Denisovan

## *H. daliensis*

Credit: Chuang Zhao.

The Denisovan or Denisova hominin (*Homo daliensis*) is an extinct species of hominin that lived in Eurasia during the Pleistocene.

## History

The first Denisovan remains were found in Denisova Cave, south-central Russia, where it's common name was coined. The term "Denisova" is named after Denis (Dyonisiy), a Russian hermit who inhabited the cave during the 1700s. Nikolai Ovodov found the first Denisovan remains when inspected for canid remains in the 1970s. Michael Shunkov (2008) investigated the cave, finding a finger bone of a 50-30,000-year-old young woman. Since, however, she has been re-dated to 76,200-51,600 years old. She was first named the X Woman since mtDNA extracted from her finger revealed she was a new species distinct from apomorphic humans and Neanderthals. Katerina Douka et al. (2019) radiocarbon dated the cave, estimating the oldest specimen, Denisova 2, lived 195,000-122,700 years ago. Older DNA from the East Chamber dates to 217,000 years old, with artifacts from the cave suggest hominin occupation began 287±41 or 203±14 ka. Neanderthals were present 193±12 ka and 97±11 ka, possibly contemporaneous with Denisovans. The Australian site of Madjedbebe has been previously estimated to 65,000 years, but is definitely older than 50 kya, despite being of complicated age. The site preserves wood, edible plants, >10,000 artifacts and cave paintings. This is older than the arrival of modern humans.

## Description

The precise anatomical features of the Denisovan are relatively unknown since little material has been identified. The finger bone sits in the variation of a apomorphic woman, contrasted by the large, robust molars similar to Middle-Late Pleistocene plesiomorphic humans. The third molar is outside the range of all *Homo* except for *Homo habilis* and *Homo rudolfensis*, more similar to australopithecines. The second molar is larger than contemporaneous lineages, more similar to *Homo erectus* and *Homo habilis*. Like Neanderthals, a gap exists behind the molars and flattened front teeth, with Denisovans lacking a high mandibular body and a receded mandibular symphasis. The parietal is similar to *Homo erectus*. Facial reconstructions found by comparing methylation at individual genetic loci suggest they have a long, broad, projecting face with a large nose, sloping forehead, protruding jaw, elongate and flatter skull and a wide chest and hips, like Neanderthals. However, they have a longer tooth row than both contemporaneous lineages. Middle/Late Pleistocene East Asian plesiomorphic human skullcaps are similar to Neanderthals. Xuchang skullcaps have prominent brow ridges, with the nuchal and angular tori near to the cranial base are reduced or absent, with the back of the skull being rounded. Xuchang 1 has a very large cranial capacity at ~1800 cc, being the high end for Neanderthals and humans but far above apomorphic humans. Denisova Cave Denisovans bear genetic variations associated with dark skin, brown hair and brown eyes in apomorphic humans. Their genome also contains a variant region of EPAS1 where most in this region would bear WARS2 or TBX15; the former allows for low-oxygen adaptation and the former two affect body fat distribution. Denisovan alleles have the highest frequency of bone and tissue genes, unlike the frequency of genes in the brain seen in introgressed Neanderthal alleles found in Papuans. The 5th phalanx bears morphology distinct from *Neanderthals* and closer to modern humans.

The Harbin skull have a low and long skull, a high and wide upper face, a large nasal opening that holds a large nose (a possible adaptation for cold air), large and square orbits, thick brow ridges that are inflated, cheekbones that are low and flat, a wide palate and large tooth sockets (which formed a large mouth) and a broad skull base. The Harbin cranium is also the largest plesiomorphic human skull in naso-occipital length and supraorbital torus length. Additionally, it has a massive brain, having ~1420 cranial capacity, which is above the range of all species of *Homo* except for *Homo sapiens* and *Homo neanderthalensis*. The post-orbital constriction is developed more than Neanderthals, but not as much as Homo heidelbergensis. The skull is 221.3×164.1 millimeters (8.71x6.46 inches) in length, has a naso-occipital length of 212.9 millimeters (8.38 inches) and has a brow ridge 145.7 millimeters (5.74 inches) long. The braincase retains many plesiomorphic features, but the occipital lacks notable protuberances, tori or massive flexion, which are all present in *Homo erectus* and *Homo heidelbergensis*, and the later *Homo neanderthalensis*. Unlike humans or Neanderthals, the parietals do not expand or protrude significantly. The face was very wide, but also exhibited reduced mid-facial prognathism like apomorphic humans, *Homo antecessor* and various Middle Pleistocene Chinese individuals. The Harbin cranium bears a mosaic of plesiomorphic and derived characteristics, which is very similar to the oldest Homo sapiens from Africa (Jebel Irhoud, Eliye Springs...). Since the Harbin skull is thought to be closely affiliated with the Xiahe mandible, the describers believe Denisovans lacked a chin. However, the specimen lacked the mandible when it was discovered. Unlike the Dali and Hualong skulls, no sagittal keel stretches across the midline. It exhibits alveolar prognathism. Additionally, the only preserved tooth is the upper left second molar, which measures a massive mesiodistal x buccolingual of 13.6x16.6 millimeters (0.54x0.65 inches), which is compared to Denisovan teeth from Denisova. It is oval-shaped, extremely worn and near-flat. It is believed the Harbin individual is a man less than 50 years of age, based on the robustness and size of the material, suture closures and tooth wearing. They speculated this individual had medium-dark skin, hair and eyes based on known genetic evidence from *Homo neanderthalensis*, other Denisovans and early apomorphic *Homo sapiens*.

