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103-431: Linguistically, Papuans speak languages from the many families of non-Austronesian languages that are found only on New Guinea and neighboring islands, as well as Austronesian languages along parts of the coast, and recently developed creoles such as Tok Pisin , Hiri Motu , Unserdeutsch , and Papuan Malay . The term "Papuan" is used in a wider sense in linguistics and anthropology. In linguistics, " Papuan languages "

206-551: A juvenile female 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, but it is unclear whether Neanderthals and Denisovans ever cohabited in the cave. Additional specimens from Denisova Cave were subsequently identified, as was a single specimen from

309-510: A marble ring, an ivory ring, an ivory pendant, a red deer tooth pendant, an elk tooth pendant, a chloritolite bracelet, and a bone needle. However, Denisovans are only confirmed to have inhabited the cave until 55 ka; the dating of Upper Paleolithic artefacts overlaps with modern human migration into Siberia (though there are no occurrences in the Altai region); and the DNA of the only specimen in

412-538: A relict H. erectus or H. erectus -like population about 53,000 years ago. Alternatively, divergent mtDNA could have also resulted from the persistence of an ancient mtDNA lineage which only went extinct in modern humans and Neanderthals through genetic drift . Modern humans contributed mtDNA to the Neanderthal lineage, but not to the Denisovan mitochondrial genomes yet sequenced. The mtDNA sequence from

515-626: A single landmass for most of their human history, having been separated by the Torres Strait only 8000 years ago, and that a deep reconstruction would likely include languages from both. However, Dixon later abandoned his proto-Australian proposal, and Foley's ideas need to be re-evaluated in light of recent research. Wurm also suggested the Sepik–Ramu languages have similarities with the Australian languages, but believed this may be due to

618-447: A 12 year old modern human child, and the middle finger length agrees with a 17 year old modern human. One of the handprints shows an impression of the forearm, and the individual was wiggling their thumb through the mud. Analyses of the modern human genomes indicate past interbreeding with at least two groups of archaic humans, Neanderthals and Denisovans, and that such interbreeding events occurred on multiple occasions. Comparisons of

721-482: 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. Denisovans may represent a new species of Homo or an archaic subspecies of Homo sapiens (modern humans), but there are too few fossils to erect a proper taxon . Proactively proposed species names for Denisovans are H. denisova or H. altaiensis . Chinese researchers suggest

824-518: A Denisovan population far west of the Altai Mountains. Genetic data suggests Neanderthals were frequently making long crossings between Europe and the Altai Mountains especially towards the date of their extinction. Using exponential distribution analysis on haplotype lengths, Jacobs calculated introgression into modern humans occurred about 29,900 years ago with the Denisovan population ancestral to New Guineans; and 45,700 years ago with

927-827: A composite of Usher's and Ross' classifications, Palmer et al. do not address the more tentative families that Usher proposes, such as Northwest New Guinea . The coherence of the South Bird's Head , East Bird's Head , Pauwasi , Kwomtari , and Central Solomons families are uncertain, and hence are marked below as "tentative." Papuan independent language families (43 families) Papuan isolates and unclassified languages (37 total) Glottolog 4.0 (2019), based partly on Usher, recognizes 70 independent families and 55 isolates. The following families are identified by Timothy Usher and Edgar Suter in their NewGuineaWorld project: In addition, poorly attested Karami remains unclassified. Extinct Tambora and

1030-427: A fairly warm and moderately humid pine and birch forest to tundra or forest-tundra landscape. Conversely, Baishiya Karst Cave is situated at a high elevation, an area characterized by low temperature, low oxygen, and poor resource availability. Colonization of high-altitude regions, due to such harsh conditions, was previously assumed to have only been accomplished by modern humans. Denisovans seem to have also inhabited

1133-537: A high percentage (roughly 5%) occurring in Melanesians , Aboriginal Australians , and Filipino Negritos . 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

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1236-406: A language, they are short and utilise a reduced set of the language's phonemic inventory . Both phenomena greatly increase the possibility of chance resemblances, especially when they are not confirmed by lexical similarities. Sorted by location north Irian : Sandaun Province : Sepik River : Bismarck Archipelago : Former isolates classified by Ross: Languages reassigned to

1339-770: A lesser extent, across Asia. Using coalescent modeling , the Denisova Cave Denisovans split from the second population about 283,000 years ago; and from the third population about 363,000 years ago. This indicates that there was considerable reproductive isolation between Denisovan populations. In a 2024 study, scientist Danat Yermakovich, of the University of Tartu , discovered that people living at different elevations in Papua New Guinea have differences in Denisovan DNA; with people living in

1442-424: A long, broad, and projecting face; large nose; sloping forehead; protruding jaw; elongated and flattened skull; and wide chest and hips. The Denisovan tooth row was longer than that of Neanderthals and anatomically modern humans. Middle-to-Late Pleistocene East Asian archaic human skullcaps typically share features with Neanderthals. The skullcaps from Xuchang feature prominent brow ridges like Neanderthals, though

