Denisova man

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The Denisova humans were a population of the genus Homo , which is closely related to the Neanderthals and how they are similar to anatomically modern humans ( Homo sapiens ), but can be genetically differentiated from both species . In the English-language specialist literature they are called Denisova hominins or Denisovans for short .

In 2010, Johannes Krause and Svante Pääbo from the Max Planck Institute for Evolutionary Anthropology in Leipzig initially succeeded in evaluating the DNA from the mitochondria (the mtDNA ) of a finger bone using DNA sequencing . The announcement of the results of this DNA analysis caused a worldwide sensation, as the fossil was interpreted as evidence of a previously unknown population of the genus Homo , which was close to the Neanderthals and anatomically modern humans . A few months later the analysis of the DNA from the cell nuclei of the bone was also published; it confirmed the relative independence of the Denisova population. Accordingly, in addition to the previously known populations of Neanderthals and Homo floresiensis, there was a third community of distant (but clearly belonging to the genus Homo ) relatives of anatomically modern humans. The Denisova fossils are most closely related to the Neanderthal finds from the Vindija Cave and the Mesmaiskaya Cave . The assignment of the finds from the Denisova cave to a new species or subspecies was expressly waived in 2010; In 2011, however, the fossils were attributed to "a previously unknown species".

Finger joint discovered in 2008 in the Denisova Cave (replica), Museum of Natural Sciences in Brussels, Belgium Molar (original) discovered in the Denisova cave in 2000, exhibited in the special exhibition "Le troisième Homme" at the Musée national de Préhistoire in Les Eyzies-de-Tayac, France
Fragment of the phalanx ( replica ), according to the technical description phalanx distalis , but placed here as phalanx media
Molar tooth Denisova 4 (original)

Denisova people lived in the Altai Mountains in southern Siberia until 76,000–52,000 years ago - during the Paleolithic Age - and in Tibet around 160,000 years ago . The existence of this population has so far only been confirmed by a few, small fossils from the Denisova cave : u. a. through the bone of a little finger , through two posterior molars, and through a lower jaw from Tibet.

Find history and dating

The location of the Altai Mountains in Central Asia

The excavations in the Denisova Cave ( Russian Денисова пещера - Denissowa peschtschera, actually: "Cave of Denis") near the border with Kazakhstan were carried out by the Natural History Museum of Novosibirsk under the direction of the two archaeologists Mikhail Schunkow and Anatoly Derewjanko from the Russian Academy Sciences . The cave had been extensively explored since the 1970s, following the uncovering of Moustérien and Levallois- style stone tools attributed to Neanderthals. Several different old references to the use of the cave by prehistoric humans ( find horizons ) could be distinguished from one another.

In 2000, employees of the Russian research group exposed the first molar, the fossil Denisova 4 ; however, one could not assign him to any particular homosexual with certainty . The 48,000-30,000-year-old phalanx (the fossil Denisova 3 , a distal phalanx), which probably comes from a five- to seven-year child was discovered of 2008. 2011, the research team was the discovery of an outer phalange bone of the left foot known that 130,000 to 90,900 year-old fossil Denisova 5 , but which was subsequently recognized to be of a Neanderthal arise.

In 2015, another molar was found, the fossil Denisova 8 .

In March 2017, Science described fragments of two skullcaps from Lingjing (灵 井), Xuchang (许昌), People's Republic of China , that belong together and that are 125,000 to 105,000 years old and, due to their characteristics, can possibly be assigned to the Denisova people. The site has been known internationally since 2008; the first skull fragments discovered in 2007 have so far been referred to as Xuchang people in China .

At the beginning of 2019, two studies with dates on the settlement of the cave based on optically stimulated luminescence and a variant of mass spectrometry (ZooMS) were published in the journal Nature . According to this, the oldest evidence for Denisova people is at least 200,000 years old. According to the analysis, the “mixed race child” Denisova 11 is around 100,000 years old. The most recent bone find (Denisova 14) is 46,300 ± 2600 years old, but its attribution to the Denisova people is not considered certain, so that the most recent evidence of the existence of the Denisova people is 76,000–52,000 years old. The cave was the According to findings not continuous but episodic - especially during the interglacials populated -.

The lower jaw from Tibet presented in 2019 is around 160,000 years old.

