User:Marshallsumter/Keynote lectures (draft)/Human DNA

"[H]uman DNA has millions of on-off switches and complex networks that control the genes' activities. ... [A]t least 80% of the human genome is active, which opposed the previously held idea that most of the DNA are useless."

"DNA contains genes, which hold the instructions for [life. But, these] take up only about 2 percent of the genome ... The human genome is made up of about 3 billion “letters” along strands that make up the familiar double helix structure of DNA. Particular sequences of these letters form genes, which tell cells how to make proteins. People have about 20,000 genes, but the vast majority of DNA lies outside of genes. ... [A]t least three-quarters of the genome is involved in making RNA [...] it appears to help regulate gene activity."

Genetics
There are "more than 4 million sites where proteins bind to DNA to regulate genetic function, sort of like a switch."

"Humans belong to the biological group known as Primates, and are classified with the great apes, one of the major groups of the primate evolutionary tree. Besides similarities in anatomy and behavior, our close biological kinship with other primate species is indicated by DNA evidence. It confirms that our closest living biological relatives are chimpanzees and bonobos, with whom we share many traits. But we did not evolve directly from any primates living today."

"DNA also shows that our species and chimpanzees diverged from a common ancestor species that lived between 8 and 6 million years ago. The last common ancestor of monkeys and apes lived about 25 million years ago."

Deoxyribonucleic acids
Deoxyribonucleic acid (DNA) is a polymer composed of nucleic acids linked together by sugar-phosphate backbone.

The nucleic acids are inorganic acids with phosphoric acid as the only acid.

Attached to each sugar is a nucleobase.

Nitrogenous bases, found in cell nuclei, are nucleobases.

In normal spiral DNA the bases form pairs between the two strands: Adenine (A) with Thymine (T) and Cytosine (C) with Guanine (G). Purines pair with pyrimidines mainly for dimensional reasons - only this combination fits the constant width geometry of the DNA spiral.

"The amount of difference in DNA is a test of the difference between one species and another – and thus how closely or distantly related they are."

"While the genetic difference between individual humans today is minuscule – about 0.1%, on average – study of the same aspects of the chimpanzee genome indicates a difference of about 1.2%. The bonobo (Pan paniscus), which is the close cousin of chimpanzees (Pan troglodytes), differs from humans to the same degree. The DNA difference with gorillas, another of the African apes, is about 1.6%. Most importantly, chimpanzees, bonobos, and humans all show this same amount of difference from gorillas. A difference of 3.1% distinguishes us and the African apes from the Asian great ape, the orangutan. How do the monkeys stack up? All of the great apes and humans differ from rhesus monkeys, for example, by about 7% in their DNA."

"Geneticists have come up with a variety of ways of calculating the percentages, which give different impressions about how similar chimpanzees and humans are. The 1.2% chimp-human distinction, for example, involves a measurement of only substitutions in the base building blocks of those genes that chimpanzees and humans share. A comparison of the entire genome, however, indicates that segments of DNA have also been deleted, duplicated over and over, or inserted from one part of the genome into another. When these differences are counted, there is an additional 4 to 5% distinction between the human and chimpanzee genomes."

"No matter how the calculation is done, the big point still holds: humans, chimpanzees, and bonobos are more closely related to one another than either is to gorillas or any other primate. From the perspective of this powerful test of biological kinship, humans are not only related to the great apes – we are one. The DNA evidence leaves us with one of the greatest surprises in biology: the wall between human, on the one hand, and ape or animal, on the other, has been breached. The human evolutionary tree is embedded within the great apes."

Archaeology
"About 14,000 years ago [14,000 before the year 2000.0, 14,000 b2k], modern humans roamed to South Florida and lived side by side with mammoths, mastodons and saber-tooth tigers."

At "the Old Vero Man site [a] large number of animal and human bones were discovered [...], providing a rare glimpse of the Florida landscape at the end of the last Ice Age."

"Based on the fossil record, here is what many archaeologists believe: Modern humans first appeared about 195,000 years ago in East Africa. Likely chasing prey, they moved across Asia and what was then a land bridge to Alaska, arriving in North America about 20,000 to 25,000 years ago. They then took 5,000 to 6,000 years to cross the continent to Florida."

"The old model basically had these folks sprinting across North America, chasing and killing big animals. We know now they moved very gradually.

"They finally reached Vero Beach about 13,000 to 14,000 years ago, as the Pleistocene Epoch and the last Ice Age were drawing to an end. At the time, Florida was almost double its current size, extending out into the Gulf of Mexico and the Atlantic, and much of the state was more than 300 feet above sea level."

"The landscape was so different than what it is today."

"Florida also was home to tapirs, sloths, camels, bison and horses — in addition to mastodons and mammoths. Many converged at what was then an "oasis" of streams and rivers about 35 miles inland from the ocean. Today, that is the Old Vero Man site and it's about five miles inland."

"It was a fairly constant source of fresh water and a tremendous draw to animals and human beings."

"Growing in numbers and becoming more skilled hunters, the humans continued forging south, evidenced by the Cutler Fossil Site on the Charles Deering Estate in south Miami-Dade County. That site [dates] back almost 12,000 years [12,000 b2k.]"

"[Bones from Paleo-Indians and] 103 species of animals, including mammoths, a saber-tooth cat, a paleo-lama, and a California condor [have been found at the Cutler Fossil Site]."

"By the time humans arrived in what is now South Florida, the Ice Age was almost over. Mastodons and their ilk were going extinct, likely because of climate change."

"With ancient DNA, you're time traveling. It provides us with a unique opportunity to look into Florida's past."

Paleogene
The Paleogene Period extends from 65.5 ± 0.3 to 23.03 ± 0.05 x 106 b2k.

The graph above shows many of the hominins that have been discovered so far in Africa and elsewhere on Earth.

Paleocene
The Paleocene dates from 65.5 ± 0.3 x 106 to 55.8 ± 0.2 x 106 b2k.

Eocene
The Eocene dates from 55.8 ± 0.2 x 106 to 33.9 ± 0.1 x 106 b2k.

Oligocene
The Oligocene dates from 33.9 ± 0.1 x 106 to 23.03 x 106 b2k.

Rukwapithecus
Rukwapithecus is "an early member of the hominoids, the group containing the great apes (gorillas, chimpanzees, bonobos, orangutans and humans) and lesser apes (gibbons)."

"The fossil remnants ... date back 25 million years ago, filling a gap in the fossil record that reveals when apes and monkeys first diverged."

"These discoveries are important because they offer the earliest fossil evidence for either of these primate groups".

"The fossils were found in a layer of the Rukwa Rift in Tanzania. The region is part of the East African Rift, a tectonic-plate boundary where the Earth's crust is being pulled apart."

“The new discoveries are particularly important for helping to reconcile a long-standing disagreement between divergence time estimates derived from analyses of DNA sequences from living primates and those suggested by the primate fossil record.”

"Studies of clock-like mutations in primate DNA have indicated that the split between apes and Old World monkeys occurred between 30 million and 25 million years ago."

Holarctic-Antarctic Ice Age
"This late Cenozoic ice age began at least 30 million years ago in Antarctica; it expanded to Arctic regions of southern Alaska, Greenland, Iceland, and Svalbard between 10 and 3 million years ago. Glaciers and ice sheets in these areas have been relatively stable, more-or-less permanent features during the past few million years."

Neogene
The Neogene dates from 23.03 x 106 to 2.58 x 106 b2k.

Miocene
The Miocene dates from 23.03 x 106 to 5.332 x 106 b2k.

Tortonian
The Tortonian lasted from 11.63 Ma to 7.246 Ma.

Gigantopithecus is an extinct genus of ape that existed from perhaps nine million years to as recently as one hundred thousand years ago, at the same period as Homo erectus would have been dispersed, in what is now India, Vietnam, China and Indonesia placing Gigantopithecus in the same time frame and geographical location as several hominin species. The primate fossil record suggests that the species Gigantopithecus blacki were the largest known primates that ever lived, standing up to 3 m and weighing as much as 540 - 600 kg,  although some argue that it is more likely that they were much smaller, at roughly 1.8 - 2 m in height and 180 - 300 kg in weight.

Prehistory
The prehistory period dates from around 7 x 106 b2k to about 7,000 b2k.

Pliocene
The Pliocene ranges from 5.332 x 106 to 2.588 x 106 b2k.

Zanclean
"The boundary-stratotype of the stage is located in the Eraclea Minoa section on the southern coast of Sicily (Italy), at the base of the Trubi Formation. The age of the Zanclean and Pliocene GSSP at the base of the stage is 5.33 Ma in the orbitally calibrated time scale, and lies within the lowermost reversed episode of the Gilbert Chron (C3n.4r), below the Thvera normal subchron."

Piacenzian
"The base of the beige marl bed of the small-scale carbonate cycle 77 (sensu Hilgen, 1991b) is the approved base of the Piacenzian Stage (that is the Lower Pliocene-Middle Pliocene boundary). It corresponds to precessional excursion 347 as numbered from the present with an astrochronological age estimate of 3.600 Ma (Lourens et al., 1996a)."

Paleolithic
The paleolithic period dates from around 2.6 x 106 b2k to the end of the Pleistocene around 12,000 b2k.

Quaternary
The "whole change elapsed just opposite the course of events that characterized the great glacial oscillations with sudden warming followed by slow cooling. Therefore, the two phenomena hardly have the same cause."

Pleistocene
The Pleistocene dates from 2.588 x 106 to 11,700 b2k.

Gelasian
"The base of the Quaternary System [shown in the image on the right] is defined by the Global Stratotype Section and Point (GSSP) of the Gelasian Stage at Monte San Nicola in Sicily, Italy, currently dated at 2.58 Ma."

Calabrian
"The [Calabrian] GSSP occurs at the base of the marine claystone conformably overlying sapropelic bed ‘e’ within Segment B in the Vrica section. This lithological level represents the primary marker for the recognition of the boundary, and is assigned an astronomical age of 1.80 Ma on the basis of sapropel calibration."

Homo erectus
""Peking Man," a human ancestor who lived in China between roughly 200,000 and 750,000 years ago, was a wood-working, fire-using, spear-hafting hominid ... these hominids, a form of Homo erectus, appear to have been quite meticulous about their clothing, using stone tools to soften and depress animal hides."

