Extinct species of the genus Homo include Homo erectus, extant during 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. 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 term anatomically modern humans (AMH) is used to distinguish H. sapiens having an anatomy consistent with the range of phenotypes seen in contemporary humans from varieties of extinct archaic humans. This is useful especially for times and regions where anatomically modern and archaic humans co-existed, for example, in Paleolithic Europe.
|Male and female H s. sapiens|
(Akha in northern Thailand,
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.
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.
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. 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).
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 and West-Central Africans have partial ancestry from an archaic H. sapiens population (termed "Basal Human" in the study) whose divergence predates even Khoisan to a significant extent and did not share much genetic drift with other Africans before admixing into them.
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, 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 cast into terms of "H. sapiens may be older than previously thought". 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, 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.
The term Middle Paleolithic is intended to cover the time between the first emergence of H. sapiens (roughly 300,000 years ago) and the emergence of full behavioral modernity (roughly 50,000 years ago).
Many of the early modern human finds, like those of Omo, Herto, Skhul, Jebel Irhoud 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 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 and Dali Man from China is dated at 260,000 years ago.
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 site Middle Awash in Ethiopia, lived about 160,000 years ago., and H. Sapiens lived at Omo Kibish in Ethiopia about 195,000 years ago.  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.
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).
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. Asia was re-populated by early modern humans in the so-called "recent out-of-Africa migration" post-dating MIS5, beginning around 70,000 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 by 65,000 years ago, while Europe, East and North Asia, and possibly the Americas, were reached by 50,000 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. 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, Papuan New Guineans, and Australian Aboriginals. 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 (Denisova hominin admixture), at 4% to 6%. Cumulatively, about 20% of the Neanderthal genome is estimated to remain present spread in contemporary populations.
In November 2018, scientists reported that nearly all extant populations of animals, including humans, may be a result of a population expansion that began between one and two hundred thousand years ago, based on genetic mitochondrial DNA studies.
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).
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.
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 require fewer jaw muscles, giving slender, more gracile jaws. Compared to archaic people, modern humans have smaller, lower faces.
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. 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 approximating the height distribution measured, e.g., among Malay people.
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 have 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.
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) 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. 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.
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.
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 earliest H. sapiens (AMH) found in Europe are the "Cro-Magnon" (named after the site of first discovery in France), beginning about 40,000 to 35,000 years ago. These are also known as "European early modern humans" in contrast to the preceding Neanderthals.
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 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, 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.
Currently available genetic and archaeological evidence is generally interpreted as supportive of a recent single origin of modern humans in East Africa."Out of Africa Revisited". Science. 308 (5724): 921g. 2005-05-13. doi:10.1126/science.308.5724.921g.
The term archaic Homo sapiens has different meanings depending on the preferred system of taxonomy.
See Human taxonomy for the question of taxonomic classification of early human varieties.
Archaic Homo sapiens may refer to:
early forms of anatomically modern humans
transitional forms of archaic humans possessing some of the derived traits of modern humansArchaic humans
A number of varieties of Homo are grouped into the broad category of archaic humans in the period contemporary to and predating the emergence of the earliest anatomically modern humans (Homo sapiens) over 315 ka. The term typically includes Homo neanderthalensis (430+–25 ka), Denisovans, Homo rhodesiensis (300–125 ka), Homo heidelbergensis (600–200 ka), and Homo antecessor.
There is no universal consensus on this terminology, and varieties of "archaic humans" are included under the binomial name of either Homo sapiens or Homo erectus by some authors.