Analysis of mtDNA from the finger bone

Johannes Krause, an expert in the analysis of Neanderthal DNA, had extracted enough DNA from mitochondria from 30 milligrams of powdered material from the finger bone to be able to completely reconstruct its blueprint (the nucleotide sequence of the mtDNA). Then this mtDNA sequence was compared with that of 54 people living today ( Homo sapiens ), furthermore with the mtDNA sequence of a Young Pleistocene person from Kostjonki 14 on the Don (southern Russia), with the complete mtDNA sequences of six Neanderthals and - as so-called Outgroup , because so far no DNA from Homo erectus / Homo heidelbergensis could be obtained - with the mtDNA of a chimpanzee and a bonobo . While Neanderthals and anatomically modern humans differ on average at 202 nucleotide positions in the mtDNA, the number of discrepancies between the find from the Denisova Cave and the anatomically modern humans is almost twice as large at 385.

By comparing these data with the deviations between humans and chimpanzees (1462 positions), it was estimated that the lines of development of Denisovans and anatomically modern humans separated as early as 1,314,000 to 779,000 years ago, while the lines of development of Homo sapiens diverged and Neanderthals finally separated only 618,000 to 321,000 years ago. From this it was concluded that in the Altai around 60,000 years ago there was, besides Homo sapiens and the Neanderthals, a third population of the genus Homo that immigrated there independently of these two species .

How reliable the dating of family relationships is based on the mtDNA alone is controversial, since the mitochondria are inherited exclusively through the mother without recombination . They are therefore particularly exposed to gene drift and gene flow , for example , which means that a relatively large number of changes can occur in a short time; In contrast to this, the cell nucleus DNA has tens of thousands of gene loci that are " evolution-neutral " and therefore change less rapidly (and less discontinuously).

In 2019 it was possible to obtain the mtDNA of a bone fragment and to compare it with the mtDNA of the fossil Denisova 3, with the result that this fragment was recognized as belonging to Denisova 3. A virtual reconstruction showed that this finger bone is very similar to that of an anatomically modern human, but not that of a Neanderthal man, so that the structure of the finger seems to be a plesiomorphic feature.

Analysis of the DNA from cell nuclei of the finger bone

The Leipzig researchers had already announced in March 2010 that, following the mtDNA, they would also sequence the complete DNA from the cell nuclei of the fossil. It was already clear at the time that the fossil unofficially named "X-Woman" and described as a "girl" by the Leipzig researchers did not have a Y chromosome , i.e. it was a female child. The Leipzig research team finally published the entire genome sequence from the nucleus of the Denisova people online on February 8, 2012, making it freely accessible to everyone. The cell nucleus DNA of the finger bone turned out to be unusually well preserved. An improvement in the investigation technique had made it possible to sequence each base within the Denisova genome thirty times. The DNA required for this was obtained from less than ten milligrams of the finger bone. The current resolution even shows the differences between the gene copies that the individual had inherited from his mother and father.

Relationship with the Neanderthals

As early as December 2010, it was reported that the DNA differences between Neanderthals and Denisovans indicated a final separation of the two populations 640,000 years ago and a final separation of their common ancestors from the ancestors of Homo sapiens around 800,000 years ago. According to these data, the Denisovans are more closely related to the Neanderthals than to the anatomically modern human, Homo sapiens , which differs significantly from the interpretation of the mtDNA findings . The results of such calculations are, however, controversial among experts, because the exact rate of the molecular clock , i.e. the frequency of mutations in past epochs, are only estimated.

A comparison of the DNA of Neanderthal finds from the Vindija Cave and the Mesmaiskaya Cave revealed an unusually large genetic proximity of both finds and a relatively large genetic distance between both finds and the Denisova fossil. From this it was concluded on the one hand that Neanderthals and Denisovans were two genetically isolated populations for a long time, but that they are more closely related to one another than to Homo sapiens ; on the other hand, that the Neanderthals went through a genetic bottleneck after separating from the ancestors of the Denisova population - severe genetic impoverishment had already been inferred from the analysis of the mtDNA of Neanderthals, as their genetic variability is much lower than the genetic variability of anatomically modern man . Due to these peculiarities, a prehistoric population of the genus Homo was separated for the first time from related populations - in analogy to Neanderthals called Denisovans in English - solely on the basis of molecular biological data .