1545-460: A low-oxygen environment, likely came from Denisovans. Genes related to phospholipid transporters (which are involved in fat metabolism ) and to trace amine-associated receptors (involved in smelling) are more active in people with more Denisovan ancestry. Denisovan genes may have conferred a degree of immunity against the G614 mutation of SARS-CoV-2 . Denisovan introgressions may have influenced

1648-464: A million years ago. The only identified Homo species of Late Pleistocene Asia are H. erectus and H. heidelbergensis , though in 2021, specimens allocated to the latter species were reclassified as H. longi and H. daliensis . Before splitting from Neanderthals, their ancestors ("Neandersovans") migrating into Europe apparently interbred with an unidentified "superarchaic" human species who were already present there; these superarchaics were

1751-490: A mutation rate of 1 × 10 or 0.5 × 10 per base pair (bp) per year, the Neanderthal/Denisovan split occurred around either 236–190,000 or 473–381,000 years ago respectively. Using 1.1 × 10 per generation with a new generation every 29 years, the time is 744,000 years ago. Using 5 × 10 nucleotide site per year, it is 616,000 years ago. Using the latter dates, the split had likely already occurred by

1854-524: A number of instances". However, he considered this not evidence of a connection between (Great) Andamanese and Trans–New Guinea, but of a substratum from an earlier migration to New Guinea from the west. Greenberg also suggested a connection to the Tasmanian languages . However, the Tasmanian peoples were isolated for perhaps 10,000 years, their disappearance wiped out their languages before much

1957-468: A population more closely related to Vindija Neanderthals. Denisova 25, dated to 200ka, is estimated to have inherited 5% of his genome from a previously unknown population of Neanderthals, and came from a different population of Denisovans than the younger samples. About 4% of the Denisovan genome derives from an unidentified archaic hominin, perhaps the source of the anomalous ancient mtDNA, indicating this species diverged from Neanderthals and humans over

2060-427: A simple sister group of Tianyuan " ("Basal East Asians"). There is evidence that the ancestors of Papuans and related groups "underwent a strong bottleneck before the settlement of the region, and separated around 20,000–40,000 years ago". Papuans display pronounced genetic diversity, explained through isolation and drift between different subgroups after the settlement of New Guinea . The most notable differentiation

2163-596: A single ancestral language... when a language is termed 'Papuan', this claims nothing more than that a language is not Austronesian. Most Papuan languages are spoken by hundreds to thousands of people; the most populous are found in the New Guinea Highlands , where a few exceed a hundred thousand. These include Western Dani (180,000 in 1993) and Ekari (100,000 reported 1985) in the western (Indonesian) highlands, and Enga (230,000 in 2000), Huli (150,000 reported 2011), and Melpa (130,000 reported 1991) in

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2266-555: A substratum effect, but nevertheless believed that the Australian languages represent a linguistic group that existed in New Guinea before the arrival of the Papuan languages (which he believed arrived in at least two different groups). The West Papuan , Lower Mamberamo , and most Torricelli languages are all left-headed , as well as the languages of New Britain and New Ireland . These languages all have SVO word order , with

2369-483: A total of 1083 languages, with 12 languages overlapping. They can be divided into two groups, the Austronesian languages , and all the others, called Papuan languages for convenience. The term Papuan languages refers to an areal grouping , rather than a linguistic one. So-called Papuan languages comprise hundreds of different languages, most of which are not related. The following indigenous peoples live within

2472-815: A variant region around the EPAS1 gene that in Tibetans assists with adaptation to low oxygen levels at high elevation, and in a region containing the WARS2 and TBX15 loci which affect body-fat distribution in the Inuit . In Papuans, introgressed Neanderthal alleles are highest in frequency in genes expressed in the brain, whereas Denisovan alleles have highest frequency in genes expressed in bones and other tissue. Early Middle Paleolithic stone tools from Denisova Cave included cores , scrapers , denticulate tools , and notched tools, deposited about 287±41 thousand years ago in

2575-678: Is a cover term for the diverse, mutually unrelated, non-Austronesian language families spoken in Melanesia , the Torres Strait Islands , and parts of Wallacea . In anthropology, "Papuan" is often used to denote the highly diverse aboriginal populations of Melanesia and Wallacea prior to the arrival of Austronesian-speakers, and the dominant genetic traces of these populations in the current ethnic groups of these areas. Ethnologue ' s 14th edition lists 826 languages of Papua New Guinea and 257 languages of Western New Guinea ,

2678-431: Is in contrast to the large, robust molars which are more similar to those of Middle to Late Pleistocene archaic humans. The third molar is outside the range of any Homo species except H. habilis and H. rudolfensis , and is more like those of australopithecines . The second molar is larger than those of modern humans and Neanderthals, and is more similar to those of H. erectus and H. habilis . Like Neanderthals,

2781-578: Is primarily derived from Ancient East Eurasians , which relates them to other mainland Asian groups such as the " AASI ", Andamanese, as well as East/Southeast Asians, although Papuans may have also received some gene flow from an earlier group (xOoA), around 2%, next to additional archaic Denisovan admixture in the Sahul region. Papuans may habor varying degrees of deep admixture from "a lineage basal to West and East-Eurasians which occurred sometimes between 45 and 38kya", although they are generally regarded "as

