انسان هوشمند، انسان خردمند، انسان نوین یا هومو ساپینس (نام علمی: Homo sapiens) نام یک گونه از سردهانسان است. بر اساس نظریه فرگشت، انسان خردمند، نمونهای فرگشت یافته از انسان راستقامت است.
از دیدگاه دیرینمردمشناختی، دو پندار بنیادین در مورد خاستگاه انسان هوشمند موجود است. نخستین دیدگاه بر آن است که انسانهای خردمند، از نیای مشترکی که از انسان راستقامت در آفریقا تمایز یافته، تکثیر گردیدهاست. فرض دیگر بر آن است که انسان خردمند مسیر تکاملی خود را از انسان راستقامت نه فقط در آفریقا که در دیگر قلمروها سپری کردهاست. از سال ۲۰۱۰ میلادی مطالعات ژنتیکی، دیدگاهی میانه را ثابت کردهاند؛ بهطوریکه خزانه ژنی انسان خردمند عمدتاً از نیای آفریقایی است، اگرچه اختلاط در طول تکامل نشان میدهد که در دیگر نقاط کره زمین نیز تکامل رخ دادهاست.
نام علمی انسان خردمند که هومو ساپینس است توسط کارل لینه (۱۷۵۸) گذاشته شده، بخش لاتین آن هومو (homo) به معنی انسان و بشر است و ساپینس (sapiens) که کلمه ای لاتین است، معنی خردمند میدهد. برخی از مترجمان و زبانشناسان معتقدند که عبارت «خردورز» ترجمهی بهتری نسبت به «خردمند» برای کلمهی لاتین sapiens است ولی با این حال در زبان فارسی بیشتر از عبارت خردمند استفاده میشود.
مدل اصلی که مورد قبول اکثریت قرار گرفته منشأ اصلی انسانهای مدرن امروزی را بر اساس مهاجرت انسانهای اولیه از آفریقا دانستهاست. این نظریه در رسانهها با نام مدل خروج-از-آفریقا (Out-of-Africa model) شناخته میشود و در دانشگاهها با نظریه جدید تک-منشأیی (recent single-origin) بودن، نظریه جایگزینی (Replacement Hypothesis) و نظریه مدل جدید منشأ آفریقایی (Recent African Origin) شناخته میشود. این نظریه که انسانها دارای یک منشأ هستند در کتاب «تبار انسان، انتخاب طبیعی در ارتباط با جنیست» توسط چارلز داروین (۱۸۷۱) منتشر شدهاست. این نظریه تا سال ۱۹۸۰ میلادی مورد قبول واقع نشد، تا اینکه در سال ۱۹۸۰ میلادی بر اساس مطالعه DNA روشهای امروزی (دیانای میتوکندریایی) و نیز با شواهد بدست آمده انسانشناسی زیستی مطابقت پیدا کرد. بر اساس شواهد بدست آمده از فسیلها و آزمایشات ژنتیک، انسان خردمند باستانی در آفریقا و به آنتومیک انسان مدرن امروزی در طی سالیان و در حدود ۲۰۰ هزار و ۱۰۰ هزار پیش تکامل یافتند و اولین دستهای که آفریقا را ترک کردند حدود ۶۰ هزار سال پیش بوده که در طی زمان با جمعیت انسانهای نئاندرتال و هومو اریکتوس (انسان راستقامت) جایگزین شدند. اخیراً، در سال ۲۰۱۷ میلادی، فسیلهایی در جبل ایرهود (مراکش) پیدا شدند که نشان میدهند ممکن است تکامل انسان خردمند در حدود ۳۰۰ هزار سال پیش بوده و همچنین دیگر شواهدی وجود دارد که نشان میدهد ممکن است انسان خردمند در ۲۷۰ هزار سال پیش آفریقا را ترک کردهاست. امروزه دانشمندان با مدل تک منشأیی بودن انسانها در آفریقای شرقی اتفاق نظر دارند. با این حال، یافتههای جدید از ژنومهای کامل نئاندرتال این فرض را ایجاد میکند که برخی از انسانهای مدرن امروزی دارای ژنهای مشترکی با برخی ژنهای نئاندرتال هستند. فردی که این مطالعه را انجام داده گفتهاست که یافتههای آنان با ترکیب بالای ۴٪ از نئاندرتال در برخی از جمعیتها مطابقت دارد. اما در همان مطالعه نیز گفته شدهاست که ممکن است دلایل دیگری برای اشتراک ژنهای انسان امروزی با ژنهای باستانی وجود داشته باشد.