However, the calculated, long-lasting genetic isolation of the Neanderthal populations from those of the Denisovans did not prevent the generation of common offspring at least 50,000 years ago. At least this is the result of a study that was published in 2018. According to this, it was possible to extract and sequence DNA from the Denisova 11 fossil - a small fragment of a long bone that was discovered in the Denisova Cave in 2012. The fossil belonged to a youngster who was probably at least 13 years old, whose mother was a Neanderthal woman and whose father was a Denisovan. Further analyzes of the genome showed that the woman's father also had at least one Neanderthal man among his ancestors. The researchers also found that the mother was genetically more closely related to Neanderthals who lived in Western Europe than to a Neanderthals who had previously lived in Denisova Cave. This shows that the Neanderthals migrated between West and East Eurasia tens of thousands of years before they disappeared. It is possible that Neanderthal-Denisova crossbreeds were mated in Asia with the groups of Homo sapiens that migrated from Africa .

Gene flow to Homo sapiens

A study was published in May 2010 that showed a gene flow from the Vindija Neanderthals to Homo sapiens . Therefore, the genetic distance of the Denisova fossil to today's living ethnic groups was analyzed, using data from 938 people from 53 populations. According to the findings, the Denisova fossil is further away from the European, Asian and African people living today than the Neanderthals. In contrast, a significant proximity to the DNA of people from Melanesia ( Papua and residents of Bougainville ) was found. This led to the statement that 2.5 ± 0.6 percent of the Melanesians' genome - like that of all non-African people - came from Neanderthals, but that an additional 4.8 ± 0.5 percent was contributed by Denisova people ; In total, according to the study, this would be 7.4 ± 0.8 percent of the genome of the Melanesians, which came from an earlier intermingling with archaic hominins . From the purely regional distribution of the Denisova DNA it was deduced that there could not have been frequent mixing.

In September 2011, further genetic findings were published, which were now based on a comparison of the DNA of 33 populations living today from Asia and Oceania with that of the Denisova fossil. According to this, DNA traces of the Denisova people could also be found in the Aborigines in Australia, the Mamanwas in the Philippines and in the east of Indonesia , but not in the west of Indonesia and not in the Onge in the Andamans , in the Jehai in Malaysia and among populations in East Asia . The authors of this study interpreted the detection of Denisova DNA in eastern Indonesia, Australia, Papua New Guinea , Fiji, and Polynesia as evidence that the genetic intermingling occurred in Southeast Asia , which would mean that the Denisova people are an area between Siberia and the tropics. This interpretation is controversial, however, since early migrations of ancestors of the ethnic groups examined cannot be ruled out and the sexual contacts could therefore also have taken place further north - in the Asian heartland.

A more detailed analysis of the Denisova DNA in 2012 showed, among other things, that alleles could be detected "which in people living today are associated with dark skin, brown hair and brown eyes". It was also possible to evaluate parts of the genetic make-up from father and mother separately. From this it was concluded that the extent of heterozygosity was very low, only 0.022% ; this corresponds to "approximately 20% of the value of today's Africans, around 26 to 33% of today's Eurasians and 36% of the Karitiana , an indigenous population living in Brazil with extremely low heterozygosity". An adaptation that has been proven in the Inuit of Greenland , which enables them to utilize fat better and convert it into body heat more easily than is possible for people in other populations today, was interpreted in 2016 as a possible introgression .

The interpretation of the findings from Neanderthal DNA as a gene flow from Neanderthals to Homo sapiens was repeatedly criticized in 2012 on the basis of model calculations: The greater correspondence of the genome of the non-African populations of Homo sapiens with the genome of Neanderthals could also be explained by the fact that coincidentally a population of Homo sapiens left Africa, which still had a particularly great genetic similarity to the common ancestor of the anatomically modern humans and the Neanderthals. These objections can be applied to the Denisova people.

In February 2020, a study was published, according to which gene flow from a previously unidentified, archaic homo- population to the common ancestors of Neanderthals and Denisovans, whose genetic markers increase today - as a result of later gene flow, was published Homo sapiens - are also detectable in anatomically modern humans.