2884-462: Is suggested to date back at least 20kya, while the sub-structure among Highlanders dates back around 10kya, with higher diversity among western Highlanders than Eastern ones. The genetic diversity is paralleled by linguistic and cultural diversity. Based on his genetic studies of the Denisova hominin , an ancient human species discovered in 2010, Svante Pääbo claims that ancient human ancestors of

2987-582: Is the earliest recorded human presence on the Tibetan Plateau. Though their remains have been identified in only these three locations, traces of Denisovan DNA in modern humans suggest they ranged across East Asia , and potentially western Eurasia. In 2019, geneticist Guy Jacobs and colleagues identified three distinct populations of Denisovans responsible for the introgression into modern populations now native to, respectively: Siberia and East Asia; New Guinea and nearby islands; and Oceania and, to

3090-478: Is the oldest, followed by Denisova 8, while Denisova 3 and Denisova 4 were roughly contemporaneous. In 2024, scientists announced the sequence of Denisova 25, which was in a layer dated to 200ka. During DNA sequencing, a low proportion of the Denisova 2, Denisova 4 and Denisova 8 genomes were found to have survived, but a high proportion of the Denisova 3 and Denisova 25 genomes were intact. The Denisova 3 sample

3193-483: The Annamite Mountains since 2008, was directed by local children to the site Tam Ngu Hao 2 ("Cobra Cave") where they recovered a human tooth. The tooth (catalogue number TNH2-1) developmentally matches a 3.5 to 8.5 year old, and a lack of amelogenin (a protein on the Y chromosome ) suggests it belonged to a girl barring extreme degradation of the protein over a long period of time. Dental proteome analysis

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3296-846: The Austronesian family : Unclassified due to lack of data: Unaccounted for: Søren Wichmann (2013) accepts the following 109 groups as coherent Papuan families, based on computational analyses performed by the Automated Similarity Judgment Program (ASJP) combined with Harald Hammarström 's (2012) classification. Some of the groups could turn out to be related to each other, but Wichmann (2013) lists them as separate groups pending further research. 9 families have been broken up into separate groups in Wichmann's (2013) classification, which are: An automated computational analysis ( ASJP 4) by Müller, Velupillai, Wichmann et al. (2013) found lexical similarities among

3399-541: The Baishiya Karst Cave on the Tibetan Plateau , and Cobra Cave in the Annamite Mountains of Laos. DNA evidence suggests they had dark skin, eyes, and hair, and had a Neanderthal-like build and facial features. However, they had larger molars which are reminiscent of Middle to Late Pleistocene archaic humans and australopithecines . Denisovans apparently interbred with modern humans, with

3502-591: The East Papuan languages have not been addressed, except to identify Yele as an Austronesian language. Joseph Greenberg proposed an Indo-Pacific phylum containing the (Northern) Andamanese languages , all Papuan languages, and the Tasmanian languages , but not the Australian Aboriginal languages . Very few linguists accept his grouping. It is distinct from the Trans–New Guinea phylum of

3605-959: The Initial Upper Paleolithic , which is "ascribed to a population movement with uniform genetic features and material culture" ( Ancient East Eurasians ), and sharing deep ancestry with modern East Asian peoples and other Asia-Pacific groups. It is estimated that people reached Sahul (the geological continent consisting of Australia and New Guinea) between 50,000 and 37,000 years ago. Rising sea levels separated New Guinea from Australia about 10,000 years ago. However, Aboriginal Australians and Papuans had diverged genetically much earlier, around 40,000 years BP. Papuans are more closely related to Melanesians than to Aboriginal Australians. The majority of Papuan Y-DNA Haplogroups belong to subclades of Haplogroup   MS , and Haplogroup   C1b2a . The frequency of each haplogroup varies along geographic clines. The genetic makeup of Papuans

3708-490: The Lower and Middle Paleolithic , and lived, based on current evidence, from 285 to 25 thousand years ago. Denisovans are known from few physical remains; consequently, most of what is known about them comes from DNA evidence. No formal species name has been established pending more complete fossil material. The first identification of a Denisovan individual occurred in 2010, based on mitochondrial DNA (mtDNA) extracted from

3811-479: The Takia language has. The Reef Islands – Santa Cruz languages of Wurm's East Papuan phylum were a potential 24th family, but subsequent work has shown them to be highly divergent Austronesian languages as well. Note that while this classification may be more reliable than past attempts, it is based on a single parameter, pronouns, and therefore must remain tentative. Although pronouns are conservative elements in

3914-566: 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

4017-859: 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

4120-444: The 40,000-year-old Chinese modern human Tianyuan Man lacking Denisovan DNA significantly different from the levels in modern-day East Asians discounts the hypothesis that immigrating modern humans simply diluted Denisovan ancestry whereas Melanesians lived in reproductive isolation. A 2018 study of Han Chinese, Japanese , and Dai genomes showed that modern East Asians have DNA from two different Denisovan populations: one similar to