Peking Man Homo erectus pekinensis, is an example of Homo erectus. A group of fossil specimens was discovered in 1923–27 during excavations at Zhoukoudian (Chou K'ou-tien) near Beijing. The finds have been dated from roughly 750,000 years ago, although a new 26Al/10Be dating suggests they may be as much as 680,000–780,000 years old. Skulls X, XI and XII (sometimes called LI, LII and LIII) were discovered at Locus L in 1936. They are thought to belong to an adult man, an adult woman and a young adult, with brain sizes of 1225 cc, 1015 cc and 1030 cc respectively. The ribonucleotide reductase RRM2P4 gene data suggests that the Chinese, while largely descending from Africa like others, nevertheless have some genetic legacy from hybridization with older Eurasian populations, consistent with limited multiregional evolution.

Homo erectus  is an extinct species of hominid that lived from the end of the Pliocene epoch to the later Pleistocene, with the earliest first fossil evidence dating to around 1.8 million years ago and the most recent to around 300,000 years ago. The species originated in Africa and spread as far as Spain, Georgia, India, China and Java.

"By the 1980s, the growing numbers of H. erectus specimens, particularly in Africa, led to the realization that Asian H. erectus (H. erectus sensu stricto), once thought so primitive, was in fact more derived than its African counterparts. These morphological differences were interpreted by some as evidence that more than one species might be included in H. erectus sensu lato (e.g., Stringer, 1984; Andrews, 1984; Tattersall, 1986; Wood, 1984, 1991a, b; Schwartz and Tattersall, 2000)." ... "Unlike the European lineage, in my opinion, the taxonomic issues surrounding Asian vs. African H. erectus are more intractable. The issue was most pointedly addressed with the naming of H. ergaster on the basis of the type mandible KNM-ER 992, but also including the partial skeleton and isolated teeth of KNM-ER 803 among other Koobi Fora remains (Groves and Mazak, 1975). Recently, this specific name was applied to most early African and Georgian H. erectus in recognition of the less-derived nature of these remains vis à vis conditions in Asian H. erectus (see Wood, 1991a, p. 268; Gabunia et al., 2000a). It should be noted, however, that at least portions of the paratype of H. ergaster (e.g., KNM-ER 1805) are not included in most current conceptions of that taxon. The H. ergaster question remains famously unresolved (e.g., Stringer, 1984; Tattersall, 1986; Wood, 1991a, 1994; Rightmire, 1998b; Gabunia et al., 2000a; Schwartz and Tattersall, 2000), in no small part because the original diagnosis provided no comparison with the Asian fossil record."

From the 1950s to 1970s, however, numerous fossil finds from East Africa yielded evidence that the oldest hominins originated there. It is now believed that H. erectus is a descendant of earlier genera such as Ardipithecus and Australopithecus, or early Homo-species such as H. habilis or H. ergaster. H. habilis and H. erectus coexisted for several thousand years, and may represent separate lineages of a common ancestor.

"[S]ome of the oldest stone hand axes on Earth ... unearthed in Ethiopia ... date to 1.75 million years ago. ... fossilized H. erectus remains were also found at the same site ... These Aucheulean tools could be up to 7.8 inches (20 centimeters) long".

"[J]awbone fossils unearthed at a site east of Lake Turkana in Kenya suggest there were two additional species of ... Homo, living alongside ... Homo erectus, nearly 2 million years ago."

When "excavating a cave in Balanica, Serbia, that contained ancient archaeological remains ... an ancient jawbone fragment with three molars still intact [was found] ... several dating techniques [determined] the fragment was definitely older than 397,000 years and perhaps older than 525,000 years. The jawbone lacked several characteristic Neanderthal features, including distinctive chewing surfaces on the teeth that show up in Western Europe at that time. Instead, the fossil resembled the more primitive Homo erectus. ... Neanderthals may not have evolved in this region of Southeastern Europe, at least during this time. ... during several ice ages, rising glaciers over the past eons cut off Western Europe from the rest of the continent, and this isolation likely contributed to the evolution of Neanderthals' distinctive features from the more primitive Homo erectus."

"Javanese specimens of Homo erectus had brains about 860 cubic cm (52 cubic inches) large".

"Researchers examined one skull from a site called the Pit of Bones, which contains the remains of at least 28 people. [...] two fractures on that skull were likely to have been caused by "multiple blows" and imply "an intention to kill"."

"As well as providing a clue as to why the bodies were in the cave, scientists say the study provides grisly evidence that violence is an intrinsic part of the earliest human culture."

"This individual was killed in an act of lethal interpersonal violence."

The "long vertical shaft of this cave was a place where these ancient people deliberately "deposited deceased members of their social groups"."

"Intentional interpersonal violence is a behaviour that accompanies humans since at least 430,000 years ago, but so does the care of sick or even the care of the dead."

"We have not changed much in the last half million years."

"I suspect the farther we push back and find straight up forensic evidence such as these authors have, we will find that violence is culturally mediated and has been with us as long as culture itself has been with us."

"We don’t see any Denisovan ancestry in populations living in mainland Southeast Asia, or Indonesia, or indeed in any population west of Wallace’s Line [which marks the eastern border of placental mammals, while Lydekker’s line marks the western border of the marsupial fauna: Wallacea lies between]. It could be that later population movements have diluted the Denisovan taint, but it is apparently undetectable even in existing populations that are thought to represent earlier strata, such as the Andaman islanders. Nor has it been seen in ancient DNA from modern humans on the Asian mainland."

"Back in the day, Indonesia was a big peninsula, called Sundaland. Let us assume that there were Denisovans there. In some way, Denisovan effective population density was higher there than in mainland Asia, or perhaps they were harder to displace.  For example, some of Sundaland seems to have been tropical rainforest, otherwise known as the Green Hell – maybe there were some potent tropical diseases (vector-borne) that the Denisovans had developed resistance to, while the invading anatomical modern humans had not [Cooper and Stringer were also thinking about jungle pathogens] – the same reason that Europeans or Middle Easterners didn’t replace sub-Saharan African populations."

"So modern humans expand into Sundaland more slowly, in something like a range expansion. The further they moved into Sundaland, the more Denisovan genes they picked up."

"So the humans living at the eastern edge of Sundaland, particularly in Borneo, have more Denisovan ancestry in this scenario than anyone else in Eurasia – and they would be the people who take the next step, crossing the narrow seas to the Philippines, Wallacea, and then Sahul (Australia/New Guinea), virgin lands in which the hand of man had never set foot [with the possible exception of Flores].   A quite small, unusually Denisovanized coastal population of hunter-gatherers then undergoes a vast expansion, becoming as numerous as the stars in the sky."

"Aboriginal skulls vary, but some look much more like archaics, more like erectus, than those from any other existing population. This was noticed a long time ago, by pros like Weidenreich."

"Actually, they look surprisingly archaic, considering that total archaic ancestry among Melanesians is under 10% [Neanderthal plus Denisovan]. Recent selective pressure are probably more important: it is possible that their archaic appearance is to some degree coincidence."

“The skulls of Australian Aborigines, although variable, in some ways look more old-fashioned, more like archaics, than any other existing human population.”

"The discovery of DNA in a 400,000-year-old human thigh bone [femur] comes from the famed "Pit of Bones" site in Spain, which gave up the remains of at least 28 ancient people."

"The early human remains from the cave site near the northern Spanish city of Burgos have been painstakingly excavated and pieced together over the course of more than two decades. It has yielded one of the richest assemblages of human bones from this stage of human evolution, in a time called the Middle Pleistocene."

"The fossils carry many traits typical of Neanderthals, and either belong to an ancestral species known as Homo heidelbergensis - or, as the British palaeoanthropologist Chris Stringer suggests - are early representatives of the Neanderthal lineage."

"Rather than showing a relationship between the Spanish specimens and Neanderthals, which might be expected based on their physical features, the mitochondrial DNA was most similar to that found in 40,000 year-old material unearthed thousands of kilometres away at Denisova Cave in Siberia."

"The Denisovans were a sister group to the Neanderthals, with distinct genetic characteristics. Identified only by DNA extracted from a tiny finger bone and tooth, they are, as some researchers have remarked, "a genome in search of a fossil" because there are no substantial remains representative of this group."

"By using missing mutations in the old DNA sequences, the researchers calculated that the Pit of Bones individual shared a common ancestor with the Denisovans about 700,000 years ago."

Interbreeding "between the Pit of Bones people (or their ancestors) and yet another early human species brought the Denisovan-like DNA into this western population. Prof Bermudez de Castro thinks there may be a candidate for this cryptic ancestor: an earlier human species known as Homo antecessor. One million years ago, antecessor inhabited the site of Gran Dolina, just a few hundred metres away from the Pit of Bones."

"mtDNA is a small and unusual component of our genetic blueprint, from which only limited conclusions can be drawn. For example, no sign of the interbreeding we now know took place between Neanderthals and modern humans remains in the mtDNA of modern people."

"That is our next big thing here, to sequence at least part of the nuclear genome from the individual in the Sima de los Huesos."

Homo heidelbergensis
"Homo heidelbergensis is known to have lived from at least 600,000 years ago in Africa and Europe to maybe as late as 250,000 years ago in some areas."

"The first fossil identified as H. heidelbergensis was a jaw discovered near Heidelberg, Germany in 1907. Since then various other finds have been made in Europe, Asia and Africa. They show a less projecting face, more prominent nose and a bigger braincase than Homo erectus, but also more primitive features than those of Neanderthals and modern humans."

"Some H. heidelbergensis individuals had brain sizes within the modern human range. However, facially they still looked very different from us, with a longer, lower shaped skull, large brow ridge and no chin."

"Evidence shows that H. heidelbergensis was an accomplished tool-maker and skillfully butchered large animals. The remains of horses, elephants, deer and rhinoceroses with butchery marks on their bones have been found alongside fossils of this hominin in Southern England and Germany. Whether they actively hunted the animals isn’t known. But, even if they scavenged the carcasses, these hominins were organised enough to drive off dangerous competing animals such as lions, hyenas and wolves."

Early Pleistocene
Early Pleistocene spans ca. 730,000-1,600,000 yr BP.

Nebraskan glacial
Nebraskan glacial spans ca. 650,000-1,000,000 yr BP.

On the right is an image showing an "analysis of hominid tooth evolution, including specimens from Spanish Neandertals (top row), pushes back the age of a common Neandertal-human ancestor to more than 800,000 years ago. The bottom row shows Homo sapiens teeth."

"During hominid evolution, tooth crowns changed in size and shape at a steady rate."

"The Neandertal teeth, which date to around 430,000 years ago, could have evolved their distinctive shapes at a pace typical of other hominids only if Neandertals originated between 800,000 and 1.2 million years ago."

Middle Pleistocene
Middle Pleistocene spans ca. 150,000-730,000 yr BP.