## Classification

It is unsure if Denisovans are an plesiomorphic subspecies of or a distinct species of human. DNA analysis classifies them as sister to Neanderthals. However, the exact taxonomic placement of them is also disputed. "H. denisova" and "H. altaiensis" are proposed epithets for Denisovans; they are apparently set to be named *H. daliensis* and "*H. longi*" will be lumped as it takes taxonomic priority.

 File:Dragon man by clumsystiggy demc6ej-fullview.jpg
The Harbin individual lived during the same time as Neanderthals and *H. sapiens*. Some researchers think this species is a transition between *Homo erectus* and *H. sapiens*, and others think The Harbin cranium may be a primitive form of *H. sapiens*. It was named "*H. longi*" in 2021 by Qiang Ji et al., who published in The Innovation. The skull is nearly complete and massive. The brain capacity is large; within the range of Neanderthals and *H. sapiens*. In some ways the skull has features that are similar to *H. sapiens* - flat, low cheekbones with shallow canine fossa and a face that is reduced and tucks underneath the braincase. This species is closer related to *H. sapiens* than Neanderthals. Thus, a third East Asian lineage is formed by "*H. longi*" (Denisovans) and other referred fossils. Chris Stringer studied the remains with the team, as he groups *H. longi* with *Homo daliensis*. It has been suggested the skull was from water-laid sediments 138,000-309,000 years old. It has been suggested "*H. longi*" evolved with the other species *Homo*, and may lead insight into the Denisovans.

### Evolution

Sequenced mtDNA fund preserved by the freezing cave climate extracted from Denisova 3 by Johannes Krause and Svante Pääbo differs from apomorphic Homo by 385 nucleotides of ~16,500, much larger than the mtDNA of the Neanderthal but much less than chimpanzees. This suggests Denisovans diverged from apomorphic humans and Neanderthals ~1,313,500-799,300 years ago, much earlier than the human/Neanderthal divergence. Krause et al. concluded Denisovans descend from an early *Homo erectus* migration out of Africa, fully distinct from contemporaneous hominins. However, the nDNA of Denisova 3 bears a unique level of preservation with low-level contamination and shows Neanderthals and Denisovans are closer to each other than to apomorphic humans. Using the *Homo*-*Pan* last common ancestor, the two split from the apomorphic lineage ~804,000 years ago, with this second lineage splitting ~640,000 years ago. With a 1x10−9 or 0.5x10−9 per base pair/year mutation rate, the second lineage split would have occurred between 236-190 thousand years ago or 473-381 thousand years ago. With 1.1x10−8 per generation with a new generation every 29 years, this date moves to 744,000 years. With 5x10−10 nucleotide site per year, it moves down to 616,000 years. Using the latter, the split probably occurred when hominins had dispersed across Europe. *Homo heidelbergensis* is sometimes an ancestor species to Denisovans. Since their dental anatomy differs strongly, they may have split before Neanderthal dentition had evolved ~300,000 years ago.

Divergent DNA of Denisovans is interpreted as admixture with an unknown plesiomorphic population, possibly relict *Homo erectus* or a similar species ~53,000 years ago. Alternatively, this would have been a persistence of an ancient lineage that went extinct through human and Neanderthal genetic drift. Apomorphic Homo sapiens may have contributed to mtDNA in Neanderthals, but not Denisovans. MtDNA of a 400,000-year-old femur of *Homo heidelbergensis* from Sima de los Huesos was found to be related to Neanderthals and Denisovans, being closer related to Denisovans. The authors posit this as an plesiomorphic sequence of mtDNA that was lost in Neanderthals due to apomorphic humans replacing it. The earliest Denisovans inhabited the Tibetan Plateau 160,000 years ago, where they evolved adaptation to altitude. It is unknown if Denisovans resided there year-round, or if their arrival at the plateau was unplanned for, and they retreated to lowlands.