بازه زمانی برای تکامل از نسل شامپانزه-انسان به گونه انسان (سرده) تقریباً ۱۰ تا ۲۰ میلیون سال پیش و از انسان راستقامت به انسان خردمند تقریباً ۱٫۸ تا ۰٫۲ میلیون سال پیش تخمین زده شدهاست. مطالعات علمی بر روی فرگشت انسان عمدتاً بر روی ژنوم انسان (سرده) تمرکز دارد (ایجاد و انقراض گونهها)، اما مطالعه بر روی دیگر گونه انسانیان را نیز شامل میشود، برای مثال «میمونهای بزرگ» که شامل جنوبیکپی به عنوان مهمترین اجداد انسانها میشود و همچنین گونههایی که در بین انسانساییان بوده و منقرض شدند: شامپانزهها، بونوبوها، گوریلها و دیگر گونههای منقرض شده. «انسانهای امروزی» به عنوان گونه انسان خردمند (هومو ساپینس) شناخته میشود که تنها زیرگونه منشعب شده از آنچه که هومو ساپینس بوده، است. انسان خردمند ایدالتو که یک گونه شناخته شده دیگر است نیز منقرض شدهاست. گونه انسان نئاندرتال که در ۳۰ هزار سال پیش منقرض شده به عنوان یک زیرگونه از «هومو ساپین نئاندرتال» طبقهبندی شده که مطالعات ژنتیک امروزه نشان دادند که DNA انسانهای امروزی از نئاندرتال در ۵۰۰ هزار سال پیش جدا شدهاست. آناتومی انسانهای امروزی ابتدا در فسیلهای پیدا شده در آفریقا که ۱۹۵ هزار سال قدمت دارند دیده شدهاست (بقایای اومو را ببینید)، که بر اساس شواهد مطالعات زیستشناسی مولکولی میتوان زمان تقریبی جدا شدن اجداد تمام جمعیت انسانها را به ۲۰۰ هزار سال پیش تخمین زد. مطالعات وسیع انجام شده بر تنوع ژنتیکی آفریقاییهای مردم بوشمن و خویسان که نشان دهنده بیشترین تنوع ژنتیکی در بین ۱۱۳ جمعیت مجزای نمونه برداری شده هستند، آنها را جز ۱۴ «خوشههای اجدادی جمعیت» قرار دادهاست. همچنین تحقیقات نشان دادهاست که مهاجرت انسانهای اولیه از جنوب غربی آفریقا نزدیک مرزهای نامیبیا و آنگولا بودهاست. نیروهای انتخاب طبیعی بر روی جمعیت انسان ادامه داشته و شواهد نشان میدهد که در برخی از مناطق ژنومها انتخاب هدایتی در طی ۱۵ هزار سال پیش داشتند.
برای داشتن دید بازتر دربارهی مفهوم خردمندی (خردورزی) در گونهی انسانها بهتر است که کتاب «انسان خردمند» اثر «نوح هراری» مطالعه شود. این کتاب تاریخ را به چهار بخش اصلی انقلاب شناختی، انقلاب کشاورزی، وحدت انسانها و انقلاب علمی مورد مطالعه قرار میدهد. نوح هراری در این کتاب علت پیشرفت انسان خردمند را در همکاری در ابعاد بالا میداند.
Homo sapiens is the only extant human species. The name is Latin for "wiseman" and was introduced in 1758 by Carl Linnaeus (who is himself the lectotype for the species).
Extinct species of the genus Homo include Homo erectus, extant from roughly 1.9 to 0.4 million years ago, and a number of other species (by some authors considered subspecies of either H. sapiens or H. erectus). The age of speciation of H. sapiens out of ancestral H. erectus (or an intermediate species such as Homo antecessor) is estimated to have been roughly 350,000 years ago.[note 1] Sustained archaic admixture is known to have taken place both in Africa and (following the recent Out-Of-Africa expansion) in Eurasia, between about 100,000 and 30,000 years ago.