Molar morphology and DNA

The almost completely preserved molar (a M3 or M2 molar from the left area of ​​an upper jaw ) discovered in 2000 was also assigned to the Denisova people in 2010 due to its mtDNA, but to a different individual than the finger bone. The tooth is extraordinarily large, larger than the molars of Neanderthals and anatomically modern humans: mesiodistal (from front to back) 13.1 mm, buccolingual 14.7 mm (from outside to inside; in Homo sapiens : mesiodistal approx. 10– 10.5 mm; buccolingually approx. 9.5–10 mm). If it was an M2 molar, it would be similar in size to the corresponding molar of Homo erectus and Homo habilis ; if it was a M3 molar, it would be similar in size to the corresponding molar of Homo habilis or Homo rudolfensis and comparable to the M3 molar of an Australopithecus . There are no similarities with the tooth finds of Middle Pleistocene hominins from China either in terms of size or shape of the tooth crown , and even the 350,000 to 600,000 year old teeth from the Sima de los Huesos in Spain have more “ modern ” features. The morphology of the tooth find thus supports the relatively large genetic distance, derived from the analysis of the mtDNA, of the Denisova fossils to other similarly old populations of the genus Homo .

In 2015, the discovery of a second molar tooth ( Denisova 8 ) was announced and at the same time its nucleus DNA and its mt-DNA were compared with the respective data of the tooth initially discovered ( Denisova 4 ). The upper tooth Denisova 8 is also quite large, but comes from a somewhat deeper layer than Denisova 4 and is therefore probably older than the tooth that was first discovered; According to the data, Denisova 8 is older than 50,000 years, Denisova 4 is a maximum of 50,000 years old. Both teeth differ significantly from all known Neanderthal finds and may in future serve as a reference for identifying Denisova fossils from other sites.

The analysis of the cell nucleus DNA of both teeth revealed a close genetic proximity to the DNA from the cell nuclei of the finger bone and also confirmed the genetic distance between the fossils and the Neanderthals. The same results were obtained from the analysis of the mt-DNA of both teeth, so that evidence for three individuals of the Denisova people is now considered to be reliable. In addition, the age difference of the teeth proves the existence of the population over a longer period of time.

Toe bone morphology and DNA

Neanderthal toe bones (original)

The distal toe bone described for the first time in 2011 comes from either the 4th or the 5th (little) toe of an adult individual. The bone is remarkably long and has a very strong, very broad shaft; the ratio of great width to comparatively low height is more like the ratio in older Pleistocene than in modern representatives of the genus Homo and exceeds the corresponding dimensions in Neanderthals. Overall, the features of the bone therefore appear ancient, but some features are in the range between the Neanderthals and early modern humans , according to the scientific description of the bone. The greatest similarity exists to the Neanderthal fossil Shanidar-4 and to the Homo sapiens fossil Tianyuan 1 .

Already in 2011 it was pointed out that only an analysis of his genetic material could provide clarity about the phylogenetic classification of the bone. In 2013, Svante Pääbo's research group reported that 60 percent of the bone's DNA corresponds to that of a Neanderthal man. Furthermore, the DNA of the homologous chromosomes is so largely identical that the former owner of the bone was probably the child of the first cousin and cousin. At the same time, it was concluded from the data that 0.5 to 8% of the DNA of Denisovans was introduced into the Denisova genome by a previously unknown population of the genus Homo around 300,000 years ago ; this population split off from the common ancestors of the Neanderthals, the Denisovans, and the anatomically modern humans more than 1 million years ago.

distribution

The distribution area of ​​the Denisova people is unclear due to the few finds known so far. However, the study published in December 2010 mentions that this population may have "lived in large parts of East Asia at the time when Neanderthals were present in Europe and western Asia". On the one hand, this assumption was derived from the finding that there was a gene flow to the ancestors of the Melanesians, which, however, “probably did not take place in southern Siberia”. On the other hand, the higher genetic variability - compared with the Neanderthals - speaks for a relatively large distribution area.

Searching for clues in China

Svante Pääbo's team has been running a laboratory in Beijing since 2008 to search for fossil DNA from Chinese stocks. As a result of this German-Chinese cooperation, it was reported at the beginning of 2013 that - comparable to the Denisova finds - the 40,000-year-old Homo sapiens fossil Tianyuan 1 from near Beijing does not contain a greater proportion of Neanderthal or Denisova DNA than the people living in northern China today. So far (as of spring 2019) no further aDNA could be detected from other finds .

Settlement of East Asia by this population may date back to 300,000 years ago. In July 2011 both Chris Stringer and Milford H. Wolpoff described it as possible that some fossils discovered in China, which so far could not be clearly assigned to Homo erectus or anatomically modern humans, could be ascribed to the Denisova humans; The Dali man and the Jinniushan man were mentioned in this context . In 2012, Chris Stringer further pointed out that in addition to the finds from Dali and Jinniushan, finds from Yunxian and Narmada in India may also be attributed to the Denisova people.