### Genetics

As much as 17% of Denisovan genome is accumulated from local Neanderthal populations. Denisova 11 is a first generation Denisovan/Neanderthal hybrid, which indicates interbreeding was possible and did happen. Their genome shares derived alleles with Altai Neanderthals than with the Vindija and Mezmaiskaya Neanderthals; this suggests gene flow originates from a population closer to the Altai Neanderthals. However, the first generation's Denisovan father has typical Altai Neanderthal introgression, with her Neanderthal mother was closer to the Vindija Neanderthals. ~4% of their genome comes from from an unknown plesiomorphic human, which could be the source of anomalous mtDNA, which indicates this species may have diverged from humans and Neanderthals ~1 million years ago. The only *Homo* from Late Pleistocene Asia are *Homo erectus* and *Homo heidelbergensis*. Before Denisovans split from their ancestor "Neandersovans" migrating from Africa interbred with a "superarchaic" species that already lived in Europe, descending from a very early Out-Of-Africa migration ~1.9 million years ago.

One 2011 study find Denisovan DNA in Aboriginal Australians, Near Oceanians, Polynesians, Eastern Indonesians and Mamanwans, but not East Asians, Western Indonesians, the Jahai or the Onge. This means Denisovans introgress within the Pacific rather than the Asian mainland, that their ancestors of the latter were not found in Southeast Asia at the time, finding East Asia was founded by two migrations of apomorphic humans. In Melanesians, ~4-6% or 1.9-3.4% comes from Denisovan introgress, with New Guineans and Aboriginal Australians having the most introgression, with Australians having less than New Guineans. In 2021, it was found 30-40% of ancestry lies in Filipinos more then Papuans, having ~5%. This team found the highs Aeto genome levels. Less Denisovan ancestry is found in the Papuans, seen in the X chromosomes and some autosomes, suggesting hybrid incapability. The former could be explained by less female introgression into apomorphic *Homo sapiens*, or more female apomorphic *Homo sapiens* immigrants that diluted Denisovan X chromosome ancestry. 0.2% of mainland Asians and Native Americans comes from Denisovans, with South Asians having a similar level to East Asians. The Tianyuan Man lacks Denisovan DNA not seen in apomorphic East Asians, discounting the dilution hypothesis.

In 2018, the Han Chinese, Japanese, and Dai genomes were found to have DNA from two Denisovan populations, with one observable similar to the population in Papuans, with the second closer to the Denisova Denisovans. This could indicate two introgression events with two populations. In South Asians, only one introgression is seen, shared with Papuans. Admixture in Papuans is thought to mold modern people's immune systems, although it is unknown how this happened. In 2019, a third wave was found with East Asians, which may have happened right when apomorphic humans immigrated to the region. In other regions, plesiomorphic introgressions stems from the Vindija Neanderthals, opposing the plesiomorphics related to them and Denisovans, About 13.1% and 3.3% of plesiomorphic DNA in Icelanders descends from the later groups, with such a high percentage indicating a west Eurasian Denisovan population introgressing into the Vindija Neanderthals or immigrating apomorphic humans. Denisovan genes may have helped the latter acclimative when coming from Africa. The distribution and divergence of HLA-B\*73 and other HLA alleles leads to introgression from Denisovans to apomorphic humans in West Asia. In 2011, it was found half of HLA alles in apomorphic Eurasian represent plesiomorphic HLA haplotypes, inferred to be Denisovan or Neanderthal. A haplotype of EPAS1 in Tibetans likely came from Denisovans, showing an adaptation to high altitudes. Phospholipis transporter and trace amine-associated receptors are more active in people with more Denisovan ancestry. Their genes may have conferred a degree of immunity against the G614 mutation of SARS-CoV-2. The Ayta Magbujon people of the Phillipines have the highest percentage of Denisovan DNA.

Denisovans are known to have lived in Eurasia, but newer studies present the idea that some populations migrated to the rainforests of southeast Asia, leading to possible new species and Asian ancestry . In 2019, genetics studies based on artificial intelligence determined a non-Neanderthal and non-Denisovan ancestor species is present in the apomorphic human genome. Such a species is unique, and was previously unknown .

### Taxonomy

Two papers were published simultaneously by Ji et al. (2021) that separated the Harbin cranium (HBSM2018-000018(A)) as a new species, named *Homo longi*. The Harbin skull is very close to the Dali skull, and was named *H. sapiens daliensis* in 1978 by Xinzhi Wu, who later abandoned it. If the Middle Pleistocene Asian *Homo* are one species, *Homo daliensis* takes priority. They recommended restricting *Homo daliensis* and argued for *H. longi*'s distinctiveness, referring the Dali and Hualong material to *Homo daliensis*. They argue two species of *Homo* lived in Middle Pleistocene China. It was also suggested H. longi is closest allied to the Xiahe mandible, which has been referred to Denisovans. They also state these specimens are closest allied to apomorphic humans than the latter is to European *Homo neanderthalensis*. However, Chris Stringer and colleagues argue for the Harbin skull being simply a Denisovan, as it sits too close to the Xiahe mandible to be distinguished. Additionally, they note the Harbin cranium bears a striking resemblance to the Dali cranium and other alleged Denisovan crania. Jie et al. (2021) find:

Supplementary data finds the following as included in the Denisovan specimen pool:

## Paleobiology

### Distribution

Denisovans lived in Siberia, with the Xiahe mandible being the earliest Tibetan Plateau occurrence. Though Denisovan material indicates a restricted range, they spanned all of East Asia and possibly western Eurasia. Guy Jacobs et al. (2019) found 3 distinct populations in Siberia, New Guinea and Oceania, across Asia. With coalescent modeling, the Denisova Denisovans split from the second population 283,000 years ago and from the third 363,000 years ago, indicting considerable reproductive isolation between them. High percentages of Denisovan DNA in Papuans and Australians suggest they may have crossed the Wallace Line, being the first *Homo* to do this. They may have also entered the Philippines and lived with *Homo luzonensis*. This may suggest they are closely related or the same species. They may have needed to cross bodies of water. Icelanders have a high amount of Denisovan DNA, either stemming from a far western population or having been sourced elsewhere and carried over as background. Neanderthals frequently made long crossings between Europe and the Altai mountains, especially late into their temporal range.

Exponential distribution analysis on haplotype lengths were used by Jacobs to find apomorphic human introgression occurring 29,900 years ago in the second and 45,700 for the third population. The late date of the second population indicates a possible persistence until 14,500 years ago, making them the latest surviving plesiomorphic species. New Guineans have introgression from the last two populations. A third wave appears to have introgressed in Eastern Asia, but not enough DNA evidence exists to find a timeframe. MtDNA from Denisova 4 is very similar to that of 3, indicating they were the same population. The genetic diversity of the Denisova Denisovans is seen as lower compared to apomorphic lineages, comparable to Neanderthals. However, its inhabitants were reproductively isolated from others across their range, with higher genetic diversity. Inhabitance of Denisova Cave swings from a fairly warm and humid pine/birch forest to a forest/tundra. Baishiya Karst, being at a high elevation, had a low temperature, low oxygen and with poor resources; inhabitance of high altitudes, due to harsh conditions, was previously assumed achieved by apomorphic humans. They also lived in the jungles of Southeast Asia.

### Artifacts

Early Middle Pleistocene discoidal and Kombewa cores are known from Denisova, with Levallois cores and flakes also found. Scrapers, denticulate and notched tools are deposited 287±41 thousand years ago in the Main Chamber, up to 170±19 thousand and 187±14 thousand years ago in the Main and East Chambers. Middle Pleistocene flat, discoid and Levallois cores and some sub-prismatic cores are present. Side scrapers, end scrapers, burins, chisel-like tools and truncate flakes and notched, denticulate tools are also found. These date to 156±15 thousand years ago in the Main, up to 170±19 thousand years ago in the South and 187±14 thousand years ago in the Main and East Chambers. Early Upper Pleistocene artifacts date to 44±5 thousand in the Main, 63±6 thousand in the East and 47±8 thousand years ago in the South Chamber; however, some East layers may have been disturbed. Blades and Levallois were made, but scrapers are dominant. Well-developed stone bladelets distinct from other scrapers start to accumulate in the Main Chamber 36±4 thousand years ago. Upper Pleistocene layers are comprised of bone tools and ornaments, including a marble ring, an ivory ring, an ivory pendant, an red deer tooth pendant, a chloritolite bracelet and a bone needle. However, Denisosvans are confirmed to have lived in the cave until 55,000 years, and these tools overlap with apomorphic human migration into Siberia, but none occur in the Altai region; Denisova 14 is too damaged to confirm which species made these, so it is unsure who made these complicated artifacts and tools. The cave was discovered from Stratum 11, which was dated to 40-50,000 years using oxygen isotope analysis.

### Art

Hand and foot print impressions from the 200,000 year-old Tibetan Plateau may represent evidence of parietal art produced by two children 7 and 12 years old. However, it is unknown which species produced these prints; the authors suggested Denisovans may have, but also note *Homo erectus* lived in the region, so their identification is left vague. These artworks are the first hominin evidence at Quesang. The prints are made of travertine, which forms a sort of sludge when first deposited. This would have allowed for the children to easily stick their hands and feet into the substance, which would harden when deprived of water.

### Palaeopathology

The Harbin skull bears shallow dents surrounding the bregma, which may have formed due to a healed injury. The second upper molar does not appear to contact the third molar, which means the third molar was either small and created a gap, or this individual did not have that tooth.

