Денисова пещера | |
 Denisova Cave in 2008 | |
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 Interactive map of Denisova Cave | |
| Alternative name | Аю-Таш |
|---|---|
| Location | Soloneshensky District,Altai Krai |
| Region | Siberian Federal District, Russia |
| Coordinates | 51°23′51″N84°40′34″E / 51.39750°N 84.67611°E /51.39750; 84.67611 |
| Altitude | 700 m (2,297 ft)[1] |
| Type | limestone,karst |
| Area | 270 m2 (2,900 sq ft) |
| History | |
| Periods | Paleolithic 30,000 to 48,000 years ago |
| Cultures | Denisovans,Neanderthals,Homo sapiens sapiens |
Denisova Cave (Russian:Денисова пещера,romanized: Denísova peshchéra,lit. 'the cave of Denis') is acave in theBashelaksky Range of theAltai Mountains in Siberia, Russia.
It is widely known for having provided items of greatpaleoarchaeological andpaleontological interest. In particular, the 2008 discovery of bone fragments that in 2010 have been conclusively established to have belonged to a separate early human species - theDenisova hominin - which is named after the cave. Other items including artifacts dated to around 40,000BP. Remains of a 32,000-year-old prehistoric species ofhorse have also been found in the cave.
The cave is located in a region thought to have been inhabited concurrently in the past byNeanderthals and by modern humans. Abone needle dated to 50,000 years ago was discovered at the archaeological site in 2016 and has been described as the most ancient needle known[2][3][4] (though another possible needle dates to about 10,000 years earlier from South Africa fromc. 61,000 years ago[5]).
Denisovans,Neanderthals andrelated hybrids may have inhabited the Denisova Cave for extended periods, but perhaps not at the same time.[6] The attribution of the needle and certain other artifacts at the cave, whether toHomo sapiens or to the Denisova hominin is uncertain.[7]
The "mystery population" ofHomo colloquially referred to as "Denisovans" was for a time also sometimes referred to by the scientific nameHomo denisova after the cave, until June 2025, whenDenisovans were confirmed to beHomo longi, makingH. denisova a junior synonym.[8][9]
Located inAltai Krai, near the border withAltai Republic, both inRussia, the cave is near the village of Chorny Anui (Чёрный Ануй), and some 150 km (93 mi) south ofBarnaul, the regional capital. The cave, which is approximately 28 m (92 ft) above the right bank of theAnuy River (a left tributary of theOb), has formed in upperSilurian limestone and contains a floor area of about 270 m2 (2,900 sq ft). The cave is composed of three galleries. The central chamber, the Main Gallery, contains a floor of 9 m × 11 m (30 ft × 36 ft) with side galleries, the East Gallery and the South Gallery.[10][11] It has been described both as akarst cave[12] and as a sandstone cave.[11]
Cave sediments are rich with remnants of animals, including extinct ones. Remains of 27 species of large and medium-sized mammals have been found, (such ascave hyena,cave lion, etc.) and 39 species of small mammals, as well as remnants of reptiles, 50 bird species and other vertebrates.[13][14] Pollen in the cave sediments is used forpalaeoclimatological research.
In the 18th century, the cave was inhabited by anOld Believer hermit, Dyonisiy (Denis), and was named after him.[12]
In the 1970s, Russian scientists discovered paleoarcheological remains in the cave that led to further explorations.[12] So far, 22strata have been identified, with archeological artifacts that cover the time from Dyonisiy back to about 125,000–180,000 years ago.[11] The dating of the strata was accomplished by the use ofthermoluminescence dating ofsediments, or, in some cases,radiocarbon dating oncharcoal.[11]
Among the archeological artifacts areMousterian- andLevallois-style tools attributed toNeanderthals.[15] Beside tools, researchers found decorative objects of bone, mammoth tusk, animal teeth, ostrich egg shell, fragments of a stone bracelet made of drilled, worked, and polished dark greenchlorastrolite, and pendants.[11] A seven-centimetre (2.8 in)sewing needle made from bird bone, estimated to be around 50,000 years-old, was found in Denisova Cave.[2] The cave also contains stone tools and bone artifacts made by modern humans, and Pääbo commented: "The one place where we are sure all three human forms have lived at one time or another is here in Denisova Cave."[16]
Denisovans,Neanderthals andrelated hybrids, may have inhabited the Denisova Cave over thousands of years, but it is unclear whether they ever shared the cave.[6]