First settlement of Tibet

In 2014, a comparison of Denisova DNA with DNA samples from Tibetans and Han Chinese living today revealed indications of a possible introgression of Denisova DNA into the DNA of the Tibetans and, to a much lesser extent, into the DNA of the Han Chinese. According to the gene analyzes, a variant of the EPAS1 gene , which is said to be identical to a variant otherwise only found in the Denisova people, caused an adaptation of the Tibetans, which made it easier for them to breathe at high altitudes.

In May 2019, the journal Nature announced that a fossil right lower jaw with two well-preserved, very large molars M1 and M2 and several anterior teeth without crowns, which was discovered in the highlands of Tibet in 1980 , can be attributed to the Denisova people. It was emphasized that the Xiahe mandible differs from Homo erectus mandibles, but has similarities with the Xujiayao and Xuchang fossils as well as the Penghu 1 fossil from the Penghu Islands . In this context, the researchers expressed the hope that, due to their similarity to the Xiahe find, further Chinese prehistoric man fossils can be assigned to the Denisova man in the future. Although no DNA samples could be obtained from the fossil, it was possible to analyze proteins from dentin , the structure of which proved to be similar to the evidence from the Denisova cave and was clearly distinguishable from modern proteins. The result was commented on by the Max Planck Society, whose experts had examined the lower jaw together with Chinese researchers, as follows: “Our protein analysis has shown that the Xiahe lower jaw belonged to a population that was closely related to the Denisova people the Denisova Cave. ”The uranium-thorium dating of the calcareous crusts on the lower jaw showed an age of approximately 160,000 years, which makes the fossil the oldest known evidence of the presence of a representative of the hominini in the highlands of Tibet. At the same time, it is the first hominine fossil whose membership of a certain population was proven solely on the basis of a protein determination. The researchers interpreted the find as evidence that Denisova people colonized the highlands of Tibet in the Middle Pleistocene and successfully adapted to the oxygen deficiency long before the region was colonized by anatomically modern humans.

The lower jaw was discovered in 1980 by a monk in Baishiya Cave at 3,280 meters above sea level in Ganjia , Xiahe ( Gansu Province , People's Republic of China ) and handed over to the 6th Gungthang Rinpoche of Labrang Monastery , who presented it to Lanzhou University in Lanzhou handed over. Researchers at Lanzhou University have been studying the fossil since 2016 in cooperation with the Max Planck Institute for Evolutionary Anthropology . During excavations in the cave, stone tools and animal bones with cut marks were also recovered.

Genetic traces in Oceania

The gene flow to the ancestors of the Melanesians and other populations in Oceania presumably occurred independently of that in East Asia. Based on DNA fragments, it has even been suggested that two lines of the Denisova people, who separated over 300,000 years ago, passed on genetic material to the ancestors of the Papuans. One of the two Denisova lines is so different from the other that it could be a separate line. In addition, the genetic traces suggest that the Denisova people only became extinct about 30,000 years ago.

Genetic traces in Spain

As proved "astonishingly" at the end of 2013, findings from a cave (the Sima de los Huesos ) in the north of Spain : From a reference to the molecular clock estimated at an age of about 400,000 years thighbone (femur XIII) of a Homo heidelbergensis had succeeded , DNA mitochondrial (mtDNA) to attract and sequencing . This mtDNA shows a high degree of similarity with the mtDNA of the Denisova people, from which it was concluded that the population to which the former owner of the bone belonged had common ancestors with the Denisova people 300,000 years earlier. The head of the mtDNA study, Matthias Meyer, therefore suspected that the Spanish population of Homo heidelbergensis had an ancestral population "from which both the Neanderthals and the Denisova humans later emerged". In this context, Chris Stringer pointed out that the fossils referred to by Spanish researchers as Homo antecessor could be possible candidates for this ancestral population.

Taxonomic classification of fossils

The relational ( taxonomic ) classification of the fossils is unclear. The discoverers initially referred to the finds in 2010 as "Denisova hominins " based on where they were found . In an article accompanying this publication of the mtDNA analysis in Nature , the evolutionary biologist Eske Willerslev, director of the Center for Ancient Genetics at the University of Copenhagen , was quoted as saying that he also advised against deriving the discovery of a new biological species from the data obtained. Even after analyzing the cell nucleus DNA in 2012, the researchers expressly refrained from naming them in accordance with the requirements of the international rules for zoological nomenclature and instead chose the term Denisovans and, in German accompanying publications, Denisova-Mensch . The researchers also expressly refused to stipulate the status of the Neanderthal in relation to anatomically modern humans ( species versus subspecies ). Instead, they left it to say that the Denisova people were a "sister group" of the Neanderthals.