4223-600: The Altai Neanderthal genome from Siberia than with the Vindija Cave Neanderthal genome from Croatia or the Mezmaiskaya cave Neanderthal genome from the Caucasus, suggesting that the gene flow came from a population that was more closely related to the local Altai Neanderthals. However, Denny's Denisovan father had the typical Altai Neanderthal introgression , while her Neanderthal mother represented

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4326-526: The Australian languages, a later migration bringing the West Papuan, Torricelli and the East Papuan languages and a third wave bringing the most recent pre-Austronesian migration, the Trans–New Guinea family. Two of Wurm's isolates have since been linked as the and since Wurm's time another isolate and two languages belonging to a new family have been discovered, Foley summarized the state of

4429-544: The Denisovan DNA found in Papuan genomes, and a second that is closer to the Denisovan genome from Denisova Cave. This could indicate two separate introgression events involving two different Denisovan populations. In South Asian genomes, DNA only came from the same single Denisovan introgression seen in Papuans. A 2019 study found a third wave of Denisovans which introgressed into East Asians. Introgression, also, may not have immediately occurred when modern humans immigrated into

4532-475: The Denisovan, Neanderthal, and modern human genomes have revealed evidence of a complex web of interbreeding among these lineages. As much as 17% of the Denisovan genome from Denisova Cave represents DNA from the local Neanderthal population. Denisova 11 was an F1 (first generation) Denisovan/Neanderthal hybrid; the fact that such an individual was found may indicate interbreeding was a common occurrence here. The Denisovan genome shares more derived alleles with

4635-727: The Denisovans based on the similarity between the type specimen's molar and that of the Xiahe mandible. KU131206 Sequenced mitochondrial DNA (mtDNA), preserved by the cool climate of the cave (average temperature is at freezing point), was extracted from Denisova 3 by a team of scientists led by Johannes Krause and Svante Pääbo from the Max Planck Institute for Evolutionary Anthropology in Leipzig , Germany. Denisova 3's mtDNA differs from that of modern humans by 385 bases ( nucleotides ) out of approximately 16,500, whereas

4738-458: The Denisovans were members of Homo longi , and the idea has been supported by the palaeontologist Chris Stringer . 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 (Dyonisiy), a Russian Old Believer hermit who lived there in the 18th century. The cave was first inspected for fossils in

4841-580: The East Chamber dates to 217,000 years ago. Based on artifacts also discovered in the cave, hominin occupation (most likely by Denisovans) began 287±41 or 203±14 ka . Neanderthals were also present 193±12 ka and 97±11 ka, possibly concurrently with Denisovans. The fossils of multiple distinct Denisovan individuals from Denisova Cave have been identified through their ancient DNA (aDNA): Denisova 2, 3, 4, 8, 11 , and 25. An mtDNA-based phylogenetic analysis of these individuals suggested that Denisova 2

4944-510: The Indonesian province of Papua include: Papuan ethnic groups/tribes in the Indonesian province of Highland Papua include: Papuan ethnic groups/tribes in the Indonesian province of Central Papua include: Papuan ethnic groups/tribes in the Indonesian province of South Papua include: The origin of Papuans is generally associated with the first settlement of Australasia by a lineage dubbed 'Australasians' or 'Australo-Papuans' during

5047-423: The Indonesian province of Southwest Papua include Abun, Ambel, Batanta, Biak (Betew, Kafdaron, Bikar, Usba, Wardo), Biga, Butlih, Domu, Fiawat, Imekko (Inanwatan-Bira, Matemani-Iwaro, Kais-Awe, Kokoda-Emeyode), Irires, Ma'ya (Kawe, Langanyan, Wawiyai), Matbat, Maybrat (Ayamaru, Mare, Karon Dori, Ayfat, Aytinyo), Meyah, Moi-Ma'ya, Moi, Mpur, Nerigo, Tehit , Tepin, Yahadian, Yaben-Konda. Papuan ethnic groups/tribes in

5150-656: The Main Chamber of the cave; and about 269±97 thousand years ago in the South Chamber; up to 170±19 thousand and 187±14 thousand years ago in the Main and East Chambers, respectively. Middle Paleolithic assemblages were dominated by flat, discoidal, and Levallois cores, and there were some isolated sub-prismatic cores. There were predominantly side scrapers (a scraper with only the sides used to scrape), but also notched-denticulate tools, end-scrapers (a scraper with only

5253-741: The Pacific region rather than on the Asian mainland, and that ancestors of the latter groups were not present in Southeast Asia at the time. In the Melanesian genome, about 4–6% or 1.9–3.4% derives from Denisovan introgression. Prior to 2021, New Guineans and Australian Aborigines were reported to have the most introgressed DNA, but Australians have less than New Guineans. A 2021 study discovered 30 to 40% more Denisovan ancestry in Aeta people in

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5356-1145: The Papuan languages are spoken on the island of New Guinea, with a number spoken in the Bismarck Archipelago , Bougainville Island and the Solomon Islands to the east, and in Halmahera , Timor and the Alor archipelago to the west. The westernmost language, Tambora in Sumbawa , is extinct. One Papuan language, Meriam , is spoken within the national borders of Australia , in the eastern Torres Strait . Several languages of Flores , Sumba , and other islands of eastern Indonesia are classified as Austronesian but have large numbers of non-Austronesian words in their basic vocabulary and non-Austronesian grammatical features. It has been suggested that these may have originally been non-Austronesian languages that have borrowed nearly all of their vocabulary from neighboring Austronesian languages, but no connection with