Archaic humans had a brain size averaging 1,200 to 1,400 cubic centimeters, which overlaps with the range of modern humans. Archaics are distinguished from anatomically modern humans by having a thick skull, prominent supraorbital ridges (brow ridges) and the lack of a prominent chin.Anatomically modern humans appear from over 160,000 years ago in Ethiopia and after 70,000 years ago (see Toba catastrophe theory), gradually supplanting the "archaic" human varieties. Non-modern varieties of Homo are certain to have survived until after 30,000 years ago, and perhaps until as recently as 12,000 years ago. Which of these, if any, are included under the term "archaic human" is a matter of definition and varies among authors. Nonetheless, according to recent genetic studies, modern humans may have bred with "at least two groups" of ancient humans: Neanderthals and Denisovans. Other studies have cast doubt on admixture being the source of the shared genetic markers between archaic and modern humans, pointing to an ancestral origin of the traits which originated 500,000–800,000 years ago.Another group may also have been extant as recently as 11,500 years ago, the Red Deer Cave people of China. Chris Stringer of the Natural History Museum in London has suggested that these people could be a result of mating between Denisovans and modern humans. Other scientists remain skeptical, suggesting that the unique features are within the variations expected for modern human populations.Denisovan
The Denisovans or Denisova hominins ( di-NEE-sə-və) are an extinct species or subspecies of archaic humans in the genus Homo. Pending its taxonomic status, it currently carries temporary species or subspecies names Homo denisova, Homo altaiensis, Homo sapiens denisova, or Homo sapiens Altai. In March 2010, scientists announced the discovery of an undated finger bone fragment of a juvenile female found in the Denisova Cave in the Altai Mountains in Siberia, a cave that has also been inhabited by Neanderthals and modern humans.
The mitochondrial DNA (mtDNA) of the finger bone showed it to be genetically distinct from Neanderthals and modern humans. The nuclear genome from this specimen suggested that Denisovans shared a common origin with Neanderthals, that they ranged from Siberia to Southeast Asia, and that they lived among and interbred with the ancestors of some modern humans, with about 3% to 5% of the DNA of Melanesians and Aboriginal Australians and around 6% in Papuans deriving from Denisovans.A 2013 comparison with the genome of another Neanderthal from the Denisova cave revealed local interbreeding with local Neanderthal DNA representing 17% of the Denisovan genome, and evidence of interbreeding with an as yet unidentified ancient human lineage. Analysis of DNA from two teeth found in layers different from the finger bone revealed an unexpected degree of mtDNA divergence among Denisovans. Two teeth belonging to different members of the Denisova cave population have been reported. In November 2015, a tooth fossil containing DNA was reported to have been found and studied.The lineage that developed into Denisovans and Neanderthals is estimated to have separated from the lineage that developed into "anatomically modern" Homo sapiens approximately 600,000 to 744,000 years ago. Denisovans and Neanderthals then significantly diverged from each other genetically a mere 300 generations after that. In January 2019, scientists reported that several types of humans, including Denisovans, Neanderthals and related hybrids, may have each dwelt in the Denisova Cave in Siberia over thousands of years, but it is unclear whether they ever co-habitated the cave.Grotte du Bichon
Grotte du Bichon is a karstic cave in the Swiss Jura, overlooking the Doubs river at an altitude of 846 m, some 5 km north of La Chaux-de-Fonds.
It is the site of the discovery of the skeleton a hunter-gatherer of the Azilian (late Upper Paleolithic to early Mesolithic), dubbed "Bichon man" (homme de Bichon), a young male about 20 to 23 years old, carbon dated to 13,770–13,560 years ago (95% CI).
The skeleton was discovered in 1956, about 15 m from the cave entrance, intermingled with the bones of a female brown bear, nine flint arrowheads and traces of charcoal.
In 1991, flint chips were found embedded in the bear's third vertebra, without indication of healing, suggesting the interpretation that the bear was wounded by arrows, retreated into the cave, and was pursued by the hunter, who made a fire to fumigate the bear from the cave, but was killed by the dying animal.A genetic analysis of the man showed membership in the "West European Hunter-Gatherer" lineage known from younger fossils of the European Mesolithic.
He was a bearer of Y-DNA haplogroup I2a and of mt-DNA haplogroup U5b1h.
Y-DNA haplogroup I2a probably arose in Europe prior to the Last Glacial Maximum.