Aftonian interglacial
"Clay deposition in the Piauí River floodplain around 436 ± 51.5 ka occurred during a warmer period of the Aftonian interglaciation, corresponding to isotope stage 12 (Ericson and Wollin, 1968)."

"Neanderthal and Homo sapiens DNA reveals that they shared a common ancestor about 400,000 years ago. Many scientists think this could have been H. heidelbergensis, giving rise to Neanderthals in Europe and to our species in Africa. And perhaps to the Denisovans in Asia."

Neanderthals
"[M]odern humans in the eastern parts of Eurasia and Native Americans actually carry more Neanderthal genetic material than people in Europe, "even though the Neanderthals mostly lived in Europe, which is really, really interesting," Reich said."

From mtDNA analysis estimates, the two species shared a common ancestor about 500,000 years ago. An article appearing in the journal Nature has calculated the species diverged about 516,000 years ago, whereas fossil records show a time of about 400,000 years ago. A 2007 study pushes the point of divergence back to around 800,000 years ago.

"Our work suggests that at present, it is unlikely that Neanderthals survived any later in this area [southern Iberia] than they did elsewhere in mainland Europe."

"Neanderthals (or Neandertals) are our closest extinct human relatives. [They may have been] a subspecies [Homo sapiens neanderthalensis]. Our well-known [...] fossil kin lived in Eurasia 200,000 to 30,000 years ago, in the Pleistocene Epoch. [...] they used tools, buried their dead and controlled fire, among other intelligent behaviors."

"Neanderthals were seen as too brutish to catch fast prey."

"Neanderthals came to Europe some 300,000 years ago. They hunted big game with stone tools. Their territory spanned Europe and Asia. They left distinctive "Mousterian" artefacts."

"The first analysis of mitochondrial DNA (mtDNA) from Neanderthals was published in 1997."

"The specimen was taken from the first Neanderthal fossil discovered, from Feldhofer Cave, in the Neander Valley, Germany. A small sample of bone was ground up to extract mtDNA."

"The Neanderthal mtDNA sequences were substantially different from modern human mtDNA (Krings et al. 1997, 1999). Researchers compared the Neanderthal to modern human and chimpanzee sequences. Most human sequences differ from each other by on average 8.0 substitutions, while the human and chimpanzee sequences differ by about 55.0 substitutions. The Neanderthal and modern human sequences differed by approximately 27.2 substitutions. Using this mtDNA information, the last common ancestor of Neanderthals and modern humans dates to approximately 550,000 to 690,000 years ago, which is about four times older than the modern human mtDNA pool. This is consistent with the idea that Neanderthals did not contribute substantially to modern human genome."

"A second mtDNA sequence, announced in 2000, was derived from a 29,000 year old Neanderthal found in Mezmaiskaya Cave, Russia (Ovchinnikov et al. 2000). Although the Mezmaiskaya Cave sequence was slightly different than the Feldhofer Neanderthal, the two Neanderthal mtDNA sequences were distinct from those of modern humans. These results confirmed the earlier findings that showed that Neanderthals were unlikely to have contributed to the modern human genome. As with the previous study of Neanderthal mtDNA, results were consistent with separation between the Neanderthal and modern human gene pools or with very low amounts of gene flow between the two groups."

"Researchers have also studied ancient DNA from anatomically modern Homo sapiens from Europe dating to the same time period as the Neanderthals. Material from two Paglicci Cave, Italy individuals, dated to 23,000 and 25,000 years old, was sequenced. The Paglicci Homo sapiens mtDNA sequences were different from all Neanderthal mtDNA sequences but were within the range of variation for modern human mtDNA sequences (Caramelli et al. 2003). Mitochondrial DNA from the Paglicci specimens as well as other ancient humans fit within the range of modern humans, but the Neanderthals remain consistently genetically distinct. This shows that early anatomically modern Homo sapiens were not very different genetically from current modern humans, but were still different from Neanderthals."

"DNA was extracted from three Neanderthal bones from Vindija Cave, Croatia. By comparing sequences from their mtDNA and their nuclear DNA, scientists determined that the three bones came from different individuals, although two of them might be related on their mother’s side."

"The Neanderthal sequence was compared to those of five modern humans from France, China, Papua New Guinea, as well as Africans from the San and Yoruba groups. Tests indicated that Neanderthals shared more derived alleles with non-African modern humans than with African modern humans. They compared parts of the Neanderthal genome with pairs of modern humans. While the European and Asian pairs had similar amounts of derived material compared with the Neanderthal, Neanderthals had more similarities with non-African humans than with Africans."

"Neanderthals have contributed approximately 1% to 4% to the genomes of non-African modern humans."

"Since the Neanderthal DNA was equally related to that of the modern samples from France, China and Papua New Guinea, admixture between moderns and Neanderthals must have occurred before the Eurasian populations split off from each other. Remains of both modern humans and Neanderthals dating to around 100,000 years ago have been found in the Middle East."

"Researchers found 78 sequence differences that would have affected proteins in which Neanderthals had the ancestral state and modern humans had a newer, derived state. Five genes had more than one sequence change that affected the protein structure. These proteins include SPAG17, which is involved in the movement of sperm, PCD16, which may be involved in wound healing, TTF1, which is involved in ribosomal gene transcription, and RPTN, which is found in the skin, hair and sweat glands. Scientists do not know the function of the CAN15 protein, which was also one of the differences."

"Nested clade phylogenetic analysis shows evidence of three expansions out of Africa at 1.9 Ma, 650,000 years, and 130,000 years, which is consistent with the admixture between ancient and modern populations rather than complete replacement (Templeton 2002, 2005, 2007)."

"Ancient DNA has been used to show aspects of Neanderthal appearance. A fragment of the gene for the melanocortin 1 receptor (MRC1) was sequenced using DNA from two Neanderthal specimens from Spain and Italy, El Sidrón 1252 and Monte Lessini (Lalueza-Fox et al. 2007). Neanderthals had a mutation in this receptor gene that has not been found in modern humans. The mutation changes an amino acid, making the resulting protein less efficient. Modern humans have other MCR1 variants that are also less active resulting in red hair and pale skin. The less active Neanderthal mutation probably also resulted in red hair and pale skin, as in modern humans."

"The specific MCR1 mutation in Neanderthals has not found in modern humans (or occurs extremely rarely in modern humans). This indicates that the two mutations for red hair and pale skin occurred independently and does not support the idea of gene flow between Neanderthals and modern humans. Pale skin may have been advantageous to Neanderthals living in Europe because of the ability to synthesize vitamin D."

"The FOXP2 gene is involved in speech and language (Lai et al. 2001). Changes in the FOXP2 gene sequence led to problems with speech, oral and facial muscle control in modern humans with a mutation in the gene. It impairs language function. Modern humans and Neanderthals share two changes in FOXP2 compared with the sequence in chimpanzees (Krause et al. 2007)."

"The gene that produces the ABO blood system is polymorphic in humans. Various selection factors may favor different alleles, leading to the maintenance of distinct blood groups in modern human populations. Though chimpanzees also have different blood groups, they are not the same as human blood types. While the mutation that causes the human B blood group arose around 3.5 Ma, the O group mutation dates to around 1.15 Ma. Lalueza-Fox and colleagues (2008) tested whether Neanderthals had the O blood group. They found that two Neanderthal specimens from Spain probably had the O blood type, though there is the possibility that they were OA or OB. Though the O allele was likely to have already appeared before the split between humans and Neanderthals, it could also have arisen in the Neanderthal genome via gene from modern humans."

"The ability to taste bitter substances is controlled by a gene, TAS2R38. Some individuals are able to taste bitter substances, while others have a different version of the gene that does not allow them to taste bitter items. Possession of two copies of alleles associated with tasting bitter substances gives the individual greater perception of bitter tastes than the heterozygous state, in which individuals have one tasting allele and one non-tasting allele. Two copies of a non-tasting allele leads to inability to taste bitter substances."

"A Neanderthal from El Sidrón, Spain, was sequenced for the TAS23R38 gene. They found that this individual was heterozygous and thus was able to perceive bitter taste, although not as strongly as a homozygous individual with two copies of the tasting allele would be able to (Lalueza-Fox et al. 2009). Since the Neanderthal sequenced was heterozygous, the two alleles (tasting and non-tasting) were probably both present in the common ancestor of Neanderthals and modern humans. Though chimpanzees also vary in their ability to taste bitterness, their abilities are controlled by different alleles than those found in humans, indicating that non-tasting alleles evolved separately in the hominin lineage."

"The microcephalin gene relates to brain size during development. A variant of this, haplogroup D, may have been positively selected for in modern humans – and may also have come from an interbreeding event with an archaic population (Evans et al. 2006). Mutations in microcephalin cause the brain to be 3 to 4 times smaller in size. All of the haplogroup D variants come from a single copy that appeared in modern humans around 37,000 years ago. However, haplogroup D itself came from a lineage that had diverged from the lineage that led to modern humans around 1.1 million years ago. Although there was speculation that the Neanderthals were the source of the microcephalin haplogroup D (Evans et al. 2006), the Neanderthal DNA recently sequenced does not contain the microcephalin haplogroup D (Green et al. 2010)."

Homo sapiens
"The first anatomically modern human fossils date back only 195,000 years ... [But the Y chromosome of a recently deceased African-American, Albert Perry,] who lived in South Carolina [is] so distinct that his male lineage probably separated from all others about 338,000 years ago. ... [There are] similarities between Perry's [Y chromosome] and those in samples taken from 11 men, all living in one village in Cameroon. ... "The oldest known fossil humans in both West Africa at Iwo Eleru and Central Africa at Ishango [in Democratic Republic of the Congo] show unexpectedly archaic features, so it certainly looks like we have a more complex scenario for the evolution of modern humans in Africa.""

Two "teeth from the Luna cave in China's Guangxi Zhuang region [based] on the proportions of the teeth, [...] at least one of them must have belonged to an early Homo sapiens [...] The teeth are clearly old. Calcite crystals, which formed as water flowed over the teeth and the cave floor, date them to between 70,000 and 125,000 years ago. [...] they are evidence of an early wave of modern humans in eastern Asia."

"I am not convinced that these teeth are diagnostic."

"Bones found in Israel, including an upper jaw from Misliya cave, could be 150,000 years old."

In "the identification of a jawbone and two molars from Zhirendong, a cave in Guizhou province [though] the bone is over 100,000 years old, [...] the shape of its chin is suggestive of modern humans."

"Though the Asian fossils are Homo sapiens-like, another species could have evolved these features in parallel."