## Paleoecology

Given the Harbin skull had a long history, the exact provenience, stratigraphy, context and age are difficult to determine. Qingfeng Shao et al. (2021) performed multiple non-destructive techniques on the skull: x-ray fluorescence, rare-earth element and strontium isotope analyses on the skull and various other mammalian material from Dongjiang Bridge. They found all of them originate from the same region and time - all occurring ~12 meters (39 feet) below the surface, correlating 30 meters (98 feet) in the Huangshan Formation, 309-138,000 years ago. Uranium-thorium dating on direct points of the skull yield many dates. For example, one being 62-296 thousand years ago, likely as a result of uranium leaching. Statistically, they determined a minimum age of ~146,000 years,  but a more exact date is difficult to find due to the unidentified provenience. Therefore, the skull is constrained to the Middle Pleistocene and is roughly contemporaneous with Xiahe, Jinniushan, Dali and Hualong Cave specimens.

Due to the Ice Ages, the earth swings from cold glacial periods to considerably warmer interglacial periods. 300-130,000 years ago marks the Penultimate Glacial Period, where permafrost may have stretched to Harbin, but evidence of glaciations this far back is lacking. Northeast China bore the *Mammuthus*-*Coelodonta* Fauna assemblage during the Middle Pleistocene. This roster of animals includes cold steppeland-adapted animals. The locality the Harbin cranium is known from preserve *Sinomegaceros ordosianus*, Przewalski's horse, elk, *Bubalus wansjock* and the brown bear.

## Cryptozoology

Some such as Ramakrishnan postulated that the Hindu god Hanuman, sometimes stated in epics to have been fluent in another language and be from a kind with a distinct culture, to be distinct species; Ramakrishnan suggested that it be assigned to *Homo erectus* (based on the Narmada cranium, now thought to be a Denisovan) or even *Homo floresiensis*. Others such as Ciochan suggest that the text does not clearly state a cultural difference and the description of the character does not match with the ape bauplan. Still, the original epic, uninterpreted, script states that Hanuman is has apelike features, not that he is a monkey.

## Synonyms

* *H. neanderthalensis mapaensis*
* *H. heidelbergensis mabaensis*
* *H. heidelbergensis daliensis*
* *H. erectus narmadensis*
* *H. erectus mapaensis*
* *H. sapiens mabaensis*
* *H. sapiens daliensis*
* "H. sapiens altaiensis" (rejected)
* *H. longi* (=Harbin)
* *H. tsaichangensis* (=Penghu)
* *H. mabaensis*/*mapaensis* (=Dali)
* "H. altaiensis" (rejected)
* "H. denisova" (rejected)
* "H. denisoviensis"
* "H. denisovensis"

