Denisovan

The Denisovans or Denisova hominins are an extinct species or subspecies of archaic human that ranged across Asia during the Middle to Late Pleistocene, approximately 200,000–32,000 years ago. Most of what is known about Denisovans comes from DNA evidence. While many recent fossils have been found and tentatively identified as Denisovan, the first Denisovans discovered were known from few physical remains. Consequently, no formal species name has been established. However, an analysis of the mitochondrial DNA and endogenous proteins from the Harbin cranium, which had been given the name Homo longi, showed with great certainty that this species represents a Denisovan.
In a study published in September 2025, remains from six additional sites in China including the 1 million year old Yunxian man were proposed to be included in the species Homo longi along with the genetically confirmed Denisovans.
The first identification of a Denisovan individual occurred in 2010, based on mitochondrial DNA extracted from a juvenile finger bone excavated from the Siberian Denisova Cave in the Altai Mountains in 2008. Nuclear DNA indicates close affinities with Neanderthals. The cave was also periodically inhabited by Neanderthals. Additional specimens from Denisova Cave were subsequently identified, as were specimens from the Baishiya Karst Cave on the Tibetan Plateau, Tam Ngu Hao 2 Cave in the Annamite Mountains of Laos, the Penghu channel between Taiwan and the mainland, and Harbin in Manchuria.
DNA evidence suggests they had dark skin, eyes, and hair, and had a Neanderthal-like build. Based on the Harbin cranium, like other archaic humans, the skull is low and long, with massively developed brow ridges, wide eye sockets, and a large mouth. The two existing Denisovan mandibles show that like Neanderthals, the Denisovans lacked a chin. Like modern humans and the much earlier Homo antecessor, but unlike Neanderthals, the face is rather flat, but with a larger nose. However, they had larger molars which are reminiscent of Middle to Late Pleistocene archaic humans and australopithecines. The cranial capacity and therefore the brain size of the Denisovans was within the range of modern humans and Neanderthals.
Denisovans interbred with modern humans, with a high percentage of Denisovan DNA occurring in Melanesians, Aboriginal Australians, and Filipino Negritos. In contrast, 0.2% derives from Denisovan ancestry in mainland Asians and Native Americans. In a 2018 study, South Asians were found to have levels of Denisovan admixture similar to that seen in East Asians. Another study found that the highest Denisovan ancestry is inferred in Oceanians, while most populations of Native Americans, East Asians, and South Asians have similar amounts. This distribution suggests that there were Denisovan populations across Asia. There is also evidence of interbreeding with the Altai Neanderthal population, with about 17% of the Denisovan genome from Denisova Cave deriving from them. A first-generation hybrid nicknamed "Denny" was discovered with a Denisovan father and a Neanderthal mother. Additionally, 4% of the Denisovan genome comes from an unknown archaic human species, which diverged from modern humans over one million years ago.

Taxonomy

Denisovans might represent a new species of Homo or an archaic subspecies of Homo sapiens, but up until the Harbin cranium was identified as a Denisovan in June 2025 through the mitochondrial DNA and autosomal proteomics there were too few fossils to erect a proper taxon. Proactively proposed species names without type specimen designation for Denisovans include H. sapiens altaiensis or H. altaiensis, H. denisoviensis, H. denisovan, and H. denisova.
In 2025, Denné Reed argued that the informal name "Denisovans" represents the better system than proactively proposed names to reference this archaic human group due to its uncertain biological status as an independent evolutionary lineage. He suggested that "H. altaiensis" represents a nomen nudum, since its description lacks differential diagnosis, does not clearly display the intent of naming a new species and lacks a fixed type specimen. He also suggested that the names "Homo daliensis" and "Homo mapaensis" are conditionally proposed which makes them unavailable based on ICZN article 15, while considered "Homo tsaichangensis" to be unpublished and unavailable, as it does not contain evidence of ZooBank registration within the published work which fails to conform to the ICZN articles 8.5.3.1 and 8.5.3.2.
Research published in 2024 proposed classifying Denisovans as part of the conditional species Homo juluensis based on the similarities between Denisovan and H. juluensis molars, prior to the classification of Denisovans as Homo longi based on DNA evidence.
Some older findings called "East Asian Archaics" have been associated in studies with the Denisovans but may or may not belong to the Denisovan line. Such findings include the Dali skull, the Xujiayao hominin, the Xuchang crania, the Jinniushan human, the Hualongdong people, Yunxian Man, Maba Man, and the Narmada Human.
In 2021, Chinese palaeoanthropologist Qiang Ji and colleagues suggested that their newly erected species, H. longi, may represent the Denisovans, based on the similarity between the type specimen's molar and that of the Xiahe mandible. In 2024, paleoanthropologists Christopher Bae and Xiujie Wu designated the Xujiayao fossils as the holotype of the species Homo juluensis with Xuchang as the paratype, and suggested sinking Denisovans into this species. They recommended relegating the Dali Man and the similar specimen Jinniushan to H. longi.
In 2025, Fu and colleagues retrieved mitochondrial DNA from the dental calculus of the Harbin cranium, reporting that it falls within the variation of seven previously sequenced Denisovan mitochondrial DNA. Fu et al. also retrieved 95 endogenous proteins from the same specimen, and suggested that H. longi can be confidently assigned to a Denisovan population.
In 2025, Feng and colleagues, the team that published the Harbin cranium in 2021 and named the new species Homo longi, ran a morphometric analysis of 104 ancient hominin cranial and mandibular specimens using 533 morphometric landmarks and grouped Yunxian Man, Dali Man, the Hualongdong people, the Jinniushan human, the Xujiayao hominins, and Maba Man under H. longi alongside the genetically confirmed Denisovans.