Morphologically, his skull was described as relatively long, with a low face and subrectangular
eye-sockets. He would have weighed just above 60 kg at a height of 1.64 m.
He was relatively slender, but muscular (based on muscle attachments visible on the skeleton), with a pronounced lateral asymmetry suggesting right-handedness.
Study of carbon and nitrogen fractionations suggests a largely meat-based diet.Homo heidelbergensis
Homo heidelbergensis is an extinct species or subspecies of archaic humans in the genus homo, which radiated in the Middle Pleistocene from about 700,000 to 300,000 years ago, known from fossils found in Southern Africa, East Africa and Europe. African H. heidelbergensis has several subspecies. The subspecies are Homo heidelbergensis heidelbergensis, Homo heidelbergensis daliensis, Homo rhodesiensis, and Homo heidelbergensis steinheimensi. The derivation of Homo sapiens from Homo rhodesiensis has often been proposed, but is obscured by a fossil gap from 400–260 kya. The species was originally named Homo heidelbergensis due to the skeleton's first discovery in Heidelberg, Germany.The first discovery—a mandible—was made in 1907 by Otto Schoetensack. The skulls of this species share features with both Homo erectus and the anatomically modern Homo sapiens; its brain was nearly as large as that of Homo sapiens. The Sima de los Huesos cave at Atapuerca in northern Spain holds particularly rich layers of deposits where excavations were still in progress as of 2018.H. heidelbergensis was dispersed throughout Eastern and Southern Africa (Ethiopia, Namibia, Southern Africa) as well as Europe (England, France, Germany, Greece, Hungary, Italy, Portugal, Spain). Its exact relation both to the earlier Homo antecessor and Homo ergaster, and to the later lineages of Neanderthals, Denisovans, and modern humans is unclear.Homo sapiens has been proposed as derived from H. heidelbergensis via Homo rhodesiensis, present in East and North Africa from around 400,000 years ago.
The correct assignment of many fossils to a particular chronospecies is difficult and often differences in opinion ensue among paleoanthropologists due to the absence of universally accepted dividing lines (autapomorphies) between Homo erectus, Homo heidelbergensis, Homo rhodesiensis and Neanderthals.
It is uncertain whether H. heidelbergensis is ancestral to Homo sapiens, as a fossil gap in Africa between 400,000 and 260,000 years ago obscures the presumed derivation of H. sapiens from H. rhodesiensis.
Genetic analysis of the Sima de los Huesos fossils (Meyer et al. 2016) seems to suggest that H. heidelbergensis in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "archaic 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 about 600,000 to 800,000 years ago, the approximate time of disappearance of Homo antecessor.The delineation between early H. heidelbergensis and H. erectus is also unclear. Given the evidence, it means there is no direct evidence that suggest the Homo heidelbergensis is related to modern-day humans.Homo rhodesiensis
Homo rhodesiensis is the species name proposed by Arthur Smith Woodward (1921) to classify Kabwe 1 (the "Kabwe skull" or "Broken Hill skull", also "Rhodesian Man"), a Middle Stone Age fossil recovered from a cave at Broken Hill, or Kabwe, Northern Rhodesia (now Zambia).H. rhodesiensis is now mostly considered a synonym of Homo heidelbergensis, or possibly an African subspecies of Homo heidelbergensis sensu lato, understood as a polymorphic species dispersed throughout Africa and Eurasia with a range spanning the Middle Pleistocene (c. 0.8–0.12 mya).