The "genomes of indigenous populations from south-east Asia [have been plugged] into a migration model. [The] genetic data was best explained by an early exodus that left Africa around 130,000 years ago, taking a coastal route along the Arabian peninsula, India and into Australia, followed by a later wave along the classic route."

Kansan glacial
Kansan glacial spans 500,000-600,000 yr BP.

Yarmouthian interglacial
"Clay deposition in the Piauí River floodplain around 436 ± 51.5 ka occurred during a warmer period of the [Yarmouthian interglaciation] Aftonian interglaciation, corresponding to isotope stage 12 (Ericson and Wollin, 1968)."

Illinois Episode glaciation
"Illinoian [is] (ca. 220,000-430,000 yr BP)".

"The [Jebel Irhoud site] Moroccan fossils [...] are roughly 300,000 years old. Remarkably, they indicate that early Homo sapiens had faces much like our own, although their brains differed in fundamental ways."

Sangamon Episode interglacial
"OSL dates also suggest that last interglacial (MIS 5; Sangamon Ep.) fluvial deposits are preserved locally."

Age "assignment of Sangamonian (sense alto = 80,000-ca. 220,000 yr BP) [is] to Illinoian (ca. 220,000-430,000 yr BP)".

"A broken skull chiselled from a lump of rock in a cave in Greece is the oldest modern human fossil ever found outside Africa. The partial skull was discovered in the Apidima cave on the Mani peninsula of the southern Peloponnese and has been dated to be at least 210,000 years old."

The "skull revealed that at least some modern humans had left Africa far earlier than previously thought and reached further geographically to settle as far away as Europe."

"Other fossils of early modern humans found in Israel already point to brief excursions out of Africa, where the species evolved, long before the mass exodus during which Homo sapiens spread from the continent about 70,000 years ago and colonised the world."

"Our results indicate that an early dispersal of Homo sapiens out of Africa occurred earlier than previously believed, before 200,000 years ago,” Karvati said. “We’re seeing evidence for human dispersals that are not just limited to one major exodus out of Africa."

"The story of the skull is unusual from the start. It was found during excavations of the Apidima cave, a hole in a limestone cliff that now towers over the sea, in the late 1970s. The fossil was encased in a lump of rock, mere inches from another skull and several bone fragments. The rock itself was wedged high up between adjacent walls of the cave."

"Once removed from the cave, the skulls were stored in a museum in Athens but received little attention until recently, in part because they are so damaged and incomplete. The second skull, which retains a face, was studied the most and identified as Neanderthal. The first skull [in the image on the right], consisting only of the back of the skull, was largely ignored."

Late Pleistocene
Late Pleistocene spans ca. 11,000-150,000 yr BP.

Eemian interglacial
The "controversially split Eemian period, the predecessor of our own warm period about 125,000 years ago."

"The Eem interglaciation […] lasted from 131 to 117 kyr B.P."

The "Neanderthal fossil [in the image on the right] was 120,000 years old and, more important, that it belonged to a branch of the Neanderthal family tree with a long history. All known Neanderthals inherited their mitochondrial DNA from an ancestor who lived 270,000 years ago."

"The common ancestors of Neanderthals and Denisovans spread across Europe and Asia over half a million years ago. Gradually the eastern and western populations parted ways, genetically speaking."

"In the east, they became Denisovans. In the west, they became Neanderthals. The 430,000-year-old fossils at Sima de los Huesos — Neanderthals with Denisovanlike genes — capture the early stage of that split."

"At some point before 270,000 years ago, African humans closely related to us moved into Europe and interbred with Neanderthals. Their DNA entered the Neanderthal gene pool."

Herning Stadial
MIS Boundary 5.5 (peak) is at 123 ka.

"Lines engraved between 125,000 and 105,000 years ago on two animal bones found in northern China held some sort of meaning for their makers."

"These two bone fragments [in the image on the right] found in northern China contain engraved lines, some marked with red pigment (red dots in left line drawing), making them the oldest examples of symbolic behavior in East Asia."

"These ancient markings provide the oldest evidence of symbolic activity by humans or our close evolutionary relatives in East Asia."

"A mysterious Stone Age population called Denisovans, which had close genetic ties to Neandertals, may have carved sets of parallel lines into the pair of bone fragments."

"Denisovans inhabited East Asia at the same time that someone carved lines into bones at northern China’s Lingjing site [...]. But either Homo sapiens or Neandertals, who also left behind Stone Age creations with apparent symbolic meanings [...], might instead have modified the Lingjing bones."

"Nonetheless, the two objects from Lingjing suggest that symbolic capacities were within the realm of cognitive abilities of [Homo] species that lived before and during the evolution of Homo sapiens in Africa.”

"Abstract markings on the Lingjing bones resemble engraved lines on roughly 100,000-year-old pigment chunks from South Africa."

"As Homo sapiens was responsible for that early symbolism in Africa, and Neandertals were responsible for such in Europe, it is a fascinating possibility that these [Chinese] examples were created by another Homo species."

"An engraved geometric design on a roughly half-million-year-old seashell found on an Indonesian island stands as the oldest example of symbolic behavior anywhere in the world [...]. Researchers suspect the now-extinct hominid species Homo erectus carved that pattern."

"Almost one-quarter of 227 animal bone fragments excavated at Lingjing between 2005 and 2015 display stone tool incisions typical of butchery [...]. But that sample included two exceptions. Seven nearly parallel lines had been cut into a partial rib from an unidentified large, adult mammal. Microscopic analysis indicated that the lines were made with a sharp point that was run across the bone’s surface after it had suffered some damage from weathering. Special care was taken to create each of the first five lines with a single pass of the engraving tool. Red residue in four engraved lines indicated that a pigment had been smeared on the pattern, possibly to increase its visibility."

"A second rib fragment from a large mammal contained 10 roughly parallel lines that had been sliced with a sharp stone point, probably in a single session [...]. Engraving of these lines also occurred after the bone had been damaged by long exposure to the air. No pigment residue appeared on this specimen."

"Estimated ages of the engraved bones relied on calculations of the time since sediment in which they were found was last exposed to sunlight."

"Whoever cut lines into the Lingjing finds also fashioned animal bones into tools. Bone and antler artifacts found in the same sediment as the engraved ribs were likely used to retouch and sharpen used stone tools."

Brørup interstadial
The "Brørup interstade [is about] 100 ka BP". It corresponds to GIS 23/24.

Rederstall Stadial
MIS Boundary 5.3 is at 96 ka.

Marine Isotope Stage 4.

"During the Middle Stone Age of Southern Africa, technological and behavioral innovations led to significant changes in the lifeways of modern humans. The glacial episode of Marine Isotope Stage 4, about 57-71,000 years ago, resulted in cooler and drier climatic conditions and the expansion of grassland vegetation. Sea level dropped by as much as 80 meters below its current level. During this period the cultural phase known as the Howieson’s Poort appeared across much of Southern Africa, peaking at about 60-65,000 years ago, and then disappeared. The lithic industry of the Howieson’s Poort is exemplified by changes in technology, such as the use of the punch technique, an increase in the selection of fine-grained silcrete, and the predominance of retouched pieces including backed tools, segments, scrapers and points. Segments are the type fossil of the Howieson’s Poort and represent multi-purpose armatures that were hafted onto wooden spear shafts. The standardized design and refined style of segments convey information about the behavior of their makers and provide insight about group identity. Increasing use of ochre, the presence of engraved ostrich eggshells, and a bone tool industry are associated with these stone artifacts. Also evident is an intensified use of space. Taken together, these behaviors suggest that the Howieson’s Poort represents a clear marker of modern human culture."

Wisconsinian glacial
Wisconsinian glacial began at 80,000 yr BP.

Odderade interstadial
The Odderade interstadial has a 14C date of 61-72 kyr B.P. and corresponds to GIS 21.

"A population that began to expand from Africa around 70 Ka reached Southeast Asia by the middle of the Upper Pleistocene. From Southeast Asia, part of this population took a southern route of expansion towards Australia, where they arrived around 50 Ka (Lahr and Foley, 1998). Sometime between 50 and 20 Ka the same Southeast Asian population took a northern route of expansion along both interior and coastal East Asia (as suggested by Fladmark, 1979), depending on the local climatic conditions. The presence of an “Australo-Melanesian-like” population in East Asia is attested by the human skeletal remains from the Zhoukoudian Upper Cave around 20 Ka (Kamminga and Wright, 1988; Wright, 1995; Neves and Pucciarelli, 1998). A late occupation of the northern zones of East Asia also agrees with the idea that major behavioral change and technological strategies necessary to exploit extreme environments were not in place until ca. 40,000 BP (Whallon, 1989; Klein, 1992, 1995)."

Karmøy stadial
The Karmøy stadial begins in the high mountains of Norway about 60 kyr B.P. and expands to the outer coast by 58 kyr B.P.

"To uncover the migratory path that the ancestors of present-day Eurasians (Europeans and Asians) took when moving out of Africa around 60,000 years ago, an international team of scientists generated 225 whole-genome sequences from six modern Northeast African populations (100 Egyptians and five Ethiopian populations each represented by 25 people)."

"The remaining masked genomic regions from Egyptian samples were more similar to non-African samples and present in higher frequencies outside Africa than the masked Ethiopian genomic regions, pointing to Egypt as the more likely gateway in the exodus to the rest of the world."

“Two geographically plausible routes have been proposed: an exit through the current Egypt and Sinai, which is the northern route, or one through Ethiopia, the Bab el Mandeb strait, and the Arabian Peninsula, which is the southern route.”

"[H]igh-quality genomes [were used] to estimate the time that the populations split from one another: people outside Africa split from the Egyptian genomes more recently than from the Ethiopians (55,000 as opposed to 65, 000 years ago), supporting the idea that Egypt was last stop on the route out of Africa."

“In our research, we generated the first comprehensive set of unbiased genomic data from Northeast Africans and observed, after controlling for recent migrations, a higher genetic similarity between Egyptians and Eurasians than between Ethiopians and Eurasians. This suggests that Egypt was most likely the last stop on the way out of Africa.”

Oerel interstadial
The Oerel interstadial has a 14C date of 53-58 kyr B.P. and corresponds to GIS 15/16 with a GIS age of 56-59 kyr B.P.

"Neanderthals had the brains and guile to catch and eat birds, a skill many had assumed was beyond them. Bones found in Gibraltar suggest Neanderthals hunted wild pigeons, possibly by climbing steep cliffs to reach their nests."

"The rock dove bones were buried in sediments laid down between 28,000 and 67,000 years ago. Most of the excavated layers date from a time when only Neanderthals lived in the area, before the arrival of modern humans around 40,000 years ago. That means only Neanderthals could have caught the rock doves."