The species was initially thought to have emerged from a predecessor within the genus Homo around 300,000 to 200,000 years ago.[note 2] A problem with the morphological classification of "anatomically modern" was that it would not have included certain extant populations. For this reason, a lineage-based (cladistic) definition of H. sapiens has been suggested, in which H. sapiens would by definition refer to the modern human lineage following the split from the Neanderthal lineage. Such a cladistic definition would extend the age of H. sapiens to over 500,000 years.[note 3]
Extant human populations have historically been divided into subspecies, but since around the 1980s all extant groups have tended to be subsumed into a single species, H. sapiens, avoiding division into subspecies altogether.[note 4]
Some sources show Neanderthals (H. neanderthalensis) as a subspecies (H. sapiens neanderthalensis). 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.
The subspecies name H. sapiens sapiens is sometimes used informally instead of "modern humans" or "anatomically modern humans". It has no formal authority associated with it.[note 5] By the early 2000s, it had become common to use H. s. sapiens for the ancestral population of all contemporary humans, and as such it is equivalent to the binomial H. sapiens in the more restrictive sense (considering H. neanderthalensis a separate species).[note 6]
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.[note 1] Neanderthals, Denisovans and unspecified archaic African hominins are shown as admixed into the H. sapiens lineage. In addition, prehistoric Archaic Human and Eurasian admixture events in modern African populations are indicated.
The speciation of H. sapiens out of archaic human varieties derived from H. erectus is estimated as having taken place over 350,000 years ago, as the Khoisan split from other populations is dated between 260,000 and 350,000 years ago.
An alternative suggestion defines H. sapienscladistically 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).
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, 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.[better source needed]H. neanderthalensis, which became extinct about 40,000 years ago, has also been classified as a subspecies, H. s. neanderthalensis.
H. heidelbergensis, dated 600,000 to 300,000 years ago, has long been thought to be a likely candidate for the last common ancestor of the Neanderthal and modern human lineages. However, genetic evidence from the Sima de los Huesos fossils published in 2016 seems to suggest that H. heidelbergensis in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "early Neanderthal", while the divergence time between the Neanderthal and modern lineages has been pushed back to before the emergence of H. heidelbergensis, to close to 800,000 years ago, the approximate time of disappearance of H. antecessor.
Many of the early modern human finds, like those of Jebel Irhoud, Omo, Herto, Florisbad, Skhul, and Peștera cu Oase exhibit a mix of archaic and modern traits. 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.[note 7] There have, however, been no reports of the survival of Y-chromosomal or mitochondrial DNA clearly deriving from archaic humans (which would push back the age of the most recent patrilinear or matrilinear ancestor beyond 500,000 years).
Overview map of the peopling of the world by anatomically modern humans (numbers indicate dates in thousands of years ago [ka])
Dispersal of early H. sapiens begins soon after its emergence, as evidenced by the North African Jebel Irhoud finds (dated to between 280,000 and 350,000 years ago). There is indirect evidence for modern human presence in West Asia around 270,000 years ago.
The Florisbad Skull from Florisbad, South Africa, dated to about 260,000 years ago, has been classified as representing a form of H. sapiens.
Among extant populations, the Khoi-San (or "Capoid") hunters-gatherers of Southern Africa may represent the human population with the earliest possible divergence within the group Homo sapiens sapiens. Their separation time has been estimated in a 2017 study to be as long as between 260,000 and 350,000 years ago, compatible with the estimated age of H. sapiens.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 195,000 years ago. Two fossils from Goumde, Kenya, dated to at least 180,000 years ago, have been tentatively assigned to H. sapiens and similarities have been noted between them 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".
Evidence for the overwhelming contribution of this "recent" (L3-derived) expansion to all non-African populations was established based on mitochondrial DNA, combined with evidence based on physical anthropology of archaic specimens, during the 1990s and 2000s.[note 9] 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.
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.
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[note 10]).
In this more narrow definition of H. sapiens, the subspecies H. s. idaltu, discovered in 2003, also falls under the umbrella of "anatomically modern". The recognition of H. s. idaltu as a valid subspecies of the anatomically modern human lineage would justify the description of contemporary humans with the subspecies name H. s. sapiens.
A further division of AMH into "early" or "robust" vs. "post-glacial" or "gracile" subtypes has since been used for convenience. The emergence of "gracile AMH" is taken to reflect a process towards a smaller and more fine-boned skeleton beginning around 50,000–30,000 years ago.