## Notable Specimens

* Denisova 2: The deciduous 2nd lower molar of an adolescent woman given Genbank accession KX663333.
* Denisova 3 "X Woman": A 13.5-year-old adolescent woman known from the fifth distal finger phalanx. Her GenBank accession is NC013993.
* Denisova 4: The permanent upper 2nd and 3rd molar of an adult man. His GenBank accession is FR695060.
* Denisova 8: The permanent upper 3rd molar of an adult man, assigned GenBank accession KT780370.
* Denisova 11 "Denny": A 13-year-old woman who was a Neanderthal x Denisovan hybrid, known from an arm/leg fragment assigned GenBank accession KU131206.
* Denisova 13: A parietal fragment, still undescribed.
* Denisova 19 (DC8846), 20 (DC7795) and 21 (DC8591): Three specimens described in 2021. They were aged and genetically examined. Though these bone scraps are undiagnostic, these methods have allowed for them to be confidently identified by Brown et al. (2021) as Denisovans.
* Xiahe Mandible: A partial Denisovan mandible from Baishiya Karst Cave.
* "Dali skull": A robust skull similar to the Harbin skull.
* HBSM2018-000018(A) "Harbin skull/cranium" or the "Dragon Man": In 1933, a contract labourer found a near-complete cranium on Songhua River's bank during the construction of the Dongjiang Bridge in Harbin, Heilongjiang. The skull's importance was recognized, likely because of the Peking Man being discovered in 1929. The labourer hid the skull in a well that had been abandoned so Manchukuo authorities would not find it. After the Soviets had defeated Manchukuo, he concealed his former employment from Manchukuo National Railway from the the Nationalist (and later Communist) authorities. Before the man had died, his third-generation family were told of the remains and they were retrieved in 2018. Anthropologist Qiang Ji persuaded them to donate the skull to Hebei GEO University where it would be studied and stored. In 2021, the remains were named as "*Homo longi*", who compared it to other *Homo* phylogenetically rather than morphologically. *Homo longi* derives from the apomorphic province rather than the city Harbin, where it was found. Heilongjiang (known as Binjiang Province of Manchukuo in 1933). The name, Hei-long-jiang (黑龍江), translates to "Black Dragon River" in Mandarin, hence the nickname "Dragon Man".
* Xujiayao hominin?: 12 parietals, a temporal, 2 occipitals, a mandibular fragment, a juvenile maxilla and 3 teeth from Xujiayao, bearing traits seen in *Homo sapiens* and *Homo erectus*. This is characteristic of the enigmatic Homo longi, which is considered to be Denisovan. Thus, the hominins at this site are likely Denisovans. A 2022 study finds brain expansion that falls into the range of *Homo sapiens* and Neanderthals and demonstrates that brain growth varied among hominin groups. XJY 6 is the posterior of a braincase made from occipital PA1486 (No.7/XJY 6a), right parietal PA1490 (No.10/XJY 6b) and left temporal PA1498 (No.17/XJY 15). Xujiayao 6 is 160,000 years old, making it the oldest cranium in the *H. daliensis* clade and thus an early Denisovan.
* "Jinniushan hominin"?: A single woman recovered from Jinniushan, initially believed to have been from a male due to their great size. A single skull, 6 vertebrae (one cervical and five thoracic), a left os coxae, the left ulna, the left patella, two left ribs and various hand and feet bones were found. She was probably similar to apomorphic women in many ways.
* "Narmada /human/Man/hominin/cranium/skull" or the "Hathnora skull/cranium"?: The Narmada skull was found on December 5, 1982 on the Narmada banks, where Arun Sonakia found the only Indian ancient human remains known aside from stone tools. However, the Naramada skull is obscure, sometimes grouped into *Homo erectus* proceeding *Homo sapiens*. It is poorly dated, but was found to be 25-30 years old and lived at least 400,000 years ago. Dam building, mining, drilling, agriculture and population pressure, however, limit study of Narmada, since many sites are being destroyed and paleontology is not well developed in India. It has also been called *Homo heidelbergensis*, but may be a Denisovan. It was first thought to have been the ancestor to apomorphic Indians.
* Penghu 1?: Penghu 1 was discovered before 2008 by fishermen who hauled it from a 60-120-meter depth ~25 kilometers off Taiwan in the Pescadores Channel. This area was once land, but sea levels rising lead to it to be submerged. In 2015, it was described and is now deposited at the National Museum of Natural Science. It is under 450,000 years old, probably 190-130,000 or 70-10,000. It is a near-complete right mandible with 4 teeth (worn molars and premolars) with a high robust index (including a robust lateral torus, molaes and a bicondylar breadth, the latter was found from 3D replication). These confirm the Middle-Late Pleistocene age. The alveoli of the incisors and right canine are well-preserved with great length. With jaw morphology it was assigned *Homo*. A receding anterior surface and a lack of a chin distinguishes it from *Homo sapiens*, with derived traits similar to *Homo habilis* (including a short jaw width) The agenesis of the M3 also place it in *Homo*. It is argued which species Penghu 1 belongs to. It was first assigned *Homo erectus*, but the authors decided to call it "The first Archaic *Homo* of Taiwan", explained by retention of plesiomorphic characters from *Homo erectus*, a robust-jawed *Homo* migration from Africa, A *Homo heidelbergensis* or an adapted *Homo erectus*. Yousuke Kaifu, a coauthor, cautioned skeletal elements would be needed to assign a species. However, Mark McMenamin argues unique dental characters should call for a new species that he called *Homo tsaichangensis*, where he found convergent evolution with *Gigantopithecus* cf. *blacki* (which he thinks may be why some early anthropologists thought it may have been a giant *Homo* or *Australopithecus*). The two have similar tooth morphology, which he suggests as a similar diet. Others think it is plesiomorphic *Homo sapiens* or even a Denisovan since it is very similar morphologically and phylogenetically to the Xiahe mandible. It is also noted it is too fragmentary to declare as new.
* Hualongdong no. 6 or HLD 6: A skull sharing a mosaic of traits, with some shared with *Homo erectus*. It was discovered between July and September 2006 by Han Ligang et al.. ~100 stone tools and ~20 bone artifacts were recovered with it. It is a 14-15-year-old boy with a cranial capacity of 1500, a low, broad skull, a large brow and no midface prognathism. They were able to reconstruct his face. Analysis finds him to be part of a sister lineage to *Homo sapiens*, closer to the latter than to Neanderthals. It bears morphology similar to suspected Denisovan specimens, so it should likely be included into *H. daliensis*.
* "Maba Man/cranium"?: A cranium found in 1958 near Maba by farmers, found to be intermediate between *Homo erectus* and *Homo sapiens*. It seems to bear the morphology of west Asian Neanderthals, but is undiagnostic. It consists of a skull cap and partial right upper face with parts of the nose attached. At first, the large brow led to a connection with *Homo erectus*. The braincase is low and thick, with a brain that was likely larger than *Homo erectus*, though the cranial capacity can not be determined due to incompleteness. At some point, it was named "*Homo mabaensis*" . A lesion on the skull shows this individual sustained a violent trauma but survived, possibly caused by another hominid. The wound is very similar to wounds caused by other individuals striking another over the head with a heavy object. This could be the first instance of human-induced trauma and intraspecific combat.
* Xuchang 1-2: A skullcap and the back of the braincase (1) and the posterior braincase (2) described in 2017 from China: it were then informally attributed to Denisovans. At the time, however, it was difficult to determine if the crania were connected to Denisovans based on limited understanding of population dynamics, no overlapping elements and range. They were discovered at Lingjing, Xuchang during excavations between 2007 and 2014. Later phylogenetic analyses would confirm Xuchang 1-2 as Denisovan.
* TNH2-1: The tooth of a female child was found in Tam Ngu Hao 2 cave and is most likely to represent a Denisovan due to morphological overlap. The tooth is about 164-131,000 years old, and suggests that Denisovans had taken mainland Asia south of the Altai region. Finding such hominin diversity in Asia at this time suggests that it was a hotspot. The tooth also has traits similar to Neanderthals, but morphology suggests otherwise.