In 2019, a team of archaeologists from the Russian Academy of Sciences' Institute of Archaeology and Ethnography in Novosibirsk discovered a 45,000-year-oldcave lion statue made from a woolly mammoth tusk, according toThe Siberian Times. This 42 mm long, 8 mm thick and 11 mm high figurine was unearthed in the eleventh layer of the southern gallery of Denisova Cave. According to Siberian archeologists, this statue made byUpper Palaeolithic artist might be the oldest animal figurine in the world. The lion's hind legs, groin, back and belly are covered with eighteen rows of notch ornaments and its head is missing. On the right side of the lion there are two extra rows with four notches.[17][18]
The average annual temperature of the cave remains at 0 °C (32 °F), which has contributed to the preservation of archaic DNA among the remains discovered.[19]
Scientists from the Russian Academy of Sciences' Institute of Archaeology and Ethnography inNovosibirsk have investigated the cave. Among the artifacts which had been left about 30,000 to 48,000 years ago (strata 9–11), bones were identified. One of these bones was a piece ofphalanx of a child found in layer 11.2 of the East Gallery. The fossil element was analyzed bySvante Pääbo and coworkers from theMax Planck Institute for Evolutionary Anthropology inLeipzig; itsmitochondrial DNA revealed a structure that differs from known human patterns and has been ascribed to "Denisova hominin".[15] Pääbo and his co-workers first intended to classify the Denisovans as a separate species but changed their minds prior to publication of the results.[20][21] Further analysis revealed the Denisovans were related to the Neanderthals and interbred with the ancestors of modernMelanesians.[22]
In 2010, a toe bone was discovered in the cave, in layer 11.4 of the East Gallery, and therefore contemporary with the Denisovan finger bone. Preliminary characterization of the bone's mitochondrial DNA suggested it belonged to a Neanderthal, not a Denisovan.[16] Later analysis confirmed the toe bone as coming from a Neanderthal.[23] The firsthigh-coverage genome of Neanderthals was taken from this toe bone.[23]
This Neanderthal is referred to as theAltai Neanderthal. TheAltai Neanderthal is estimated to be around 120,000 years old. Other Neanderthals for whichnuclear DNA has been recovered are all genetically closer to each other than to theAltai Neanderthal. Modern humans andUst'-Ishim man share more alleles with all other Neanderthals than with theAltai Neanderthal, which shows that theintrogression event from Neanderthals into humans likely took place after the split of the lineage of theAltai Neanderthal from that of other Neanderthals.[24]
The use of collagenpeptide mass fingerprinting of ancient bone samples, calledZooarchaeology by Mass Spectrometry (ZooMS), for species identification allowed the sorting of 2,315 unidentified bone fragments from 2012[25] or 2014 excavation.[26][27] One sample, DC1227 (Denisova 11), taken from layer 12 of the East Gallery, carried human traits. This was the first time that this technique was used to successfully identify the presence of an extinct hominid.[27] DC1227 was a bone fragment weighing 1.68 g (0.059 oz), measuring in with a maximum length of 24.7 mm (0.97 in) and maximum width of 8.39 mm (0.330 in).[26] Further analysis showed the bone fragment to have Neanderthalmitochondrial DNA (mtDNA).[26][27]
Whole genome sequencing and other characterization ofDenisova 11 to 2.6-foldcoverage showed this specimen belonged to a female at least thirteen years old at the time of death.Denisova 11 was found to be the hybrid progeny of a Neanderthal mother and a Denisovan father.[25] Comparing the entire genome against all archaic hominin genomes on record,Denisova 11 shares the most genetic affinity withDenisova 3, indicating that her Denisovan father is more closely related toDenisova 3 than her mother was to any of the sequenced Neanderthals. LikeDenisova 3, this father carries someintrogressed Neanderthal DNA from an admixture event far in the past, estimated at more than 300 generations earlier, and similar in sequence to the Altai Neanderthal genome. However, the mother ofDenisova 11 was genetically closer to Neanderthal specimenVindija 33.19 fromVindija Cave in Croatia and to other sequenced Neanderthal individuals than to theAltai Neanderthal. This suggests a migration or population turnover involving the Neanderthal populations of the region surrounding the Denisova cave.[25][28]