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.[note 11]
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 can be 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.[note 12] By comparison, contemporary national averages range between 158 to 184 cm (62 to 72 in) in males and 147 to 172 cm (58 to 68 in) in females. Neanderthal ranges approximate the height distribution measured among Malay people, for one.[note 13]
Physiological or phenotypical changes have been traced to Upper Paleolithic mutations, such as the East Asian variant of the EDAR gene, dated to c. 35,000 years ago.[note 15]
Recent divergence of Eurasian lineages was sped up significantly during the Last Glacial Maximum, the Mesolithic and the Neolithic, due to increased selection pressures and due to founder effects associated with migration. 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[update]) 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 (adult male averages). Larger cranial volume is associated with climatic region, the largest averages being found in populations of Siberia and the Arctic.[note 16] 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.
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.
Lithic Industries of early Homo sapiens at Blombos Cave (M3 phase, MIS 5), Southern Cape, South Africa (c. 105,000 – 90,000 years old)
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.
In January 2018, it was announced that modern human finds at Misliya cave, Israel, in 2002, had been dated to around 185,000 years ago, the earliest evidence of their out of Africa migration.
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 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. 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 60,000-70,000 years ago, 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 increased 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. 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 has 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.
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 approximately contemporary to 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 80,000-100,000 years ago, in the form of points considered likely to belong to darts delivered by spear throwers.
^ abBased on Schlebusch et al., "Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago",Fig. 3 (H. sapiens divergence times) and Stringer (2012), (archaic admixture).
^This is a matter of convention (rather than a factual dispute), and there is no universal consensus on terminology. Some scholars include humans of up to 600,000 years ago under the same species. See Bryant (2003), p. 811. 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," [...]).
^The history of claimed or proposed subspecies of H. sapiens is complicated and fraught with controversy. The only widely recognized archaic subspecies is H. sapiens idaltu (2003). The name H. s. sapiens is due to Linnaeus (1758), and refers by definition the subspecies of which Linnaeus himself is the type specimen. However, Linnaeus postulated four other extant subspecies, viz. H. s. afer, H. s. americanus, H. s. asiaticus and H. s. ferus for Africans, Americans, Asians and Malay. This classification remained in common usage until the mid 20th century, sometimes alongside H. s. tasmanianus for Australians. See, for example, Bailey, 1946; Hall, 1946. The division of extant human populations into taxonomic subspecies was gradually given up in the 1970s (for example, Grzimek's Animal Life Encyclopedia).
^Homo sapiens sapiens is first used in the 1940s as a synonym of Linnaeus' H. s. europaeus, i. e. Caucasoids. This usage is abandoned by the 1970s, and H. s. sapiens was now used for Cro-Magnon by authors who wished to classify Neanderthals as subspecies of H. sapiens taken in a wider sense, for example Seely.
^For example, "DMA studies have revealed that the first anatomically modern humans (H. s. sapiens) arose in Africa between 200,000 and 140,000 years ago". This usage persists alongside H. s. sapiens designating Upper Paleolithic Cro Magnon, for example. "About 200,000 years ago our own species, Homo sapiens (the thinking human), evolved [...] About 60,000 years ago we became elaborate artisans, building boats and intricate shelters; at this stage, scientists refer to us as Homo sapiens sapiens."
^"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."
^"Currently available genetic and archaeological evidence is generally interpreted as supportive of a recent single origin of modern humans in East Africa."
^It is important to note that this is a question of conventional terminology, not one of a factual disagreement. Pääbo (2014) frames this as a debate that is unresolvable in principle, "since there is no definition of species perfectly describing the case."
^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). 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).
^"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."
^Malay, 20–24 (N= m:749 f:893, Median= m:166 cm (5 ft 51⁄2 in) f:155 cm (5 ft 1 in), SD= m:6.46 cm (21⁄2 in) f:6.04 cm (21⁄2 in))
^"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."
^Traits affected by the mutation are sweat glands, teeth, hair thickness and breast tissue.
^"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)
^Thieme, H (2007). "Der große Wurf von Schöningen: Das neue Bild zur Kultur des frühen Menschen". Die Schöninger Speere – Mensch und Jagd vor 400 000 Jahren. Konrad Theiss Verlag. pp. 224–28. ISBN978-3-89646-040-0.
^Haidle, M.N. (2006). "Menschenaffen? Affenmenschen? Mensch! Kognition und Sprache im Altpaläolithikum". In Conard, N.J. (ed.). Woher kommt der Mensch. Attempto Verlag. pp. 69–97. ISBN3-89308-381-2.