Discovery

The Denisova Cave is located in Altai Krai, Russia, in south-central Siberia, on the western edges of the Altai Mountains. It is named after Denis, a Russian Old Believer hermit who lived there in the 18th century. The cave was first inspected for fossils in the 1970s by Soviet paleontologist Nikolai Ovodov, who was looking for remains of canids.
In 2008, Michael Shunkov from the Russian Academy of Sciences and other Russian archaeologists from the Institute of Archaeology and Ethnography of the Siberian Branch of the Russian Academy of Sciences in Novosibirsk Akademgorodok investigated the cave and found the finger bone of a juvenile female hominin originally dated to 50–30,000 years ago. The estimate has changed to 76,200–51,600 years ago. The specimen was originally named X-woman because matrilineal mitochondrial DNA extracted from the bone demonstrated it to belong to a novel ancient hominin, genetically distinct both from contemporary modern humans and from Neanderthals.
In 2019, Greek archaeologist Katerina Douka and colleagues radiocarbon dated specimens from Denisova Cave, and estimated that Denisova 2 lived 195,000–122,700 years ago. Older Denisovan DNA collected from sediments in the East Chamber dates to 217,000 years ago. Based on artifacts also discovered in the cave, hominin occupation began 287±41 or 203±14 ka. Neanderthals were also present 193±12 ka and 97±11 ka, possibly concurrently with Denisovans.

Specimens

The fossils of multiple distinct Denisovan individuals from Denisova Cave have been identified through their ancient DNA : Denisova 2, 3, 4, 8, 11, 19, 20, 21 and 25. An mtDNA-based phylogenetic analysis of these individuals suggested that Denisova 19, 20, and 21 are the oldest, followed by Denisova 2, then Denisova 8; while Denisova 3 and Denisova 4 were roughly contemporaneous. The mtDNA from Denisova 4 bore a high similarity to that of Denisova 3, indicating that they belonged to the same population.
Denisova Cave contained the only known examples of Denisovans until 2019, when a research group led by Fahu Chen, Dongju Zhang, and Jean-Jacques Hublin described a partial mandible discovered in 1980 by a Buddhist monk in the Baishiya Karst Cave on the Tibetan Plateau in China. Known as the Xiahe mandible, the fossil became part of the collection of Lanzhou University, where it remained unstudied until 2010. It was determined by ancient protein analysis to contain collagen that by sequence was found to have close affiliation to that of the Denisovans from Denisova Cave, while uranium decay dating of the carbonate crust enshrouding the specimen indicated it was more than 160,000 years old. The identity of this population was later confirmed through study of environmental DNA, which found Denisovan mtDNA in sediment layers ranging in date from 100,000 to 60,000 years before present, and perhaps more recent. A 2024 reanalysis identified a partial Denisovan rib fragment dating to between 48,000 and 32,000 BP.
In 2018, a team of Laotian, French, and American anthropologists, who had been excavating caves in the Laotian Annamite Mountains since 2008, was directed by local children to the site Tam Ngu Hao 2 where they recovered a human tooth. The tooth developmentally matches a 3.5 to 8.5 year old, and a lack of amelogenin suggests it belonged to a girl, barring extreme degradation of the protein over a long period of time. Dental proteome analysis was inconclusive for this specimen, but the team found it anatomically comparable with the Xiahe mandible, and so they categorized it as a Denisovan. The tooth probably dates to 164,000 to 131,000 years ago.
In 2022, a team from Germany, Austria, Russia and the UK found three Denisovans from layer 15 of the East Chamber, in Denisova Cave. It turns out that the mtDNA sequences of Denisova 19 and 21 are identical, indicating that they may belong to the same individual or be maternal relatives. The divergence date for the mtDNAs of the three new and the four previously published Denisovans is 229 ka during the Interglacial period MIS 7. The genetic diversity among the Denisovans from Denisova Cave is on the lower range of what is seen in modern humans, and is comparable to that of Neanderthals. However, it is possible that the inhabitants of Denisova Cave were more or less reproductively isolated from other Denisovans, and that, across their entire range, Denisovan genetic diversity may have been much higher.
In 2024, scientists announced the sequence of Denisova 25, which was in a layer dated to 200 ka. During DNA sequencing, low proportions of the Denisova 2, Denisova 4 and Denisova 8 genomes were found to have survived, but high proportions of the Denisova 3 and Denisova 25 genomes were intact. The Denisova 3 sample was cut into two, and the initial DNA sequencing of one fragment was later independently confirmed by sequencing the mtDNA from the second.
In 2008, a Taiwanese citizen purchased a fossil Homo mandible, dredged from the sea floor of the Taiwan Strait, from an antique shop and donated it to the Taiwan National Museum of Natural Science. Attempts to extract DNA were unsuccessful, but in 2025 protein analysis of the specimen, designated Penghu 1, was published showing that it belonged to a male Denisovan.
In 2018, a relatively complete skull was reported from Harbin, China, and was described in 2021 as H. longi. In 2025, mtDNA and proteomic analysis confirmed that this skull is a Denisovan.
The following list is the currently known specimens of Denisovans, with colored specimens being proposed through morphometric analyses only without mitogenome or proteomic analyses:

Name	Fossil elements	Age	Discovery	Place	Sex and age	Publication	Image	GenBank / Genebase accession
Denisova 3	Distal phalanx of the fifth finger	76.2–51.6 ka	2008	Denisova cave	13.5-year-old adolescent female	2010
Denisova 4	Permanent upper 2nd or 3rd molar	84.1–55.2 ka	2000	Denisova cave	Adult male	2010
Denisova 8	Permanent upper 3rd molar	136.4–105.6 ka	2010	Denisova cave	Adult male	2015
Denisova 2	Deciduous 2nd lower molar	194.4–122.7 ka	1984	Denisova cave	Adolescent female	2017
Xiahe mandible	Partial mandible	> 160 ka	1980	Baishiya Cave		2019
Penghu 1	Partial mandible	130 to 190 kya or 10 to 70 kya	2008	Penghu Channel	Adult male	2015
Denisova 11	Arm or leg bone fragment	118.1–79.3 ka	2012	Denisova cave	13 year old adolescent female	2016
Denisova 13	Parietal bone fragment	Layer 17, 202-167 ka	2019	Denisova cave		pending
TNH2–1	Permanent lower left 1st or 2nd molar	164–131 ka	2018	Tam Ngu Hao 2 cave	3.5 to 8.5 year old female	2022
Denisova 19	Undiagnostic bone fragment	Layer 15, 217–187 ka	2012–13	Denisova cave		2022		-
Denisova 20	Undiagnostic bone fragment	Layer 15, 217–187 ka	2012–13	Denisova cave		2022		-
Denisova 21	Undiagnostic bone fragment	Layer 15, 217–187 ka	2012–13	Denisova cave		2022		-
BSY-19-B896-1	Distal rib fragment	48-32 ka	1980	Baishiya Cave	Unknown	2024		-
Denisova 25	Molar	200 ka	2024	Denisova cave	Male	pending		-
Harbin cranium	Complete skull	>146 ka	1933?	Harbin	Adult male aged approximately 50 years	2021
Dali cranium	Complete skull	260±20 ka	1978	Dali County	Adult	1981
Jinniushan human	Fragments of the skull cap, ribs, hand, pelvis, and leg bones	260±20 ka	1984	Jinniushan	Adult female	1985
Xujiayao hominin	12 parietal bones, 1 temporal bone, 2 occipital bones, 1 mandibular bone fragment, 1 juvenile maxilla, and 3 isolated teeth.	130–71 ka	1976–1979	Xujiayao village in Yanggao County	Adult and juvenile	2011
HLD 6	11 fossil parts belong to a single individual	300 ka	2014–2016, 2020	Hualong Cave	Juvenile 12–13 years old	2019
Maba 1	Partial skull, a skull cap and parts of the right upper face, with parts of the nose also still attached	300–130 ka	1958	Lion Cave, Maba, near Shaoguan city in the northern part of Guangdong province	Adult middle-aged male	1959