5459-659: The Papuan languages of Timor has been found. In general, the Central–Eastern Malayo-Polynesian languages are marked by a significant historical Papuan influence, lexically, grammatically, and phonologically, and this is responsible for much of the diversity of the Austronesian language family. The "Papuan languages" are a strictly geographical grouping, and does not imply a genetic relationship . The concept of Papuan (non-Austronesian) speaking Melanesians as distinct from Austronesian -speaking Melanesians

5562-481: The Papuan people have among the highest rate of the newly evolved ASPM Haplogroup   D, at 59.4% occurrence of the approximately 6,000-year-old allele . While it is not yet known exactly what selective advantage is provided by this gene variant, the haplogroup   D allele is thought to be positively selected in populations and to confer some substantial advantage that has caused its frequency to rapidly increase. Papuan language The Papuan languages are

5665-567: The Papuan region is the Trans–New Guinea phylum , consisting of the majority of Papuan languages and running mainly along the highlands of New Guinea. The various high-level families may represent distinct migrations into New Guinea, presumably from the west. Since perhaps only a quarter of Papuan languages have been studied in detail, linguists' understanding of the relationships between them will continue to be revised. Statistical analyses designed to pick up signals too faint to be detected by

5768-482: The Papuans interbred in Asia with these humans . He has found that people of New Guinea share 4%–7% of their genome with the Denisovans, indicating this exchange. Denisovan introgressions may have influenced the immune system of present-day Papuans and potentially favoured "variants to immune-related phenotypes" and "adaptation to the local environment". In a 2005 study of ASPM gene variants , Mekel-Bobrov et al. found that

5871-1030: The Philippines than in Papuans , estimated as about 5% of the genome. The Aeta Magbukon in Luzon have the highest known proportion of Denisovan ancestry of any population in the world. In Papuans, less Denisovan ancestry is seen in the X chromosome than autosomes , and some autosomes (such as chromosome 11 ) also have less Denisovan ancestry, which could indicate hybrid incompatibility . The former observation could also be explained by less female Denisovan introgression into modern humans, or more female modern human immigrants who diluted Denisovan X chromosome ancestry. In contrast, 0.2% derives from Denisovan ancestry in mainland Asians and Native Americans . South Asians were found to have levels of Denisovan admixture similar to that seen in East Asians. The discovery of

5974-528: The Philippines, living alongside H. luzonensis which, if this is the case, may represent the same or a closely related species. These Denisovans may have needed to cross large bodies of water. Alternately, high Denisovan DNA admixture in modern Papuan populations may simply represent higher mixing among the original ancestors of Papuans prior to crossing the Wallace line. Icelanders also have an anomalously high Denisovan heritage, which could have stemmed from

6077-503: The South Chamber, though some layers of the East Chamber seem to have been disturbed. There was blade production and Levallois production, but scrapers were again predominant. A well-developed, Upper Paleolithic stone bladelet technology distinct from the previous scrapers began accumulating in the Main Chamber around 36±4 thousand years ago. In the Upper Paleolithic layers, there were also several bone tools and ornaments:

6180-456: The Tibetan Plateau, and since the Xiahe mandible is the oldest human fossil from the region (though younger than the Quesang impressions), these may have been made by Denisovan children. The impressions were printed onto a small panel of space, and there is little overlap between all the prints, so they seem to have been taking care to make new imprints in unused space. If considered art, they are

6283-482: The ZooMS analysis was there was no evidence of any other human group having occupied the cave. Other bone fragments included a large number of blue sheep , wild yaks , woolly rhino , spotted hyena , marmots , and other small mammals and birds. Examination of the animal bone surfaces indicates the Denisovans removed meat and bone marrow from the bones and also show the humans used them as raw material to make tools. There

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6386-404: The archaic DNA in the modern Icelandic genome descends from the Denisovans, and such a high percentage could indicate a western Eurasian population of Denisovans which introgressed into either Vindija-related Neanderthals or immigrating modern humans. Denisovan genes may have helped early modern humans migrating out of Africa to acclimatize . Although not present in the sequenced Denisovan genome,

6489-472: The cave dating to the time interval (Denisova 14) is too degraded to confirm species identity, so the attribution of these artefacts is unclear. In 1998, five child hand- and footprint impressions were discovered in a travertine unit near the Quesang hot springs in Tibet; in 2021, they were dated to 226 to 169 thousand years ago using uranium decay dating. This is the oldest evidence of human occupation of

6592-524: The classifications below. Joseph Greenberg proposed that the Andamanese languages (or at least the Great Andamanese languages ) off the coast of Burma are related to the Papuan or West Papuan languages. Stephen Wurm stated that the lexical similarities between Great Andamanese and the West Papuan and Timor–Alor families "are quite striking and amount to virtual formal identity [...] in