^ Scerri, M.L., et al., "Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter?" Trends in Ecology & Evolution 33(8), August 2018, 582–594, doi:10.1016/j.tree.2018.05.005.
^Posth C, Renaud G, Mittnik M, Drucker DG, Rougier H, Cupillard C, Valentin F, Thevenet C, Furtwängler A, Wißing C, Francken M, Malina M, Bolus M, Lari M, Gigli E, Capecchi G, Crevecoeur I, Beauval C, Flas D, Germonpré M, van der Plicht J, Cottiaux R, Gély B, Ronchitelli A, Wehrberger K, Grigorescu D, Svoboda J, Semal P, Caramelli D, Bocherens H, Harvati K, Conard NJ, Haak W, Powell A, Krause J (2016). "Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe". Current Biology. 26 (6): 827–833. doi:10.1016/j.cub.2016.01.037. PMID26853362.
^Ding, Q.; Hu, Y.; Xu, S.; Wang, J.; Jin, L. (2014) [Online 2013]. "Neanderthal Introgression at Chromosome 3p21.31 was Under Positive Natural Selection in East Asians". Molecular Biology and Evolution. 31 (3): 683–695. doi:10.1093/molbev/mst260. PMID24336922.
^Tattersall, Jeffrey H; Schwartz, Ian (2003). The human fossil record Craniodental Morphology of Genus Homo (Africa and Asia) (vol 2). Wiley-Liss. pp. 327–328. ISBN978-0471319283.
^Steegmann, A. Theodore; Cerny, Frank J.; Holliday, Trenton W. (2002). "Neandertal cold adaptation: Physiological and energetic factors". American Journal of Human Biology. 14 (5): 566–583. doi:10.1002/ajhb.10070. PMID12203812.
^Stock, J.T. (October 2006). "Hunter-gatherer postcranial robusticity relative to patterns of mobility, climatic adaptation, and selection for tissue economy". American Journal of Physical Anthropology. 131 (2): 194–204. doi:10.1002/ajpa.20398. PMID16596600.
^Helmuth H (1998). "Body height, body mass and surface area of the Neanderthals". Zeitschrift für Morphologie und Anthropologie. 82 (1): 1–12. PMID9850627.
^Ingram, Catherine J. E.; Mulcare, Charlotte A.; Itan, Yuval; Thomas, Mark G.; Swallow, Dallas M. (2008-11-26). "Lactose digestion and the evolutionary genetics of lactase persistence". Human Genetics. 124 (6): 579–591. doi:10.1007/s00439-008-0593-6. ISSN0340-6717. PMID19034520.
^Ilardo, M. A.; Moltke, I.; Korneliussen, T. S.; Cheng, J.; Stern, A. J.; Racimo, F.; de Barros Damgaard, P.; Sikora, M.; Seguin-Orlando, A.; Rasmussen, S.; van den Munckhof, I. C. L.; ter Horst, R.; Joosten, L. A. B.; Netea, M. G.; Salingkat, S.; Nielsen, R.; Willerslev, E. (2018-04-18). "Physiological and Genetic Adaptations to Diving in Sea Nomads". Cell. 173 (3): 569–580.e15. doi:10.1016/j.cell.2018.03.054. PMID29677510.
^Gislén, A; Dacke, M; Kröger, RH; Abrahamsson, M; Nilsson, DE; Warrant, EJ (2003). "Superior Underwater Vision in a Human Population of Sea Gypsies". Current Biology. 13 (10): 833–836. doi:10.1016/S0960-9822(03)00290-2. PMID12747831.
^Klein, Richard (1995). "Anatomy, behavior, and modern human origins". Journal of World Prehistory. 9 (2): 167–98. doi:10.1007/bf02221838.
^Shea, John (2011). "Homo sapiens Is As Homo sapiens Was". Current Anthropology. 52 (1): 1–35. doi:10.1086/658067.
^McBrearty, Sally; Brooks, Allison (2000). "The revolution that wasn't: a new interpretation of the origin of modern human behavior". Journal of Human Evolution. 39 (5): 453–563. doi:10.1006/jhev.2000.0435. PMID11102266.
^Henshilwood, Christopher; Marean, Curtis (2003). "The Origin of Modern Human Behavior: Critique of the Models and Their Test Implications". Current Anthropology. 44 (5): 627–651. doi:10.1086/377665.