### Artifacts

* Pandado: A site in discovered in central Narmada Valley, India in 2022. From here, Acheulean industry tools were described by Vivek Singh and Shantanu Katiyar. This makes it the second site in India representing Denisovan habitation since the discovery of the Narmada hominin.

## Gallery

Narmada cranium.Narmada cranium.A reconstruction of the Harbin skull, "*Homo longi*".
**Credit: Chuang Zhao.**Harbin cranium.Harbin cranium.Harbin cranium.Harbin cranium (left) and Dali cranium (right).Harbin cranium.Penghu 1.Tsaichangensis.jpg**Credit: Clumsystiggy on DeviantArt.**Hualongdong no. 6.On the top row is HLD 6, including a restored model. In the middle is a facial reconstruction of HLD6. The bottom row are crania with similar morphology used to restore the skull and face.Maba cranium.Denisova 11 "Denny".Denisova 17 is a Neanderthal, Denisova 18 is *H.* sp. and Denisova 19-21 are Denisovan.Xujiayao 6.Xuchang 1.Xuchang 2.Dali cranium.Tooth from Laos.
## References

1. ↑ https://www.nhm.ac.uk/events/nature-live-online-dragon-man.html
2. ↑ https://the-past.com/news/ancient-human-relative-identified-in-china/
3. ↑ https://worldofpaleoanthropology.org/2021/08/16/dragon-man-with-chris-stringer/?fbclid=IwAR3RsL2OHcaRoIq82JesVEGOrKUpknB8juv2yfyBDPPCQVdkWm1SmHio-xY
4. ↑ 4.0 4.1 4.2 4.3 4.4 4.5 4.6 https://en.wikipedia.org/wiki/Denisovan
5. ↑ https://en.wikipedia.org/wiki/Madjedbebe
6. ↑ https://australian.museum/learn/science/human-evolution/the-spread-of-people-to-australia/
7. ↑ https://www.science.org/doi/10.1126/sciadv.aaw3950
8. ↑ 8.0 8.1 8.2 8.3 8.4 8.5 https://www.nhm.ac.uk/discover/news/2021/june/ancient-skull-from-china-could-be-new-species-dragon-man.html?utm\_source=tw-link-post-20210625-kp&utm\_medium=social&utm\_campaign=news
9. ↑ 9.0 9.1 9.2 9.3 9.4 9.5 https://www.cell.com/the-innovation/fulltext/S2666-6758(21)00055-2#.YNX324UW9vo.twitter
10. ↑ 10.0 10.1 10.2 10.3 10.4 10.5 https://twitter.com/ChrisStringer65/status/1408440483039301635
11. ↑ 11.0 11.1 11.2 11.3 11.4 11.5 https://www.cell.com/the-innovation/fulltext/S2666-6758(21)00057-6
12. ↑ 12.0 12.1 12.2 12.3 12.4 12.5 https://www.cell.com/the-innovation/fulltext/S2666-6758(21)00056-4#%20
13. ↑ 13.0 13.1 13.2 13.3 13.4 13.5 https://www.sciencemag.org/news/2021/06/stunning-dragon-man-skull-may-be-elusive-denisovan-or-new-species-human
14. ↑ 14.0 14.1 14.2 14.3 14.4 14.5 https://kar.kent.ac.uk/74280/1/Xiahe\_Main.pdf
15. ↑ 15.0 15.1 15.2 15.3 15.4 15.5 https://www.nature.com/articles/s41586-019-1139-x
16. ↑ 16.0 16.1 16.2 16.3 16.4 16.5 https://repositori.upf.edu/bitstream/handle/10230/25596/Marques\_nat\_gen.pdf;jsessionid=B41688C4A3F277E6CC161E2B93B42B5A?sequence=1
17. ↑ 17.0 17.1 17.2 17.3 17.4 17.5 https://www.nature.com/articles/nature09710
18. ↑ 18.0 18.1 18.2 18.3 18.4 18.5 http://www.inderscience.com/offer.php?id=16747
19. ↑ 19.0 19.1 19.2 19.3 19.4 19.5 https://www.mdpi.com/2571-550X/3/4/32
20. ↑ 20.0 20.1 20.2 20.3 20.4 20.5 https://twitter.com/Tweetisaurus/status/1441445516903026697
21. ↑ 21.0 21.1 21.2 21.3 21.4 21.5 https://en.wikipedia.org/wiki/Homo\_longi