In his book Die Neandertaler und Wir, Svante Pääbo reported in 2014 that the name "Homo altaiensis" was initially considered; however, it was rejected because - as in the case of Neanderthals and anatomically modern humans - there are no reliable criteria for delimiting or grouping these species.

See also

literature

Web links

Individual evidence

  1. ↑ The spelling Denisova follows the international designation of the " Denissowa Cave " in professional circles . The lemma follows the relevant German designation by the researchers of the Max Planck Society , see on the trail of human evolution. Max Planck Society of December 19, 2013, accessed on December 29, 2016.
  2. ↑ a b c d e David Reich et al .: Genetic history of an archaic hominin group from Denisova Cave in Siberia. In: Nature. Volume 468, No. 7327, 2010, pp. 1053-1060 doi: 10.1038 / nature09710
  3. ↑ "belonging to a hitherto unknown species"
  4. ↑ Katerina Douka et al .: Age estimates for hominin fossils and the onset of the Upper Palaeolithic at Denisova Cave. In: Nature. Volume 565, 2019, pp. 640-644, doi: 10.1038 / s41586-018-0870-z
  5. ↑ Andreas Ziemons: "Everything points to a new human species" . In: Deutsche Welle , March 25, 2010. Retrieved January 11, 2013. 
  6. ↑ Michael Balter: Researchers Discover New Lineage of Ancient Human (en) . In: news.sciencemag.org , March 24, 2010. Archived from the original on August 20, 2011. Retrieved on December 30, 2010. 
  7. ↑ Robin Dennell: Dating of hominin discoveries at Denisova. In: Nature. Volume 565, 2019, pp. 571-572, doi: 10.1038 / d41586-019-00264-0
  8. ↑ a b c Maria Mednikova: A proximal pedal phalanx of a Paleolithic hominin from denisova cave, Altai. In: Archeology, Ethnology and Anthropology of Eurasia. Volume 39, No. 1, 2011, pp. 129-138, doi: 10.1016 / j.aeae.2011.06.017
  9. ↑ a b Kay Prüfer, Fernando Racimo u. a .: The complete genome sequence of a Neanderthal from the Altai Mountains. In: Nature. Volume 505, No. 7481, 2014, pp. 43-49, doi: 10.1038 / nature12886 .
  10. ↑ a b Susanna Sawyer et al .: Nuclear and mitochondrial DNA sequences from two Denisovan individuals. In: PNAS . Volume 112, No. 51, 2015, pp. 15696–15700, doi: 10.1073 / pnas.1519905112
  11. ^ Zhan-Yang Li et al .: Late Pleistocene archaic human crania from Xuchang, China. In: Science. Volume 355, No. 6328, 2017, pp. 969–972, doi: 10.1126 / science.aal2482
  12. ↑ Ancient skulls may belong to elusive humans called Denisovans. On: sciencemag.org of March 2, 2017
    Des crânes de Denisoviens? On: hominides.com of March 3, 2017
  13. ↑ China: Researchers find a 100,000 year old skull. On: spiegel.de from January 23, 2008
  14. ^ Zenobia Jacob et al .: Timing of archaic hominin occupation of Denisova Cave in southern Siberia. In: Nature. Volume 565, 2019, pp. 594-599, doi: 10.1038 / s41586-018-0843-2
  15. ↑ Katerina Douka et al .: Age estimates for hominin fossils and the onset of the Upper Palaeolithic at Denisova Cave. In: Nature. Volume 565, 2019, pp. 640-644, doi: 10.1038 / s41586-018-0870-z
  16. ↑ Robin Dennell: Dating of hominin discoveries at Denisova. In: Nature. Volume 565, 2019, pp. 571-572, doi: 10.1038 / d41586-019-00264-0
  17. ↑ a b Fahu Chen, Frido Welker, Chuan-Chou Shen, Shara E. Bailey, Inga Bergmann, Simon Davis, Huan Xia, Hui Wang, Roman Fischer, Sarah E. Freidline, Tsai-Luen Yu, Matthew M. Skinner, Stefanie Stelzer, Guangrong Dong, Qiaomei Fu, Guanghui Dong, Jian Wang, Dongju Zhang, Jean-Jacques Hublin : A late Middle Pleistocene Denisovan mandible from the Tibetan Plateau. In: Nature. Online pre-publication from May 1, 2019, doi: 10.1038 / s41586-019-1139-x