^Henshilwood, Christopher S.; d'Errico, Francesco; Watts, Ian (2009). "Engraved ochres from the Middle Stone Age levels at Blombos Cave, South Africa". Journal of Human Evolution. 57 (1): 27–47. doi:10.1016/j.jhevol.2009.01.005. PMID19487016.
^Texier PJ, Porraz G, Parkington J, Rigaud JP, Poggenpoel C, Miller C, Tribolo C, Cartwright C, Coudenneau A, Klein R, Steele T, Verna C. (2010). "A Howiesons Poort tradition of engraving ostrich eggshell containers dated to 60,000 years ago at Diepkloof Rock Shelter, South Africa". Proceedings of the National Acadademy of Science U S A. 107: 6180–6185. doi:10.1073/pnas.0913047107PMID20194764
^ abMcBrearty, Sally; Brooks, Allison (2000). "The revolution that wasn't: a new interpretation of the origin of modern human behavior". Journal of Human Evolution. 39 (5): 453–563. doi:10.1006/jhev.2000.0435. PMID11102266.
^d'Errico, Francesco, et al. (2005) Nassarius kraussianus shell beads from Blombos Cave: evidence for symbolic behaviour in the Middle Stone Age. Journal of Human Evolution, 48, 3-24.
^Vanhaeren, Marian, et al. (2013) Thinking strings: Additional evidence for personal ornament use in the Middle Stone Age at Blombos Cave, South Africa. Journal of Human Evolution, 64, 500-517.
^Backwell L, d'Errico F, Wadley L.(2008). Middle Stone Age bone tools from the Howiesons Poort layers, Sibudu Cave, South Africa. Journal of Archaeological Science, 35:1566–1580. doi:10.1016/j.jas.2007.11.006
^Wadley, Lyn (2008). "The Howieson's Poort industry of Sibudu Cave". South African Archaeological Society Goodwin Series. 10.
^Lombard M, Phillips L (2010). "Indications of bow and stone-tipped arrow use 64,000 years ago in KwaZulu-Natal, South Africa". Antiquity. 84 (325): 635–648. doi:10.1017/S0003598X00100134.
^Lombard M (2011). "Quartz-tipped arrows older than 60 ka: further use-trace evidence from Sibudu, Kwa-Zulu-Natal, South Africa". Journal of Archaeological Science. doi:10.1016/j.jas.2011.04.001.
^Backwell L, Bradfield J, Carlson KJ, Jashashvili T, Wadley L, d'Errico F.(2018). The antiquity of bow-and-arrow technology: evidence from Middle Stone Age layers at Sibudu Cave. Journal of Archaeological Science, 92:289-303. doi:10.15184/aqy.2018.11
^Brown, Kyle S.; Marean, Curtis W.; Herries, Andy I.R.; Jacobs, Zenobia; Tribolo, Chantal; Braun, David; Roberts, David L.; Meyer, Michael C.; Bernatchez, J. (14 August 2009), "Fire as an Engineering Tool of Early Modern Humans", Science, 325 (5942): 859–862, Bibcode:2009Sci...325..859B, doi:10.1126/science.1175028, PMID19679810
^Brown, Kyle S.; Marean, Curtis W.; Jacobs, Zenobia; Schoville, Benjamin J.; Oestmo, Simen; Fisher, Erich C.; Bernatchez, Jocelyn; Karkanas, Panagiotis; Matthews, Thalassa (2012). "An early and enduring advanced technology originating 71,000 years ago in South Africa". Nature. 491 (7425): 590–3. Bibcode:2012Natur.491..590B. doi:10.1038/nature11660. PMID23135405.
^Brooks AS, Yellen JE, Potts R, Behrensmeyer AK, Deino AL, Leslie DE, Ambrose SH, Ferguson JR, d'Errico F, Zipkin AM, Whittaker S, Post J, Veatch EG, Foecke K, Clark JB (2018). "Long-distance stone transport and pigment use in the earliest Middle Stone Age". Science. 360 (6384): 90–94. Bibcode:2018Sci...360...90B. doi:10.1126/science.aao2646. PMID29545508.
^Sahle Y, Brooks AS (2018). "Assessment of complex projectiles in the early Late Pleistocene at Aduma, Ethiopia". PLOS ONE. 14 (5): e0216716. doi:10.1371/journal.pone.0216716.