"They couldn't have picked up the skills to catch the birds from modern humans."

"This provides the first evidence for sustained and significant use of birds for food by Neanderthals."

"The more we can show similarities between our ancestors and Neanderthals, the more the barriers between us are broken down."

"The sustained use of pigeons provides even more evidence that Neanderthal hunting and foraging abilities were on a par with those of modern humans."

"We know that they climbed up cliffs to hunt ibex, so maybe they also climbed to the ledges where the birds nested. I think they might have had snares or netting made from grasses, but we'll never know as it's all perishable."

"Neanderthals may have started with eating [the birds], then moved to other purposes such as clothing or ornamentation."

Ebersdorf Stadial
"Genetics suggests Neanderthal numbers dropped sharply around 50,000 years ago. This coincides with a sudden cold snap, hinting climate struck the first blow."

"There is a surprising genetic unity between the earliest known Europeans and contemporary Europeans, ancient DNA reveals. This finding suggests that a complex network of sexual exchange may have existed across Europe over the past 50,000 years, and also helps to pinpoint when modern humans interbred with Neanderthals, the closest extinct relatives of modern humans."

"There is solid evidence of modern humans at Tam Pa Ling [in Laos] around 50,000 or 60,000 years ago, and the Zhirendong mandible has modern features. So yes, modern humans were present in at least south-east Asia and south China by somewhere in this time range."

Glinde interstadial
The Glinde interstadial has a 14C date of 48-50 kyr B.P. and corresponds to GIS ?13/14 with a GIS age of 49-54.5 kyr B.P.

"It is recognized that the human population that arrived in Australia around 50,000 BP (Bowler et al., 2003) was the product of an expansive movement out of Africa, following the tropical areas of Southern Asia. This route of expansion represents one of the first offshoots of modern humans out of Africa (Stringer and Andrews 1988; Stringer 1992; Lahr, 1996; Lahr and Foley, 1998). This is why Australians and Africans still form a supra-population unit of morphological affinity (Howells, 1973, 1989; Lahr, 1996). Therefore, the similarities of the first South Americans with Australians are easily explained if we accept that both populations shared a common ancestral population in mainland East Asia (most probably Southeast Asia). A population expansion from Asia to the New World before the Mongoloid traits were fully developed in the Old World was actually predicted more than fifty years ago by Birdsell (1951:63). The similarities of the first South Americans with sub-saharan Africans may result from the fact that the non-Mongoloid Southeast Asian ancestral population came, ulti- mately, from Africa, with no major modification in the original cranial bau plan of the first moderns. Neves et al. (1999c) have already suggested a possible historical connection among early modern humans (Skhul/Kafzeh), UC 101, Paleoindians and recent Africans and Australians based on their cranial morphology."

Marine Isotope Stage 3
"One bone, which came from a wild goat, was found in Zafarraya Cave in a similar layer as Neanderthal fossils. The bone was previously estimated as 33,300 years in age. However, using an ultrafiltration technique that cleansed the bone of modern carbon impurities that can give inaccurate younger dates, ... the bone was more than 46,700 years old."

Moershoofd interstadial
The Moershoofd interstadial has a 14C date of 44-46 kyr B.P. and corresponds to GIS 12 at 45-47 kyr B.P.

"Unearthed by an ivory carver from a Siberian riverbank, a man's 45,000-year-old thigh bone reveals when people first mated with Neanderthals [...] The Ust'-Ishim man's thigh bone is the oldest human bone found so far outside of Africa and the Middle East [...] It's nearly twice as old as the next oldest from a modern human, which comes from a boy who died elsewhere in Siberia some 24,000 years ago."

"A bone found by chance on the banks of a Siberian river has yielded the oldest modern human genome yet recovered, according to a new study that sheds light on when people left Africa and first interbred with Neanderthals living in Europe and Asia."

"The DNA narrows down the time when mating first brought Neanderthal genes into the human gene pool: from 50,000 to 60,000 years ago."

"The man, who lived 45,000 years ago, was definitely related to both humans and Neanderthals [...]. His DNA showed that the two human groups first mated around 60,000 years ago."

"The Ust'-Ishim man had about 2.3 percent Neanderthal genes, but modern people typically have less than 2.1 percent."

The "Siberian man belonged to a population that was closely related to the ancestors of today’s Europeans and Asians. He carried only slightly more Neanderthal DNA than they do."

“But his genomic segments of Neanderthal ancestry are on average about three times the length of those found in genomes today.”

This is highly informative “as the chunks of Neanderthal DNA have been gradually broken up each generation since the time of interbreeding.”

The "idea of a single time of human mating with Neanderthals is almost certainly [...] an oversimplification. The contacts could have extended over a longer period."

"The femur shaft turned up on the banks of the Irtysh River near Ust'-Ishim, Russia, in 2008. A Russian ivory carver and historian named Nikolay Peristov collected the bone after it eroded from a bluff above the river in western Siberia. It was identified as human, based on its teardrop-shaped cross section, in 2010."

“Thus the ancestors of Australasians (people from Australia, New Zealand, the island of New Guinea, and neighboring islands in the Pacific Ocean), with their similar input of Neanderthal DNA to Eurasians, must have been part of a late, rather than early, dispersal through Neanderthal territory.”

“While it is still possible that modern humans did traverse southern Asia before 60,000 years ago, those groups could not have made a significant contribution to the surviving modern populations outside of Africa, which contain evidence of interbreeding with Neanderthals.”

"Work on material from Italy seems to show human settlers pushing Neanderthals out (see maps). Mousterian tools were common there 45,000 years ago, when human-made Uluzzian material first appeared. By 44,000 years ago, humans were sharing Italy with a dwindling Neanderthal population. By 42,000 years ago, the Neanderthals were gone."

"Mousterian and Châtelperronian [artefacts] are probably Neanderthal. [...] Uluzzian, were once attributed to late Neanderthals, but recent work suggests they were made by humans (Nature, doi.org/bxh255)."

Denisovans
"The genome of ... the Denisovans that once interbred with us has been sequenced ... [A]bout 100,000 recent changes in our genome [have] occurred after the split from the Denisovans. ... DNA from fossils unearthed in Denisova Cave in southern Siberia in 2008 revealed a lineage unlike us and closely related to Neanderthals. The precise age of the Denisovan material remains uncertain — anywhere from 30,000 to 80,000 years of age. ... [T]he [Denisovans] lived in a vast range from Siberia to Southeast Asia. The Denisovans share more genes with people from Papua New Guinea than any other modern population studied."

"The picture of her genome [the girl from Denisova Cave] is as accurate as that of modern day human genomes, and shows she had brown eyes, hair and skin."

"The Denisovans have mysterious origins. They appear to have left little behind for palaeontologists save a tiny finger bone and a wisdom tooth found in Siberia's Denisova cave in 2010."

"This is an extinct genome sequence of unprecedented accuracy."

"For most of the genome we can even determine the differences between the two sets of chromosomes that the Denisovan girl inherited from her mother and father."

About "3% of the genes of people living today in Papua New Guinea come from Denisovans, with a trace of their DNA lingering in the Han and Dai people from mainland China."

A genetic "contribution from Denisovans is found exclusively in island Southeast Asia and Oceania (6)."

"Assuming 6.5 million years of sequence divergence between humans and chimpanzees, the shortening of the Denisovan branch allows the bone to be tentatively dated to between 74,000 and 82,000 years before present, in general agreement with the archaeological dates (2)."

Less "Denisovan allele-sharing [occurred] with the Dai than with the Han (although nonsignificantly so, Z = –0.9) [...]. Further analysis shows that if Denisovans contributed any DNA to the Dai, it represents less than 0.1% of their genomes today [...]."

"Denisovans share more alleles with the three populations from eastern Asia and South America (Dai, Han, and Karitiana) than with the two European populations (French and Sardinian) (Z = 5.3)."

Def. "the fraction of nucleotide sites that are different between a person’s maternal and paternal genomes" is called heterozygosity.

"The great apes have 24 pairs of chromosomes whereas humans have 23. This difference is caused by a fusion of two acrocentric chromosomes that formed the metacentric human chromosome 2 (25) and resulted in the unique head-to-head joining of the telomeric hexameric repeat GGGGTT. A difference in karyotype would likely have reduced the fertility of any offspring of Denisovans and modern humans. We searched all DNA fragments sequenced from the Denisovan individual and identified 12 fragments containing joined repeats. By contrast, reads from several chimpanzees and bonobos failed to yield any such fragments (8). We conclude that Denisovans and modern humans (and presumably Neandertals) shared the fused chromosome 2."

"In total, we identified 111,812 single-nucleotide changes (SNCs) and 9499 insertions and deletions where modern humans are fixed for the derived state, whereas the Denisovan individual carried the ancestral, i.e., ape-like, variant (8). This is a relatively small number. We identified 260 human-specific SNCs that cause fixed amino acid substitutions in well-defined human genes, 72 fixed SNCs that affect splice sites, and 35 SNCs that affect key positions in well-defined motifs within regulatory regions."

"One way to identify changes that may have functional consequences is to focus on sites that are highly conserved among primates and that have changed on the modern human lineage after separation from Denisovan ancestors. We note that, among the 23 most conserved positions affected by amino acid changes (primate conservation score of ≥0.95), eight affect genes that are associated with brain function or nervous system development (NOVA1, SLITRK1, KATNA1, LUZP1, ARHGAP32, ADSL, HTR2B, and CNTNAP2). Four of these are involved in axonal and dendritic growth (SLITRK1 and KATNA1) and synaptic transmission (ARHGAP32 and HTR2B), and two have been implicated in autism (ADSL and CNTNAP2). CNTNAP2 is also associated with susceptibility to language disorders (27) and is particularly noteworthy as it is one of the few genes known to be regulated by FOXP2, a transcription factor involved in language and speech development as well as synaptic plasticity (28). It is thus tempting to speculate that crucial aspects of synaptic transmission may have changed in modern humans."

"Of the 34 genes with clear associations with human diseases that carry fixed substitutions changing the encoded amino acids in present-day humans, four (HPS5, GGCX, ERCC5, and ZMPSTE24) affect the skin and six (RP1L1, GGCX, FRMD7, ABCA4, VCAN, and CRYBB3) affect the eye. Thus, particular aspects of the physiology of the skin and the eye may have changed recently in human history. Another fixed difference occurs in EVC2, which when mutated causes Ellis–van Creveld syndrome. Among other symptoms, this syndrome includes taurodontism, an enlargement of the dental pulp cavity and fusion of the roots, a trait that is common in teeth of Neandertals and other archaic humans. A Denisovan molar found in the cave has an enlarged pulp cavity but lacks fused roots (2). This suggests that the mutation in EVC2, perhaps in conjunction with mutations in other genes, has caused a change in dental morphology in modern humans."