6695-440: The comparative method, though of disputed validity, suggest five major Papuan stocks (roughly Trans–New Guinea , West , North , East , and South Papuan languages); long-range comparison has also suggested connections between selected languages, but again the methodology is not orthodox in historical linguistics. The Great Andamanese languages may be related to some western Papuan languages, but are not themselves covered by

6798-631: The descendants of a very early migration out of Africa around 1.9 mya. A 2011 study found that Denisovan DNA is prevalent in Papuans , Aboriginal Australians , Near Oceanians , Polynesians , Fijians , Eastern Indonesians , and Aeta (from the Philippines); but not in East Asians , western Indonesians, Jahai people (from Malaysia), or Onge (from the Andaman Islands ). This may suggest that Denisovan introgression occurred within

6901-567: The descendants of an earlier migration of H. erectus out of Africa, completely distinct from modern humans and Neanderthals. However, according to the nuclear DNA (nDNA) of Denisova 3—which had an unusual degree of DNA preservation with only low-level contamination—Denisovans and Neanderthals were more closely related to each other than they were to modern humans. Using the percent distance from human–chimpanzee last common ancestor , Denisovans/Neanderthals split from modern humans about 804,000 years ago, and from each other 640,000 years ago. Using

7004-448: The difference between modern humans and Neanderthals is around 202 bases. In comparison, the difference between chimpanzees and modern humans is approximately 1,462 mtDNA base pairs. This suggested that Denisovan mtDNA diverged from that of modern humans and Neanderthals about 1,313,500–779,300 years ago; whereas modern human and Neanderthal mtDNA diverged 618,000–321,200 years ago. Krause and colleagues then concluded that Denisovans were

7107-606: The distribution pattern and divergence of HLA-B*73 from other HLA alleles (involved in the immune system 's natural killer cell receptors ) has led to the suggestion that it introgressed from Denisovans into modern humans in West Asia . In a 2011 study, half of the HLA alleles of modern Eurasians were shown to represent archaic HLA haplotypes, and were inferred to be of Denisovan or Neanderthal origin. A haplotype of EPAS1 in modern Tibetans, which allows them to live at high elevations in

7210-629: The eastern (PNG) highlands. To the west of New Guinea, the largest languages are Makasae in East Timor (100,000 in 2010) and Galela in Halmahera (80,000 reported 1990). To the east, Terei (27,000 reported 2003) and Naasioi (20,000 reported 2007) are spoken on Bougainville. Although there has been relatively little study of these languages compared with the Austronesian family, there have been three preliminary attempts at large-scale genealogical classification, by Joseph Greenberg , Stephen Wurm , and Malcolm Ross . The largest family posited for

7313-410: The ends used to scrape), burins , chisel -like tools, and truncated flakes. These dated to 156±15 thousand years ago in the Main Chamber, 58±6 thousand years ago in the East Chamber, and 136±26–47±8 thousand years ago in the South Chamber. Early Upper Paleolithic artefacts date to 44±5 thousand years ago in the Main Chamber, 63±6 thousand years ago in the East Chamber, and 47±8 thousand years ago in

7416-498: The exception of the language isolate Kuot , which has VSO word order . All other Papuan languages are right-headed . Tonal Papuan languages include the Sko , Lepki , Kaure , Kembra , Lakes Plain , and Keuw languages. Denisovan The Denisovans or Denisova hominins ( / d ə ˈ n iː s ə v ə / də- NEE -sə-və ) are an extinct species or subspecies of archaic human that ranged across Asia during

7519-402: The families that appear when comparing pronouns may be due to pronoun borrowing rather than to genealogical relatedness. However, Ross argues that Papuan languages have closed-class pronoun systems, which are resistant to borrowing, and in any case that the massive number of languages with similar pronouns in a family like Trans–New Guinea preclude borrowing as an explanation. Also, he shows that

7622-664: The femur of a 400,000-year-old H. heidelbergensis from the Sima de los Huesos Cave in Spain was found to be related to those of Neanderthals and Denisovans, but closer to Denisovans, and the authors posited that this mtDNA represents an archaic sequence which was subsequently lost in Neanderthals due to replacement by a modern-human-related sequence. Denisovans are known to have lived in Siberia, Tibet, and Laos. The Xiahe mandible

7725-472: The following language groups. Note that some of these automatically generated groupings are due to chance resemblances. Bill Palmer et al. (2018) propose 43 independent families and 37 language isolates in the Papuasphere, comprising a total of 862 languages. A total of 80 independent groups are recognized. While Pawley & Hammarström 's internal classification of Trans-New Guinea largely resembles

7828-502: The highlands having variants for early brain development and those living in the lowlands having variants for the immune system. Based on the high percentages of Denisovan DNA in modern Papuans and Australians, Denisovans may have crossed the Wallace Line into these regions (with little back-migration west), the second known human species to do so, along with earlier Homo floresiensis . By this logic, they may have also entered