22. ↑ 22.0 22.1 22.2 22.3 22.4 22.5 https://arstechnica.com/science/2021/07/is-the-dragon-man-skull-actually-from-a-new-hominin-species/
23. ↑ https://twitter.com/ChrisStringer65/status/1512005997883240457
24. ↑ https://www.sciencedaily.com/releases/2021/12/211207152519.htm
25. ↑ https://www.scientificamerican.com/article/dna-from-extinct-human-relative-may-have-shaped-modern-papuans-immune-system/
26. ↑ https://t.co/XSWf9TfVZJ?amp=1
27. ↑ https://scitechdaily.com/new-dna-evidence-in-search-for-the-mysterious-denisovans/
28. ↑ https://www.nature.com/articles/s41559-021-01408-0
29. ↑ http://www.sci-news.com/othersciences/anthropology/southeast-asians-denisovan-dna-09480.html
30. ↑ https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6335398
31. ↑ https://www.sciencealert.com/artificial-intelligence-identifies-unknown-ghost-ancestor-in-the-human-genome
32. ↑ https://twitter.com/ChrisStringer65/status/1409418572984729600
33. ↑ https://johnhawks.net/weblog/an-enormous-sample-sheds-light-on-the-denisovan-ancestry-of-people-in-iceland/
34. ↑ https://www.sciencedirect.com/science/article/abs/pii/S1563011008000378
35. ↑ https://www.scientificamerican.com/article/kids-fossilized-handprints-may-be-some-of-the-worlds-oldest-art/
36. ↑ https://www.sciencedirect.com/science/article/pii/S2095927321006174
37. ↑ https://www.ancientpages.com/2020/03/03/hindu-monkey-god-hanuman-may-have-been-homo-erectus-scientist-says/
38. ↑ https://www.scmp.com/week-asia/people/article/3048479/hindu-monkey-god-hanuman-really-homo-erectus
39. ↑ 39.0 39.1 https://nutcrackerman.com/2021/10/25/nuevo-grupo-hermano-linaje-sapiens-craneos-hualongdong-harbin/
40. ↑ https://fr.wikipedia.org/wiki/Homme\_de\_Denisova
41. ↑ https://www.nature.com/articles/s41559-021-01581-2
42. ↑ https://en.wikipedia.org/wiki/Dali\_(fossil)
43. ↑ https://en.wikipedia.org/wiki/Xujiayao
44. ↑ https://www.sciencedirect.com/science/article/abs/pii/S0047248421001718?via%3Dihub
45. ↑ https://www.newscientist.com/article/2305830-160000-year-old-fossil-may-be-the-first-denisovan-skull-weve-found/#ixzz7JCja1v7w
46. ↑ https://twitter.com/ChrisStringer65/status/1487351120628486148
47. ↑ https://en.wikipedia.org/wiki/Jinniushan
48. ↑ https://twitter.com/ChrisStringer65/status/1451459685182173184
49. ↑ https://www.downtoearth.org.in/coverage/loneliness-of-narmada-human-39817
50. ↑ https://www.sciencedirect.com/science/article/pii/S1631068305001272
51. ↑ https://en.wikipedia.org/wiki/Penghu\_1
52. ↑ http://www.ivpp.cas.cn/xwdt/kyjz/202110/t20211008\_6219503.html
53. ↑ https://www.britannica.com/topic/Maba-cranium
54. ↑ https://en.wikipedia.org/wiki/Maba\_Man
55. ↑ https://www.teachinganthropology.org/2018/08/06/teaching-complexity-uncertainty-and-lies-the-case-of-homo-heidelbergensis/
56. ↑ https://www.nbcnews.com/id/wbna45392341
57. ↑ https://www.science.org/doi/full/10.1126/science.aal2482
58. ↑ https://www.bbc.com/news/science-environment-39118752
59. ↑ https://www.sciencealert.com/this-ancient-tooth-once-belonged-to-the-mysterious-denisovans-scientists-think
60. ↑ https://t.co/mpq0ltmye0