Hasselo stadial
The "Hasselo stadial [is] at approximately 40-38,500 14C years B.P. (Van Huissteden, 1990)."

The "Hasselo Stadial [is a glacial advance] (44–39 ka ago)".

"Analysis of the jawbone of a man who lived about 40,000 years ago reveals the closest direct descendant of a Neanderthal who mated with a modern human."

"A modern human who lived in what is now Romania between 37,000 and 42,000 years ago had at least one Neanderthal ancestor as little as four generations back—which is to say, a great-great-grandparent."

“I could hardly believe that we were lucky enough to hit upon an individual like this.”

"The specimen, known as Oase 1, consists only of a male jawbone, and from the moment it was discovered in 2002 its shape suggested that it might belong to a hybrid between Homo sapiens and Neanderthal."

"The genome they sequenced from the samples was incomplete, but it was enough for the scientists to conclude that between 6% and 9% of Oase 1’s genome is Neanderthal in origin. People living today have 4% at most."

“We found seven huge pieces of chromosomes that seemed to be purely of Neanderthal origin. That means pieces had to come from a relatively recent ancestor, since they hadn’t yet been broken up by the reshuffling that happens in each generation as parents' chromosomes combine.”

"The non-Neanderthal genome sequences, meanwhile, show that Oase 1 isn’t related to humans living today. His genealogical line died out at some point."

Instead "of dying out 23,000 years ago, Neanderthals were gone as early as 39,000 years ago. It also looks like we shared their territory for 5000 years, steadily replacing them as we spread across Europe."

"Neanderthals had largely, and perhaps entirely, vanished from their known range by 39,000 years ago."

Hengelo interstadial
Preliminary DNA sequencing from a 38,000-year-old bone fragment of a femur found at Vindija Cave, Croatia, in 1980 showed Neanderthals and modern humans share about 99.5% of their DNA.

"DNA from the left shinbone of a skeleton, known as K14, which was excavated in 1954 [was analyzed]. K14 is one of the oldest fossils of a European modern human — a man who lived between 36,200 and 38,700 years ago in the area that's now Kostenki, in western Russia [whose skull is shown on the right]."

Kostenski is known for its mammoth structures, "circles made of mammoth bones that would have been the base of tents, huts, hearths, lithic and bone artifacts, as well as personal ornaments and figurines."

"K14's complete genome [was sequenced], making it the second-oldest modern human genome ever sequenced. The oldest yet was from the 45,000-year-old thighbone of a man found in western Siberia."

Contemporary "Europeans shared genetic continuity with ancient Europeans."

"Virtually all the major genetic components you find in contemporary Europeans are present among the earliest Europeans."

For "millennia, Europe may have been home to a so-called "metapopulation" of modern humans — a group of distinct, separate populations that regularly mixed, grew and fragmented. The genetic contributions of the earliest Eurasians to modern European populations may not have arrived through a few distinct migrations from Asia to Europe, but instead through gene flow in various directions."

"We have to revise our understanding of how the genetic diversity in contemporary Europeans came about. Early Europeans were part of a metapopulation stretching all the way to Central Asia, and through a complex network of sexual exchange, contemporary European populations were created."

Cro-Magnon
"Cro-magnons are, in informal usage, a group among the late Ice Age peoples of Europe. The Cro-Magnons are identified with Homo sapiens sapiens of modern form, in the time range ca. 35,000-10,000 b.p. [...] The term "Cro-Magnon" has no formal taxonomic status, since it refers neither to a species or subspecies nor to an archaeological phase or culture. The name is not commonly encountered in modern professional literature in English, since authors prefer to talk more generally of anatomically modern humans. They thus avoid a certain ambiguity in the label "Cro-Magnon", which is sometimes used to refer to all early moderns in Europe (as opposed to the preceding Neanderthals), and sometimes to refer to a specific human group that can be distinguished from other Upper Paleolithic humans in the region. Nevertheless, the term "Cro-Magnon" is still very commonly used in popular texts because it makes an obvious distinction with the Neanderthals, and also refers directly to people rather than to the complicated succession of archaeological phases that make up the Upper Paleolithic. This evident practical value has prevented archaeologists and human paleontologists - especially in continental Europe - from dispensing entirely with the idea of Cro-Magnons."

The earliest known remains of Cro-Magnon-like humans are radiocarbon dated to 43,000 years before present.

Cro-Magnons were robustly built and powerful. The body was generally heavy and solid with a strong musculature. The forehead was fairly straight rather than sloping like in Neanderthals, and with only slight browridges. The face was short and wide. Like other modern humans, Cro-Magnons had a prominent chin.

The brain capacity was about 1,600 cc, larger than the average for modern humans. However, recent research suggests that the physical dimensions of so-called "Cro-Magnon" are not sufficiently different from modern humans to warrant a separate designation.

"‘Cro-Magnon Man’ is commonly used for the modern humans that inhabited Europe from about 40,000 to 10,000 years ago."

Heinrich Event 4
Heinrich Event 4 "33-39.93 ka BP".

Huneborg interstadial
The Huneborg interstadial is a Greenland interstadial dating 36.5-38.5 kyr B.P. GIS 8.

Stadial
"Stadial duration 0.642 ka".

GIS 7 interstadial
"GIS 7 (start) 32.896 [to] GIS 7 (end) 32.15 ka BP".

Stadial
"Stadial duration 0.932 ka".

Ålesund Interstadial
The Ålesund interstadial began with GIS 6 and ended after GIS 8.

Stadial
Stadial duration 0.836 ka""

GIS 5
{{main|}History/Paleolithic/GIS 5|GIS 5}} GIS 5 interstadial occurred during the Klintholm advance about 33.5 kyr B.P.

"GIS 5 (start) 30.013 [to] GIS 5 (end) 29.526 ka BP".

Klintholm advance
This advance occurred after the Møn and ended with GIS 6.

"Stadial duration 2.899 ka".

Møn interstadial
The Møn interstadial corresponds to GIS 4.

Stadial
Heinrich Event 3 (H3) "occurs at 26.74 ka BP, coincident with the start of the transition into GIS 4."

MIS Boundary 2/3 is at 29 ka.

"Stadial duration 0.768 ka".

GIS 3
The stronger GIS 3 interstadial occurred about 27.6 kyr B.P.

It begins abruptly at 29 ka and ends about 26 ka.

"GIS 3 (start) 25.571 [to] GIS 3 (end) 25.337 ka BP".

Laugerie Interstadial
The weak interstadial corresponding to GIS 2 occurred about 23.2 kyr B.P.

"GIS 2 (start) 21.556 [to] GIS 2 (end) 21.407 ka BP".

Heinrich Event 2 (H2) extends "22-25.62 ka BP".

The δ18O values from GISP-2 follow the diagram of Wolfgang Weißmüller. The positions of the Dansgaard-Oeschger events DO1 to DO4 and the Heinrich events H1 to H3 are also indicated. DV 3-4 and DV 6-7 are cold events marked by ice wedges in the upper loess of Dolní Veštonice.

Jylland stade
"After c. 22 ka BP [which is] during the Jylland stade (Houmark-Nielsen 1989)".

Homo floresiensis
"The 18,000-year-old fossils of the extinct type of human officially known as Homo floresiensis were first discovered on the remote Indonesian island of Flores in 2003. Its squat, 3-foot-tall (1 meter) build led to the hobbit nickname."

"[T]he hobbit's brain was larger than previously suggested — 426 cubic cm (nearly 26 cubic inches), instead of the commonly cited figure of 400 cubic cm. (The modern human brain is 1,300 cubic centimeters, or 79 cubic inches, large on average.)"

Lascaux interstadial
The Lascaux interstadial begins about 21 ka and extends to about 18 ka.

Heinrich event H1
This stadial starts about 17.5 ka, extends to about 15.5 ka and is followed after a brief warming by H1.

Meiendorf Interstadial
The period spans starting at the far right of the image on the right from Lascaux interstadial to Heinrich event H1, and to Meiendorf/Bölling warm stage, and Allegöd warm stage, to Younger dryas and early holocene.

The Meiendorf Interstadial is typified by a rise in the pollens of dwarf birches (Betula nana), willows (Salix sp.), sandthorns (Hippophae), junipers (Juniperus) and Artemisia.

The beginning of the Meiendorf Interstadial is around 14,700 b2k.

Oldest Dryas
"During the Late Weichselian glacial maximum (20-15 ka BP) the overriding of ice streams eventually lead to strong glaciotectonic displacement of Late Pleistocene and pre-Quaternary deposits and to deposition of till."

The "beginning of Oldest Dryas time [is] (~14,600 14C yr BP)".

"More than 15,000 years ago, humans began crossing a land bridge called Beringia that connected their native home in Eurasia to modern-day Alaska. Who knows what the journey entailed or what motivated them to leave, but once they arrived, they spread southward across the Americas."

"The prevailing theory is that the first Americans arrived in a single wave, and all Native American populations today descend from this one group of adventurous founders. But now there’s a kink in that theory. The latest genetic analyses back up skeletal studies suggesting that some groups in the Amazon share a common ancestor with indigenous Australians and New Guineans. The find hints at the possibility that not one but two groups migrated across these continents to give rise to the first Americans."

“Our results suggest this working model that we had is not correct. There’s another early population that founded modern Native American populations.”

"Genetic studies have since connected both these ancient and modern humans to ancestral populations in Eurasia, adding to the case that a single migratory surge produced the first human settlers in the Americas. Aleutian Islanders are a notable exception. They descend from a smaller second influx of Eurasians 6,000 years ago that bear a stronger resemblance to modern populations, and some Canadian tribes have been linked to a third wave."

The "Suruí and Karitiana people of the Amazon had stronger ties to indigenous groups in Australasia—Australians, New Guineans and Andaman Islanders—than to Eurasians."

They "scrutinized the genomes of 30 Native American groups in Central and South America. Using four statistical strategies, they compared the genomes to each other and to those of 197 populations from around the world. The signal persisted. Three Amazonian groups—Suruí, Karitiana and Xavante—all had more in common with Australasians than any group in Siberia."

"The DNA that links these groups had to come from somewhere. Because the groups have about as much in common with Australians as they do with New Guineans, the researchers think that they all share a common ancestor that lived tens of thousands of years ago in Asia but that doesn’t otherwise persist today. One branch of this family tree moved north to Siberia, while the other spread south to New Guinea and Australia. The northern branch likely migrated across the land bridge in a separate surge from the Eurasian founders. The researchers have dubbed this hypothetical second group “Population y” for ypykuéra, or “ancestor” in Tupi, a language spoken by the Suruí and Karitiana."