7931-404: The influence of contact and bilingualism . Similarly, several groups that do have substantial basic vocabulary in common with Trans–New Guinea languages are excluded from the phylum because they do not resemble it grammatically. Wurm believed the Papuan languages arrived in several waves of migration with some of the earlier languages (perhaps including the Sepik–Ramu languages ) being related to

8034-500: The introgression event did not take place in Sundaland, or Denisovan ancestry was diluted with gene flow from the mainland Asian Hòabìnhian culture and subsequent Neolithic cultures. In other regions of the world, archaic introgression into humans stems from a group of Neanderthals related to those which inhabited Vindija Cave , Croatia, as opposed to archaics related to Siberian Neanderthals and Denisovans. However, about 3.3% of

8137-429: The jungles of Southeast Asia. The Tam Ngu Hao 2 site might have been a closed forest environment. Little is known of the precise anatomical features of the Denisovans since the only physical remains discovered so far are a finger bone, four teeth, long bone fragments, a partial jawbone, a parietal bone skull fragment, and a rib bone. The finger bone is within the modern human range of variation for women, which

8240-757: The literature. Besides Trans–New Guinea and families possibly belonging in TNG ( see ), he accepted the proposals for, Malcolm Ross re-evaluated Wurm's proposal on purely lexical grounds. That is, he looked at shared vocabulary, and especially shared idiosyncrasies analogous to English I and me vs. German ich and mich . The poor state of documentation of Papuan languages restricts this approach largely to pronouns . Nonetheless, Ross believes that he has been able to validate much of Wurm's classification, albeit with revisions to correct for Wurm's partially typological approach. (See Trans–New Guinea languages .) Ethnologue (2009) largely follows Ross. It has been suggested that

8343-421: The lowest levels of his classification, most of which he inherited from prior taxonomies. Foley (1986) divides Papuan languages into over sixty small language families, plus a number of isolates. However, more recently Foley has accepted the broad outline if not the details of Wurm's classification, as he and Ross have substantiated a large portion of Wurm's Trans–New Guinea phylum. According to Ross (see below),

8446-451: The main problem with Wurm's classification is that he did not take contact-induced change into account. For example, several of the main branches of his Trans–New Guinea phylum have no vocabulary in common with other Trans–New Guinea languages, and were classified as Trans–New Guinea because they are similar grammatically . However, there are also many Austronesian languages that are grammatically similar to Trans–New Guinea languages due to

8549-458: The mandible had a gap behind the molars, and the front teeth were flattened; but Denisovans lacked a high mandibular body, and the mandibular symphysis at the midline of the jaw was more receding. The parietal is reminiscent of that of H. erectus . A facial reconstruction has been generated by comparing methylation at individual genetic loci associated with facial structure. This analysis suggested that Denisovans, much like Neanderthals, had

8652-461: The modern borders of Indonesia and Papua New Guinea. Austronesian-speaking (AN) groups are given in italics . Papuan ethnic groups / tribes in the Indonesian province of West Papua include Arfak, Borai, Doreri, Hatam, Irarutu, Koiwai, Kuri, Madewana, Mairasi, Maniwak, Mbaham, Matta, Meiah, Miere, Meyah, Moire, Moru, Moskona, Napiti, Oburauw, Roon, Roswar, Sebyar, Sougb, Soviar, Sumuri, Wamesa, Warumba, Waruri, Wondama. Papuan ethnic groups / tribes in

8755-666: The more egalitarian New Guinea societies.) Ross has proposed 23 Papuan language families and 9–13 isolates. However, because of his more stringent criteria, he was not able to find enough data to classify all Papuan languages, especially many isolates that have no close relatives to aid in their classification. Ross also found that the Lower Mamberamo languages (or at least the Warembori language—he had insufficient data on Pauwi) are Austronesian languages that have been heavily transformed by contact with Papuan languages, much as

8858-434: The mud, or dug their finger into the toe prints. The footprints average 192.3 mm (7.57 in) long, which roughly equates to a 7 or 8 year old child by modern human growth rates. There are two sets of handprints (from a left and right hand), which may have been created by an older child unless one of the former two individuals had long fingers. The handprints average 161.1 mm (6.34 in), which roughly equates with

8961-625: The non- Austronesian languages spoken on the western Pacific island of New Guinea , as well as neighbouring islands in Indonesia , Solomon Islands , and East Timor . It is a strictly geographical grouping, and does not imply a genetic relationship . New Guinea is the most linguistically diverse region in the world. Besides the Austronesian languages, there arguably are some 800 languages divided into perhaps sixty small language families, with unclear relationships to each other or to any other languages, plus many language isolates . The majority of

9064-507: The nuchal and angular tori near the base of the skull are either reduced or absent, and the back of the skull is rounded off like in early modern humans . Xuchang 1 had a large brain volume of approximately 1800 cc, on the high end for Neanderthals and early modern humans, and well beyond the present-day human average. The Denisovan genome from Denisova Cave has variants of genes which, in modern humans, are associated with dark skin, brown hair, and brown eyes. The Denisovan genome also contains

9167-427: The oldest known examples of rock art . Similar hand stencils and impressions do not appear again in the archeological record until roughly 40,000 years ago. The footprints comprise four right impressions and one left superimposed on one of the rights. They were probably left by two individuals. The tracks of the individual who superimposed their left onto their right may have scrunched up their toes and wiggled them in