Sequencing of DNA from soil samples taken from Denisova Cave showed the presence of Neanderthal and Denisovan mtDNA from several samples, as well as the DNA of several animals. Neanderthal mtDNA was present in soil samples from layer 15 of the Main Gallery, a layer associated with Paleolithic artifacts where no Neanderthal fossils have been found. Neanderthal and Denisovan mtDNA were present in samples from layers 14 and 15, respectively, from the East Gallery, lower than any previous fossil finds.[10]
MtDNA has also been recovered from anequine fossil, dating to around 32,000 years ago, taken from Denisova Cave. The equid fossil was identified as coming fromEquus ovodovi an extinct species first described based on a 40,000 year old fossil taken from Proskuryakova Cave inKhakassia, Russia. The mtDNA of the Denisova sample shows close affinity for that taken from Proskuryakova Cave. DNA analysis placesEquus ovodovi as aphylogenetically basal group fornon-caballine horses, with closer genetic affinities withzebras andasses.[29]
A deer tooth pendant impregnated with the genetic material of anAncient North Eurasian (ANE) woman was found in the Denisova Cave, and dated to circa 24,700 years before present. The woman was found to be closely related toMal'ta andAfontova Gora specimens found further east.[30]
So far, the fossils of seven distinct individuals from Denisova Cave have been identified through their DNA. Four of the individuals,Denisova 2,Denisova 3,Denisova 4,Denisova 8, andDenisova 25 are classified as Denisovans.[31] Denisova 2 and Denisova 3 are young females, while Denisova 4 and Denisova 8 are adult males.[31][32] mtDNA analysis of the Denisovan individuals suggests that Denisova 2 is the oldest, followed by Denisova 8, while Denisova 3 and Denisova 4 are roughly contemporaneous.[31] Denisova 25 is estimated to be from 200ka.[33]
One of the individuals, theAltai Neanderthal, is a Neanderthal woman.[23] Before its DNA was sequenced, the Altai Neanderthal had been given the provisional name ofDenisova 5.[32] In 2018,Denisova 11 was identified as a Neanderthal/Denisova hybrid, based onwhole genome sequencing and comparisons.[28]
During DNA sequencing, Denisova 2, Denisova 4 and Denisova 8 yieldedlow-coverage genomes, while Denisova 3, Denisova 25, and the Altai Neanderthal yielded high-coverage genomes.[32][31]
| Name | Element | Layer | Location | Type | Age | Sex | Discovery | First public. | Image | Notes |
|---|---|---|---|---|---|---|---|---|---|---|
| Denisova 2[31] | deciduous lower molar | 22.1 | Main Gallery | Denisovan | >100 ka[31] | Female | 1984 | |||
| Denisova 3 orX Woman[34][35] | finger phalanx | 11.2 | East Gallery | Denisovan | 30–50 ka | Female | 2008 Team ofMichael Shunkov | Johannes Krause,et al.[36] |  | Cut in two, one piece partially destroyed to investigate the mtDNA. |
| Denisova 4[34][37][38][39] | upper molar | 11.1 | South Gallery | Denisovan | 30–50 ka | Male | 2000 |  | ||
| Altai Neanderthal orDenisova 5[23] | proximal toe phalanx | 11.4 | East Gallery | Neanderthal | 30–50 ka (disputed)[40] | Female | 2010 | M.B. Mednikova (2011)[41] | [42] | Molecular clock analysis of mtDNA suggested this specimen instead dates to ~120 ka.[40] |
| Denisova 8[32] | upper molar | 11.4–12 | East Gallery | Denisovan | Male | |||||
| Denisova 11[26][25][28] | long bone fragment | 12 | East Gallery | Neanderthal/Denisovan hybrid | ~90 ka | Female | 2014 | Samantha Brown,et al. (2016) | ||
| Denisova 25[33] | molar | Denisovan | ~200 ka | Male | 2024 | 2024 |
The cave also preserves remains ofwoolly mammoth,woolly rhinoceros,wild horse,Equus ovodovi,Irish elk,Siberian roe deer,red deer,moose,reindeer,wild yak,steppe wisent,snow leopard,Eurasian cave lion,Eurasian lynx,manul,cave hyena,steppe fox,red fox,grey wolf,dhole,brown bear,Pleistocene small cave bear,wolverine,kolonok,least weasel,pale weasel,steppe polecat,stoat,sable,Eurasian beaver, andAltai marmot.[43][44]
Application of the method to an Upper Palaeolithic deer tooth pendant from Denisova Cave, Russia, resulted in the recovery of ancient human and deer mitochondrial genomes, which allowed us to estimate the age of the pendant at approximately 19,000–25,000 years. Nuclear DNA analysis identifies the presumed maker or wearer of the pendant as a female individual with strong genetic affinities to a group of Ancient North Eurasian individuals who lived around the same time but were previously found only further east in Siberia. Our work redefines how cultural and genetic records can be linked in prehistoric archaeology. (...) Nuclear DNA analysis identifies the presumed maker or wearer of the pendant as a female individual with strong genetic affinities to a group of Ancient North Eurasian individuals who lived around the same time but were previously found only further east in Siberia.
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