Studies "of ancient skulls unearthed in Brazil and Colombia [...] bear stronger resemblance to those of Australasians than the skulls of other Native Americans. Based on the skeletal remains, some anthropologists had previously pointed to more than one founding group".

"We postulate a putative time of entry of this “Australo-Melanesian-like” population in the New World around 14 Ka (see also Dillehay, 2000; Dixon, 2001). This population expanded southward in the New World using the Pacific coast. Eventually, when the recession of the ice sheets in North America permitted, this population expanded towards the interior. A rapid expansion along the coast would explain why the Australo-Melanesian cranial bau plan of the first settlers was not altered during the expansion into South America. If we see the Australo-Melanesian cranial morphology as the result of selection under tropical climates since its inception in Africa (Lahr, 1996), a relatively fast migration along the Pacific Rim could explain why this “tropical pattern” was not significantly altered by colder climates."

Bølling Oscillation
The "intra-Bølling cold period [IBCP is a century-scale cold event and the] Bølling warming [occurs] at 14600 cal [calendar years, ~ b2k] BP (12700 14C BP)".

Older Dryas
"Older Dryas [...] events [occurred about 13.4 b2k]".

Mesolithic
The mesolithic period dates from around 13,000 to 8,500 b2k.

"These hunter-gatherers were some of the earliest known residents of South America and they chose to live at this extreme altitude — higher than any Ice Age encampment found thus far in the New World. Despite the thin air and sub-freezing night-time temperatures, this plain would have seemed a hospitable neighbourhood to those people."

“The basin has fresh water, camelids, stone for toolmaking, combustible fuel for fires and rock shelters for living in. Basically, everything you need to live is here. This is one of the richest basins I've seen, and it probably was then, too.”

"150 kilometres away from the Andes cave, on Peru's arid coast at Quebrada Jaguay [is] a site that dated to the end of the last Ice Age, between 13,000 and 11,000 years ago."

At "the end of the last Ice Age, glaciers were mainly continued to alpine valleys, and [the] Pucuncho [plateau] and other areas were not glaciated."

"Paleoclimate data indicate that the environment was probably wetter then, so there might have been more plants and animals available for the early residents."

“These Palaeo-Indians were able to live in one of the most extreme environments on Earth, at the end of an ice age, and they seem to have done so quite successfully. This tells us that Palaeo-Indians were capable of living just about anywhere.”

An "ancient encampment, [is] dated to around 13,000 years ago4. [It is] called Quebrada Santa Julia [see the image above.] Some of the stone tools at the site [...] were made of translucent quartz that is not found in coastal deposits."

Beneath "Huaca Prieta, a 32-metre-high mound on the coast of northern Peru [...] traces of Ice Age settlements [...] Radiocarbon dating indicates5 that humans had lived there as much as 14,200 years ago, when the area was surrounded by wetlands."

"Some tools found at a site called Pay Paso are made of translucent agate, which apparently came from quarries near the border with Brazil about 150 kilometres away. And other tools from Uruguay have been found 500 kilometres to the south in Argentina's Buenos Aires province6."

"12,800-year-old artefacts at Cueva Bautista, a rock shelter 3,930 metres above sea level in southwestern Bolivia. [A] similarly aged site exists at the same latitude in Chile on the western slope of the Andes."

There is "an outcrop of translucent quartz at a site where people had lived and quarried between 12,600 and 11,400 years ago. The similarity with Quebrada Santa in terms of age and tool-making techniques suggests that the coastal tools came from these mountain outcrops."

People "built a fire in the rock shelter, named Cuncaicha [image at the right], about 12,400 years ago."

"At the Panama Isthmus, the coastal expansion trifurcated, with one migration following down the Pacific coast, another the Atlantic coast, and a third one inward into the Amazon basin. These multiple axes south of Panama would explain the presence of humans in Southern Chile around 12.3 Ka (Dillehay, 1989, 1997), the presence of humans in Lagoa Santa and elsewhere in eastern Central Brazil around 12 Ka (Kipnis, 1998, Prous and Fogac ̧a, 1999), and in the Amazon around 11.2 Ka (Roosevelt et al., 1996). A Pacific and an Atlantic expansion would also explain the presence of Paleoindians with Australo-Melanesian morphology on opposite sides of South America, in eastern Central Brazil and the Colombia Highlands, by the end of the Pleistocene (Neves and Pucciarelli, 1989; 1991; Munford, 1999)."

Neolithic
The base of the Neolithic is approximated to 12,200 b2k.

“What we're seeing is that 12,000 years ago or more, these groups already had networks, knew the landscape and moved between the coast and the interior.”

At the left is a reconstruction of the face of one of the oldest human remains found in the Americas. It is dated to 11,500 years ago.

Allerød Oscillation
The "Allerød Chronozone, 11,800 to 11,000 years ago".

"Kamminga and Wright (1988), Wright (1995) and Neves and Pucciarelli (1998) have demonstrated, however, that the Zhoukoudian Upper Cave (UC) cranium 101 display marked similarities with Australo-Melanesians. Cunningham and Wescott (2002) has shown that although highly variable, none of the three specimens from this site (UC 101, UC 102, UC 103) resembles modern Asian populations. Matsumura and Zuraina (1999:333) reported the presence of the “Australo-Melanesian lineage” in Malaysia as late as the terminal Pleistocene. If we consider that UC is dated to between 32,000 BP and 11,000 BP, the fixation of the classical Mongoloid morphology in North Asia could have been a recent phenomenon (terminal Pleistocene/early Holocene), a hypoth- esis favored by several authors (see Cunningham and Wescott, 2002 for a review)."

"Accordingly, an Australo-Melanesian-like population present in North Asia by the end of the Pleistocene could have been the source of the first Americans. This would explain the presence of a non-Mongoloid morphology in the New World without invoking a direct transpacific route departing from Australia, as suggested by Rivet (1943)."

"Lahr (1995) has argued that human diversity in northern Asia was probably higher in the final moments of the Pleistocene than today, at least as far as cranial morphology is concerned. Therefore, non-Mongoloid Asians could have arrived in the Americas using the Behring Strait as the gate of entry following either the shore of Beringia or a land bridge."

Holocene
The Holocene starts at ~11,700 b2k and extends to the present.

Younger Dryas
The "Alleröd/Younger Dryas transition [occurred] some 11,000 years ago [11,000 b2k]."

Pre-Boreal transition
The last glaciation appears to have a gradual decline ending about 12,000 b2k. This may have been the end of the Pre-Boreal transition.

There is "a cranial morphology for terminal Pleistocene and Early Holocene populations in the Americas outside the range of variation of modern Amerindians, both in North and South America (Jantz and Owsley, 2001). While in North America the early human remains are, in general, but not exclusively, more similar to South Asians, Ainu/ Polynesians or Europeans (Steele and Powell, 1992, 1994, 1999; Chatters et al., 1999; Brace et al., 2001; Jantz and Owsley, 2001), in South America they are closer to Australians and sub-Saharan Africans (Neves and Pucciarelli, 1989, 1990, 1991; Munford et al., 1995; Neves et al., 1998, 1999a,b)."

"About 9000 years ago the temperature in Greenland culminated at 4°C warmer than today. Since then it has become slowly cooler with only one dramatic change of climate. This happened 8250 years ago [...]. In an otherwise warm period the temperature fell 7°C within a decade, and it took 300 years to re-establish the warm climate. This event has also been demonstrated in European wooden ring series and in European bogs."

"The last remains of the American ice sheet disappeared about 6000 years ago [6,000 b2k], the Scandinavian one 2000 years earlier [8,000 b2k]."

"Santana do Riacho is a late Paleoindian burial site where approximately 40 individuals were recovered in varying states of preservation. The site is located at Lagoa Santa/Serra do Cipó, State of Minas Gerais. The first human activities in this rockshelter date back to the terminal Pleistocene, but the burials are bracketed between circa 8200 and 9500 BP. The collection contains only six skulls well-enough preserved to be measured. The Santana do Riacho late Paleoindians present a cranial morphology characterized by long and narrow neurocrania, low and narrow faces, with low nasal apertures and orbits. The multivariate analyses show that they exhibit strong morphological affinities with present day Australians and Africans, showing no resemblance to recent Northern Asians and Native Americans."

The image on the right shows two of these skulls. On the left is a female and the right is a male.

"The similarities of the first South Americans with sub-Saharan Africans may result from the fact that the non-Mongoloid Southeast Asian ancestral population came, ultimately, from Africa, with no major modification in the original cranial bau plan of the first modern humans."

Ancient history
The ancient history period dates from around 8,000 to 3,000 b2k.

"All in all, Europeans maintained genetic continuity from their earliest establishment out of Africa until Middle Eastern farmers arrived in the last 8,000 years, bringing with them agriculture and a lighter skin color."

"Ancient DNA has been retrieved and analyzed from Egyptian mummies".

Copper Age
The copper age history period began from 6990 b2k.

The "oldest securely dated evidence of copper making, from 7,000 years ago [6990 b2k], at the archaeological site of Belovode, Serbia."

The "Scandinavian one 2000 years earlier [8,000 b2k]."

Boreal transition
"In some cores a narrow band of clay interrupts the organic muds, at the horizon of the Boreal Atlantic transition."

Atlantic history
The "Atlantic period [is] 4.6–6 ka [4,600-6,000 b2k]."

Bronze Age
A general world-wide use of bronze occurred between 5300 and 2600 b2k.

"The first (purely typological) studies on Early Bronze Age (EBA) assemblages in the Jordan Valley settled on the turn of the 4th/3rd millennium BC [mark] the beginnings of the earliest Bronze Age culture (Albright 1932; Mallon 1932)."

"In the Chalcolithic/earliest Bronze Age I period (c. 4500±3000 cal BC), copper was mined in open galleries from the massive brown sandstone deposit, which consisted of thick layers of the copper carbonate malachite and chalcocite, a copper sulphide."

Iron Age
The iron age history period began between 3,200 and 2,100 b2k.

"The earliest known iron artefacts are nine small beads securely dated to circa 3200 BC, from two burials in Gerzeh, northern Egypt."

"Since both tombs are securely dated to Naqada IIC–IIIA, c 3400–3100 BC (Adams, 1990: 25; Stevenson, 2009: 11–31), the beads predate the emergence of iron smelting by nearly 2000 years, and other known meteoritic iron artefacts by 500 years or more (Yalçın 1999), giving them an exceptional position in the history of metal use."