Other designations such as Homo sapiens arcaicus and Homo sapiens rhodesiensis have also been proposed. White et al. (2003) suggested Rhodesian Man as ancestral to Homo sapiens idaltu (Herto Man).The derivation of Homo sapiens from Homo rhodesiensis has often been proposed, but is obscured by a fossil gap during 400–260 kya.Homo sapiens idaltu
Homo sapiens idaltu (Afar: Idaltu; "elder" or "first born"), also called Herto Man, is the name given to a number of early modern human fossils found in 1997 in Herto Bouri, Ethiopia. They date to around 160,000 years ago.Paleoanthropologists determined that the skeletal finds belong to an extinct subspecies of Homo sapiens who lived in Pleistocene Africa. In the narrow definition of H. sapiens, the subspecies H. s. idaltu falls under the umbrella of Anatomically modern humans. 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. Because of their early dating and unique physical characteristics, they represent the immediate ancestors of anatomically modern humans, as suggested by the Out-of-Africa theory.Human evolution
Human evolution is the evolutionary process that led to the emergence of anatomically modern humans, beginning with the evolutionary history of primates—in particular genus Homo—and leading to the emergence of Homo sapiens as a distinct species of the hominid family, the great apes. This process involved the gradual development of traits such as human bipedalism and language, as well as interbreeding with other hominins, which indicate that human evolution was not linear but a web.The study of human evolution involves several scientific disciplines, including physical anthropology, primatology, archaeology, paleontology, neurobiology, ethology, linguistics, evolutionary psychology, embryology and genetics. Genetic studies show that primates diverged from other mammals about 85 million years ago, in the Late Cretaceous period, and the earliest fossils appear in the Paleocene, around 55 million years ago.Within the Hominoidea (apes) superfamily, the Hominidae family diverged from the Hylobatidae (gibbon) family some 15–20 million years ago; African great apes (subfamily Homininae) diverged from orangutans (Ponginae) about 14 million years ago; the Hominini tribe (humans, Australopithecines and other extinct biped genera, and chimpanzee) parted from the Gorillini tribe (gorillas) between 8–9 million years ago; and, in turn, the subtribes Hominina (humans and biped ancestors) and Panina (chimps) separated 4–7.5 million years ago.Human taxonomy
Human taxonomy is the classification of the human species (systematic name Homo sapiens, Latin: "knowing man") within zoological taxonomy. The systematic genus, Homo, is designed to include both anatomically modern humans and extinct varieties of archaic humans. Current humans have been designated as subspecies Homo sapiens sapiens, differentiated from the direct ancestor, Homo sapiens idaltu.
Since the introduction of systematic names in the 18th century, knowledge of human evolution has increased drastically, and a number of intermediate taxa have been proposed in the 20th to early 21st century. The most widely accepted taxonomy groups takes the genus Homo as originating between two and three million years ago, divided into at least two species, archaic Homo erectus and modern Homo sapiens, with about a dozen further suggestions for species without universal recognition.
The genus Homo is placed in the tribe Hominini alongside Pan (chimpanzees). The two genera are estimated to have diverged over an extended time of hybridization spanning roughly 10 to 6 million years ago, with possible admixture as late as 4 million years ago. A subtribe of uncertain validity, grouping archaic "pre-human" or "para-human" species younger than the Homo-Pan split is Australopithecina (proposed in 1939).
A proposal by Wood and Richmond (2000) would introduce Hominina as a subtribe alongside Australopithecina, with Homo the only known genus within Hominina. Alternatively, following Cela-Conde and Ayala (2003), the "pre-human" or "proto-human" genera of Australopithecus, Ardipithecus, Praeanthropus, and possibly Sahelanthropus may be placed on equal footing alongside the genus Homo. An even more radical view rejects the division of Pan and Homo as separate genera, which based on the Principle of Priority would imply the re-classification of chimpanzees as Homo paniscus (or similar).Prior to the current scientific classification of humans, philosophers and scientists have made various attempts to classify humans. They offered definitions of the human being and schemes for classifying types of humans. Biologists once classified races as subspecies, but today anthropologists reject the concept of race and view humanity as an interrelated genetic continuum. Taxonomy of the hominins continues to evolve.Jebel Irhoud
Jebel Irhoud (Moroccan Arabic: جبل إيغود, translit. žbəl iġud pronounced [ˈd͡ʒabal ˈʔiːɣuːd]) is an archaeological site located just north of the locality known as Tlet Ighoud, about 50 km (30 mi) south-east of the city of Safi in Morocco. It is noted for the hominin fossils that have been found there since the site's discovery in 1960. Originally thought to be Neanderthals, the specimens have since been assigned to Homo sapiens and have been dated at 315,000 years old.List of human evolution fossils
The following tables give an overview of notable finds of hominin fossils and remains relating to human evolution, beginning with the formation of the Hominini tribe (the divergence of the human and chimpanzee lineages) in the late Miocene, roughly 7 to 8 million years ago.