9270-526: The population ancestral to both New Guineans and Oceanians. Such a late date for the New Guinean group could indicate Denisovan survival as late as 14,500 years ago, which would make them the latest surviving archaic human species. A third wave appears to have introgressed into East Asia, but there is not enough DNA evidence to pinpoint a solid timeframe. The mtDNA from Denisova 4 bore a high similarity to that of Denisova 3, indicating that they belonged to

9373-685: The region. The timing of introgression into Oceanian populations likely occurred after Eurasians and Oceanians split roughly 58,000 years ago, and before Papuan and Aboriginal Australians split from each other roughly 37,000 years ago. Given the present day distribution of Denisovan DNA, this may have taken place in Wallacea, though the discovery of a 7,200 year old Toalean girl (closely related to Papuans and Aboriginal Australians) from Sulawesi carrying Denisovan DNA makes Sundaland another potential candidate. Other early Sunda hunter gatherers so far sequenced carry very little Denisovan DNA, which either means

9476-469: The same population. 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. Denisova Cave, over time of habitation, continually swung from

9579-461: 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. In 2018, a team of Laotian, French, and American anthropologists, who had been excavating caves in the Laotian jungle of

9682-581: The term Papuan. The most widely used classification of Papuan languages is that of Stephen Wurm , listed below with the approximate number of languages in each family in parentheses. This was the scheme used by Ethnologue prior to Ross's classification (below). It is based on very preliminary work, much of it typological , and Wurm himself has stated that he does not expect it to hold up well to scrutiny. Other linguists, including William A. Foley , have suggested that many of Wurm's phyla are based on areal features and structural similarities, and accept only

9785-490: The time hominins spread out across Europe. H. heidelbergensis is typically considered to have been the direct ancestor of Denisovans and Neanderthals, and sometimes also modern humans. Due to the strong divergence in dental anatomy, they may have split before characteristic Neanderthal dentition evolved about 300,000 years ago. The more divergent Denisovan mtDNA has been interpreted as evidence of admixture between Denisovans and an unknown archaic human population, possibly

9888-559: The two cases of alleged pronoun borrowing in New Guinea are simple coincidence, explainable as regular developments from the protolanguages of the families in question: as earlier forms of the languages are reconstructed, their pronouns become less similar, not more. (Ross argues that open-class pronoun systems, where borrowings are common, are found in hierarchical cultures such as those of Southeast Asia and Japan , where pronouns indicate details of relationship and social status rather than simply being grammatical pro-forms as they are in

9991-800: Was also evidence of stone artefacts in each layer excavated. Some older findings may or may not belong to the Denisovan line, but Asia is not well mapped in regards to human evolution . Such findings include the Dali skull , the Xujiayao hominin, Maba Man , the Jinniushan hominin, and the Narmada Human . The Xiahe mandible shows morphological similarities to some later East Asian fossils such as Penghu 1 , but also to Chinese H. erectus . In 2021, Chinese palaeoanthropologist Qiang Ji suggested his newly erected species, H. longi , may represent

10094-401: Was cut into two, and the initial DNA sequencing of one fragment was later independently confirmed by sequencing the mtDNA from the second. 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

10197-461: Was first suggested and named by Sidney Herbert Ray in 1892. In accordance with William A. Foley (1986): The term 'Papuan languages' must not be taken in the same sense as 'Austronesian languages'. While all Austronesian languages are genetically related in one family, in the sense that they all descend from a common ancestral language called Proto-Austronesian spoken some 6,000 years ago... [Papuan languages] do not all trace their origins back to

10300-468: Was found to be between Highlanders and Lowlanders. Papuan Highlanders fall into three clusters, but form a single clade compared against Lowlanders. The Highlanders underwent a population bottleneck around 10,000 years ago, associated with the adoption of Neolithic lifestyles. Papuan Lowlanders display increased diversity and can be broadly differentiated into a Southern Lowlander cluster and a Northern Lowlander cluster. The genetic differentiation among Papuans

10403-538: Was inconclusive for this specimen, but the team found it anatomically comparable with the Xiahe mandible, and so tentatively categorized it as a Denisovan, although they could not rule out it being Neanderthal. The tooth probably dates to 164,000 to 131,000 years ago. In 2024 a ZooMS analysis of more than 2,500 bones found in Baishiya Karst Cave revealed a further  bone fragment; a rib bone dating from between 48,000 BP and 32,000 BP. The conclusion of

10506-487: Was originally named X-woman because matrilineal mitochondrial DNA (mtDNA) 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 (the oldest specimen) lived 195,000–122,700 years ago. Older Denisovan DNA collected from sediments in

10609-496: Was recorded of them, and few linguists expect that they will ever be linked to another language family . William A. Foley (1986) noted lexical similarities between R. M. W. Dixon 's 1980 reconstruction of proto- Australian and the languages of the East New Guinea Highlands . He believed that it was naïve to expect to find a single Papuan or Australian language family when New Guinea and Australia had been

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