  18. ^ A b Johannes Krause et al .: The complete mitochondrial DNA genome of an unknown hominin from southern Siberia. In: Nature . Volume 464, No. 7290, 2010, pp. 894-897. doi: 10.1038 / nature08976 full text (PDF; 298 kB)
  19. ↑ J. Krause et al .: A complete mtDNA genome of an early modern human from Kostenki, Russia. In: Current Biology . Volume 20, No. 3, 2010, pp. 231-236, doi: 10.1016 / j.cub.2009.11.068
  20. ^ E. Andrew Bennett, Isabelle Crevecoeur, Bence Viola et al .: Morphology of the Denisovan phalanx closer to modern humans than to Neanderthals. In: Science Advances. Volume 5, No. 9, 2019, eaaw3950, doi: 10.1126 / sciadv.aaw3950
  21. ↑ Sonja Kastilan : Sensational find “X-Woman”: Did researchers discover a new species of human being? . In: faz.net , March 25, 2010. Retrieved January 11, 2013.  
  22. ↑ Ulrich Bahnsen: The Alien of Altai: The little finger of evolution . In: Zeit Online , March 24, 2010. Retrieved January 11, 2013. 
  23. ↑ Index of denisova , see: Max Planck Society: A High Coverage Denisovan Genome.
  24. ^ Max Planck Institute for Evolutionary Anthropology , accessed January 11, 2013: A High Coverage Denisovan Genome.
  25. ↑ idw-online of February 7, 2012: Entire genome of an extinct human form decoded from a fossil.
  26. ↑ See also the specialist publication: Matthias Meyer et al .: A High-Coverage Genome Sequence from an Archaic Denisovan Individual. In: Science . Volume 338, No. 6104, 2012, pp. 222-226, doi: 10.1126 / science.1224344
  27. ↑ cf. the most recent revision of the data on mutation rates in great apes in: Kevin E. Langergraber et al .: Generation times in wild chimpanzees and gorillas suggest earlier divergence times in great ape and human evolution. In: PNAS . Volume 109, No. 39, 2012, pp. 15716-15721, doi: 10.1073 / pnas.1211740109 ; For examples of the multitude of competing data, see also Human Tribal History # Molecular biological and paleoanthropological findings on the origin of the human species
  28. ↑ In the original wording : “ We call the group to which this individual belonged Denisovans in analogy to Neanderthals, as Denisovans are described for the first time based on molecular data from Denisova Cave just as Neanderthals were first described based on skeletal remains retrieved in the Neander Valley in Germany. ”
    “ We call the group to which this individual belonged, Denisovans, in analogy to the Neanderthals, because just as the Neanderthals were first described using skeletal finds from the Neanderthal in Germany, the Denisovans were first described using molecular data from the Denisova -Cave."
  29. ↑ Viviane Slon, Fabrizio Mafessoni, Benjamin Vernot et al .: The genome of the offspring of a Neandertal mother and a Denisovan father. In: Nature. Volume 561, 2018, pp. 113–116, doi: 10.1038 / s41586-018-0455-x
  30. ↑ Samantha Brown, Thomas Higham , Viviane Slon, Svante Pääbo et al .: Identification of a new hominin bone from Denisova Cave, Siberia using collagen fingerprinting and mitochondrial DNA analysis. In: Scientific Reports. Volume 6, Article No. 23559, 2016, doi: 10.1038 / srep23559
  31. ↑ Mother Neanderthals, father Denisovans. On: mpg.de from August 22, 2018
  32. ↑ Mayukh Mondal, Jaume Bertranpetit and Oscar Lao: Approximate Bayesian computation with deep learning supports a third archaic introgression in Asia and Oceania. In: Nature Communications. Volume 10, 2019, Article No. 246, doi: 10.1038 / s41467-018-08089-7
  33. ↑ Richard E. Green et al .: A draft sequence of the Neandertal Genome. In: Science . Volume 328, No. 5979, 2010, pp. 710-722. doi: 10.1126 / science.1188021 PDF
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