The image on the left uses neutron radiography to show the metal underneath the corrosion.

"Bead UC10738 [in the image on the right] has a maximum length of 1.5 cm and a maximum diameter of 1.3 cm, bead UC10739 is 1.7 cm by 0.7 cm, and bead UC10740 is 1.7 cm by 0.3 cm. All three beads are of rust-brown colour with a rough surface, indicative of heavy iron corrosion. Initial analysis by [proton–induced X–ray fluorescence] pXRF indicated an elevated nickel content of the surface of the beads, in the order of a few per cent, and their magnetic property suggested that iron metal may be present in their body (Jambon, 2010)."

Early history
The early history period dates from around 3,000 to 2,000 b2k.

In the image on the right are Guanches engravings in a rock cave on La Palma island of the Canary Islands.

The Guanches are believed to be the original inhabitants of the Canary Islands perhaps as early as 3,000 b2k.

Subboreal history
The "period around 850-760 BC [2850-2760 b2k], characterised by a decrease in solar activity and a sharp increase of Δ 14C [...] the local vegetation succession, in relation to the changes in atmospheric radiocarbon content, shows additional evidence for solar forcing of climate change at the Subboreal - Subatlantic transition."

The "Holocene climatic optimum in this interior part of Asia [Lake Baikal] corresponds to the Subboreal period 2.5–4.5 ka".

Subatlantic history
The "calibration of radiocarbon dates at approximately 2500-2450 BP [2500-2450 b2k] is problematic due to a "plateau" (known as the "Hallstatt-plateau") in the calibration curve [...] A decrease in solar activity caused an increase in production of 14C, and thus a sharp rise in Δ 14C, beginning at approximately 850 cal (calendar years) BC [...] Between approximately 760 and 420 cal BC (corresponding to 2500-2425 BP [2500-2425 b2k]), the concentration of 14C returned to "normal" values."

Imperial Antiquity
Imperial Antiquity lasts from 2,000 to 1,700 b2k.

In Felix Romuliana, "the construction [...] is [...] Imperial Antique (1st-3rd c.), and sometimes even late Hellenistic, [in] appearance."

High Middle Ages
The High Middle Ages date from around 1,000 b2k to 700 b2k.

Mitochondrial "DNA analysis (HVRI sequences and RFLPs) [have been performed from] aborigine remains around 1000 years old. The sequences retrieved show that the Guanches possessed U6b1 lineages that are in the present day Canarian population, but not in Africans. In turn, U6b, the phylogenetically closest ancestor found in Africa, is not present in the Canary Islands. Comparisons with other populations relate the Guanches with the actual inhabitants of the Archipelago and with Moroccan Berbers. This shows that, despite the continuous changes suffered by the population (Spanish colonisation, slave trade), aboriginal mtDNA lineages constitute a considerable proportion of the Canarian gene pool. Although the Berbers are the most probable ancestors of the Guanches, it is deduced that important human movements have reshaped Northwest Africa after the migratory wave to the Canary Islands."

The "sublineage U6b1 is the most prevalent of the U6 subhaplogroup in the Canarian population,4 and has still not been detected in North Africa."

"This survey includes 131 teeth, corresponding to 129 different individuals, belonging to 15 archaeological sites sampled from four of the seven Canary Islands and dated around 1000 years old [image on the right]."

"The Canarian-specific U6b1 sequences are also found in high frequency (8.45%), corroborating the fact that these lineages were already present in the aboriginal population. Three additional founder haplotypes4 were also detected (260, 069 126 and 126 292 294), all of them showing equal or higher frequencies than in the present day Canarian population."

"The detection in the Guanches of the most abundant haplotype of the U6b1 branch, also found in present day islanders,4 points to a significant continuity of the aboriginal maternal gene pool."

"The [...] estimated age of the [U6b1] subgroup is around 6000 years,29 which predates the arrival of the first human settlers to the Islands.1"

Guanches in 1496
The image on the right hangs in the interior of the ayuntamiento of San Cristobal de La Laguna, Tenerife.

The painting on the right shows the surrender of the Guanches kings of Tenerife to Ferdinand and Isabella. This appears to have occurred c. 504 b2k.

The painting on the left was painted in 1764. It depicts the surrender of the Guanches leaders Bencomo mencey with Tacoronte, Anaga and Tegueste to Governor Alonso Fernández de Lugo with his captains and noble friends, by bringing gifts to the governor.

Ainu in 1860s
The Ainu are a people inhabiting the Northern island of Yesso. They differ from the Japanese in language and race. Their origin is lost in a wild and fabulous tradition. The legend runs thus—“That the race owes its preservation to a doll which swam across from Corea to the uninhabited island of Yesso.” They were conquered some three hundred years ago by the Japanese.

The colorized photograph on the right is from between 1863 and 1870 (137-130 b2k).

The map shows their widest expanse historically.

Noncoding DNA
More than 98% of the human genome does not encode protein sequences, including most sequences within introns and most intergenic DNA.

Fully 98% of the human genome is noncoding DNA.

Over 80% of DNA in the human genome "serves some purpose, biochemically speaking".

Non-coding repetitive sequences
Over 50% of human DNA consists of non-coding repetitive sequences.

Non-coding RNA sequences
Some DNA sequences that do not code protein may still encode functional non-coding RNA molecules, which are involved in the regulation of gene expression.

Pseudogenes
An abundant form of noncoding DNA in humans are pseudogenes, which are copies of genes that have been disabled by mutation. These sequences are usually just molecular fossils, although they can occasionally serve as raw genetic material for the creation of new genes through the process of gene duplication and divergence.

Genes
Def. a "unit of heredity; a segment of DNA or RNA that is transmitted from one generation to the next, and that carries genetic information such as the sequence of amino acids for a protein" is called a gene.

The genetic information in a genome is held within genes, and the complete set of this information in an organism is called its genotype. A gene is a unit of heredity and is a region of DNA that influences a particular characteristic in an organism. Genes contain an open reading frame that can be transcribed, as well as regulatory sequences such as promoters and enhancers, which control the transcription of the open reading frame.

Only about 1.5% of the human genome consists of protein-coding exons.

Telomeres
Some noncoding DNA sequences such as telomeres and centromeres play structural roles in chromosomes.

Telomeres are usually lengths of single-stranded DNA containing several thousand repeats of a simple TTAGGG sequence.

Telomeres and centromeres typically contain few genes, but are important for the function and stability of chromosomes.

Centromeres
Centromeres are chromosomal loci that ensure delivery of a copy of a chromosome to each daughter upon cell division. On the Spindle Apparatus, chromosome movement is run and maintained by the centromere during meiosis and mitosis.

Introns
An intron is any nucleotide sequence within a gene that is removed by RNA splicing while the final mature RNA product of a gene is being generated. The term intron refers to both the DNA sequence within a gene and the corresponding sequence in RNA transcripts.

There are several families of internal nucleic acid sequences that are not present in the final gene product, including inteins, untranslated sequences (Untranslated region UTR), and nucleotides removed by RNA editing, in addition to introns.

Introns are extremely common within the nuclear genome of higher vertebrates (e.g. humans and mice), where protein-coding genes almost always contain multiple introns.

Some introns themselves encode specific proteins or can be further processed after splicing to generate noncoding RNA molecules. Alternative splicing is widely used to generate multiple proteins from a single gene. Furthermore, some introns represent mobile genetic elements and may be regarded as examples of selfish DNA.

The human genome contains an average of 8.4 introns/gene (139,418 in the genome).

Some introns are known to enhance the expression of the gene that they are contained in by a process known as intron-mediated enhancement (IME).

Geography
The specifics of Paleo-Indian migration to and throughout the Americas, including the exact dates and routes traveled, are subject to ongoing research and discussion.

Americas
The map at the right is a schematic illustration of maternal geneflow in and out of Beringia. Colours of the arrows correspond to approximate timing of the events and are decoded in the coloured time-bar. The initial peopling of Berinigia (depicted in light yellow) was followed by a standstill after which the ancestors of indigenous Americans spread swiftly all over the New World while some of the Beringian maternal lineages–C1a-spread westwards. More recent (shown in green) genetic exchange is manifested by back-migration of A2a into Siberia and the spread of D2a into north-eastern America that post-dated the initial peopling of the New World.

South America
Genetic history of indigenous peoples of the Americas primarily focus on Human Y-chromosome DNA haplogroups and Human mitochondrial DNA haplogroups. "Y-DNA" is passed solely along the patrilineal line, from father to son, while "mtDNA" is passed down the matrilineal line, from mother to offspring of both sexes. Neither recombines, and thus Y-DNA and mtDNA change only by chance mutation at each generation with no intermixture between parents' genetic material. Autosomal "atDNA" markers are also used, but differ from mtDNA or Y-DNA in that they overlap significantly. AtDNA is generally used to measure the average continent-of-ancestry genetic admixture in the entire human genome and related isolated populations.

"The traditional Western theory has been that these early migrants moved into the Beringia land bridge between eastern Siberia and present-day Alaska around 40,000—16,500 years ago,   when sea levels were significantly lowered due to the Quaternary glaciation.  These people are believed to have followed herds of now-extinct Pleistocene megafauna along ice-free corridors that stretched between the Laurentide and Cordilleran ice sheets. Another route proposed is that, either on foot or using primitive boats, they migrated down the Pacific Northwest coast to South America. Evidence of the latter would since have been covered by a sea level rise of hundreds of meters following the last ice age. Some recent DNA studies suggest additional migration from Europe around the northern fringe of the Atlantic possibly as long ago as either 36,000 to 23,000 years ago or between 17,000 to 12,000 years. However, this is also attributed to admixture of Europeans into northern Asia before the Beringian migration. Recent genetic studies have shown that that Paleolithic Europeans and Native Americans share a genetic founder population and that there is strong evidence that the "population that crossed the Bering Strait from Siberia into the Americas more than 15,000 years ago was likely related to the ancient population of Europe."

"Canada's oldest known home is a cave in Yukon occupied not 12,000 years ago like the U.S. sites, but at least 20,000 years ago"

"However, despite the lack of this conclusive and widespread evidence, there are suggestions of human occupation in the northern Yukon about 24,000 years ago, and hints of the presence of humans in the Old Crow Basin as far back as about 40,000 years ago."

A "site in southern Chile called Monte Verde [has evidence of] human occupation1 that [is] dated to about 14,500 years ago."

Hypotheses

 * 1) There is at least one isoform in hominin DNA that makes Homo sapiens sapiens unique.