As there are thousands of fossils, mostly fragmentary, often consisting of single bones or isolated teeth with complete skulls and skeletons rare, this overview is not complete, but does show some of the most important finds. The fossils are arranged by approximate age as determined by radiometric dating and/or incremental dating and the species name represents current consensus; if there is no clear scientific consensus the other possible classifications are indicated.
Most of the early fossils shown are not considered direct ancestors to Homo sapiens but are closely related to direct ancestors and are therefore important to the study of the lineage. After 1.5 million years ago (extinction of Paranthropus), all fossils shown are human (genus Homo). After 11,500 years ago (11.5 ka, beginning of the Holocene), all fossils shown are Homo sapiens (anatomically modern humans), illustrating recent divergence in the formation of modern human sub-populations.Loschbour man
The Loschbour man (also Loschbur man) is a skeleton of Homo sapiens from the European Mesolithic discovered in 1935 in Mullerthal, in the commune of Waldbillig, Luxembourg.Love, Simon
Love, Simon is a 2018 American romantic teen comedy-drama film directed by Greg Berlanti, written by Isaac Aptaker and Elizabeth Berger, and based on the novel Simon vs. the Homo Sapiens Agenda by Becky Albertalli. The film stars Nick Robinson, Josh Duhamel, and Jennifer Garner. It centers on Simon Spier, a closeted gay high school boy who is forced to balance his friends, his family, and the blackmailer threatening to out him to the entire school, while simultaneously attempting to discover the identity of the anonymous classmate with whom he has fallen in love online.
Love, Simon premiered at the Mardi Gras Film Festival on February 27, 2018, and was released in the United States on March 16, 2018, by 20th Century Fox. Critics praised the film for its "big heart, diverse and talented cast, and revolutionary normalcy", describing it as "tender, sweet, and affecting" and a "hugely charming crowd-pleaser" that is "funny, warm-hearted and life-affirming", with reviews comparing it to the romantic comedy-drama films of John Hughes. It is notable as the first film by a major Hollywood studio to focus on a gay teenage romance, it grossed $66 million worldwide.Manot 1
Manot 1 is a fossil specimen designated to a skullcap that represents an archaic modern human discovered in Manot Cave, Western Galilee, Israel.
It was discovered in 2008 and the scientific description was published in 2015. Radiometric dating indicates that it is about 54,700 years old (the late Mousterian), and thought to be directly ancestral to the
Upper Paleolithic populations of the Levant and Europe.Misliya cave
Misliya cave (Hebrew: מערת מיסליה, also known as Brotzen Cave after Fritz Brotzen who first described it in 1927) at Mount Carmel, Israel, is a collapsed cave containing archaeological layers from the Lower Paleolithic and Middle Paleolithic periods. The site is significant in paleoanthropology for the discovery of the earliest known remains attributed to Homo sapiens outside of Africa, dated to 185,000 years ago.Excavations by teams of University of Haifa and University of Tel Aviv were conducted in the 2000/1 season, yielding finds dated to between 300,000 and 150,000 years ago.
Of special interest is the Misliya-1 fossil, an upper jawbone discovered in 2002 and at first dated to "possibly 150,000 years ago" and classified as "early modern Homo sapiens" (EMHS).
In January 2018, the date of the fossil has been revised to between 177,000 and 194,000 years ago (95% CI).
