago.

Expanding subsistence strategies beyond big-game hunting and the consequential diversity in tool types have been noted as signs of behavioral modernity. A number of South African sites have shown an early reliance on aquatic resources from fish to shellfish. Pinnacle Point, in particular, shows exploitation of marine resources as early as 120,000 years ago, perhaps in response to more arid conditions inland. Establishing a reliance on predictable shellfish deposits, for example, could reduce mobility and facilitate complex social systems and symbolic behavior. Blombos Cave and Site 440 in Sudan both show evidence of fishing as well. Taphonomic change in fish skeletons from Blombos Cave have been interpreted as capture of live fish, clearly an intentional human behavior.

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

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.

In 2019, further evidence of Middle Stone Age complex projectile weapons in Africa was found at Aduma, Ethiopia, dated 100,000–80,000 years ago, in the form of points considered likely to belong to darts delivered by spear throwers.

Pace of progress during Homo sapiens history

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

Operator (computer programming)

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