This qualifies Misliya-1 as one of the oldest known fossil of H. sapiens, of comparable age to the Omo remains (identified as "archaic Homo sapiens", or Homo sapiens idaltu), and the oldest modern humans ever found outside of Africa.Mutant (Marvel Comics)
In American comic books published by Marvel Comics, a mutant is a human being that possesses a genetic trait called the X-gene. It causes the mutant to develop superhuman powers that manifest at puberty. Human mutants are sometimes referred to as a human subspecies Homo sapiens superior, or simply Homo superior. Mutants are the evolutionary progeny of Homo sapiens, and are generally assumed to be the next stage in human evolution. The accuracy of this is the subject of much debate in the Marvel Universe.
Unlike Marvel's mutates, which are characters who develop their powers only after exposure to outside stimuli or energies (such as the Hulk, Spider-Man, The Fantastic Four, and Absorbing Man), mutants have actual genetic mutations.Neanderthal
Neanderthals (UK: , also US: ;) (Homo neanderthalensis or Homo sapiens neanderthalensis) are an extinct species or subspecies of archaic humans in the genus Homo, who lived within Eurasia from circa 400,000 until 40,000 years ago.Currently the earliest fossils of Neanderthals in Europe are dated between 450,000 to 430,000 years ago, and thereafter Neanderthals expanded into Southwest and Central Asia. They are known from numerous fossils, as well as stone tool assemblages. Almost all assemblages younger than 160,000 years are of the so-called Mousterian techno-complex, which is characterised by tools made out of stone flakes.
The type specimen is Neanderthal 1, found in Neander Valley in the German Rhineland, in 1856.
Compared to modern humans, Neanderthals were stockier, with shorter legs and bigger bodies. In conformance with Bergmann's rule, this likely was an adaptation to preserve heat in cold climates. Male and female Neanderthals had cranial capacities averaging 1,600 cm3 (98 cu in) and 1,300 cm3 (79 cu in), respectively,
within the range of the values for anatomically modern humans.
Average males stood around 164 to 168 cm (65 to 66 in) and females 152 to 156 cm (60 to 61 in) tall.Since 2010, there has been growing evidence for admixture between Neanderthals and anatomically modern humans. This admixture is reflected in the genomes of modern European, Asian, Papua New Guinean, Australian Aboriginal, Native American and other non-African populations, but not in the genomes of most sub-Saharan 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 100,000 years ago.Solo Man
Solo Man (Homo erectus soloensis) is a subspecies of Homo erectus., identified based on fossil evidence discovered between 1931 and 1933 by Gustav Heinrich Ralph von Koenigswald, from sites along the Solo River, on the Indonesian island of Java, dated to between 550,000 and 143,000 years old.
The remains are also commonly referred to as Ngandong (now at Kradenan district, Blora Regency), after the village near where they were first recovered.
It is a late variant of H. erectus, dated to after 550,000 years ago, overlapping with Homo heidelbergensis and possibly with early Homo sapiens.
Though its morphology was, for the most part, typical of Homo erectus, its cranial capacity of 1,013–1,251 cm³ places it amongst the larger-brained representatives of its species (compared to 900 cm³ for the older Java Man), and its culture was also unusually advanced.Due to the tools found with the fossils and many of their more gracile anatomical features, Solo Man was first classified as a subspecies of Homo sapiens (dubbed Homo sapiens soloensis)
and long thought to be the ancestor of modern Australo-Melanesians. More rigorous studies in the 1990s have concluded that this is not the case. Analysis of 18 crania from Sangiran, Trinil, Sambungmacan, and Ngandong show chronological development from the Bapang-AG to Ngandong periods.Yamashita Cave Man
The Yamashita Cave People (山下洞人, Yamashita Dōjin) are the prehistoric humans known from many bones found in the Yamashita limestone cave near Naha, in Okinawa, Japan. The remains have been dated at 32,000±1000 years ago. The most important bones found in the cave in Yamashita are those of an approximately 6 to 8-year-old girl.