Denisovan

Denisovans or Denisova hominins are an extinct hominid species or subspecies closely related to the Neanderthals and identified from the DNA sequence of the about 40,000 year old fossils found in Siberia. The fossils unearthed from Denisova Cave in the Altai Mountains of southern Siberia. were quite limited ni number: However, the entire genome has been sequenced and this DNA sequence shows

extinct hominin species or homo sapien subspecies

believed to have separated from the neanderhtals and one of three migrations ut f Africa since that time: **8 and about 60,000 yearsago homo sapiens.

(Homo sapiens ssp. 'Denisova) Neanderthals (Homo sapiens neanderthalensis or Homo neanderthalensis)

cousins of Neanderthals hominins known from a (fossilized pinkie bone) finger bone and two teeth but also becasue the entire genome has been sequenced about 40,000 years ago discoveredin Siberia may have "ranged widely across Asia" nuclear DAN from pinkie bone Siberian cave relatedt o Neanderhals, but "distinct enought" national geographic "merit clssification as a new species of archaic humans" names Deniosvan after ave where pinkie bone found

genetic overlap with some present-day east Asians and partricular the melanesians living in papua New Guinea, where about 3 to 5 percent overlap of geneticmateiral like interbred with modern human ancestors of Melanesians

also genome of Den shows snippes that seem to be from another group, either a previously unknown species or from teh many species that "are known only fromfossils" new scientists never genetically analyzed maybe inbreed iwth h.hedelbergenesis — (600,000 to 250,000 years ago, and spread from Africa into Europe and Western Asia) or maybe H. erectus (also widespread)... eno and neaderthals in cold climiate sogeneomes preserved, those in hot, wet regions, the DNA breaks down — new scientists

our ancestors inbreed with boht neaderthals nd denisovans

Paleolithic-era members of a species of Homo or subspecies of Homo sapiens. In March 2010, scientists announced the discovery of a finger bone fragment of a juvenile female who lived about 41,000 years ago, found in the remote Denisova Cave in the Altai Mountains in Siberia, a cave which has also been inhabited by Neanderthals and modern humans. Two teeth and a toe bone belonging to different members of the same population have since been reported.

in addition to Homo erectus

Neanderthals, Denisovans, and Homo sapiens all believedto have migrated out of aftrica, homo spaiens about 60,000 years ago, neanderhals about 300,000 years ago (settlining in Euorpe and western Asia) and Denisovans

National geographic: One theory, Nean. den, modernhumans all desended form ancient human Homo heidelbergenesis. onebranch left Africa 300,000 to 400,000 years ago and splie wiht one branch becma eneanderhals (West Asia and Europe) and one Denisovans (more east0 130,000 years ago H. heidel... became Homo sapiens and exodus 60,000 years ago.

branched off about 400,000 years ago and then split into neand and denosovians

snippets different, meaning likely inbreed fwith uniidentified species unrelated to Neanderhals

Notes: http://www.sciencemag.org/content/342/6163/1156.summary https://genographic.nationalgeographic.com/denisovan/ http://www.newscientist.com/article/dn24603-mystery-human-species-emerges-from-denisovan-genome.html#.UqyGK-JFP6o http://www.usatoday.com/story/news/nation/2013/10/17/skull-sheds-light-human-species/2994613/ http://www.nytimes.com/2013/10/18/science/fossil-skull-may-rewrite-humans-evolutionary-story.html?_r=0 http://www.bbc.co.uk/news/science-environment-19184370 http://www.theguardian.com/science/2013/oct/17/skull-homo-erectus-human-evolution http://online.wsj.com/news/articles/SB10001424052702304384104579141600675336982

Analysis of the mitochondrial DNA (mtDNA) of the finger bone showed it to be genetically distinct from the mtDNAs of Neanderthals and modern humans. Subsequent study of the nuclear genome from this specimen suggests that this group shares a common origin with Neanderthals, that they ranged from Siberia to Southeast Asia, and that they lived among and interbred with the ancestors of some present-day modern humans, with up to 6% of the DNA of Melanesians and Australian Aborigines deriving from Denisovans. A comparison with the genome of a Neanderthal from the same cave revealed significant local interbreeding, with local Neanderthal DNA representing 17% of the Denisovan genome, while evidence was also detected of interbreeding with an as yet unidentified ancient human lineage. Similar analysis of a toe bone discovered in 2011 is underway, while analysis of DNA from two teeth found in different layers than the finger bone revealed an unexpected degree of mtDNA divergence among Denisovans. In 2013, mitochondrial DNA from a 400,000 year old hominin femur bone from Spain, which had been seen as either Neanderthal or Homo heidelbergensis, was found to be closer to Denisovan mtDNA than to Neanderthal mtDNA.

Discovery

Tourists in front of the Denisova Cave, where "X woman" was found

In 2008, Russian archaeologists from the Institute of Archaeology and Ethnology of Novosibirsk, working at the site of Denisova Cave in the Altai Mountains of Siberia, uncovered a small bone fragment from the fifth finger of a juvenile hominin, dubbed the "X woman" (referring to the maternal descent of mitochondrial DNA,) or the Denisova hominin. Artifacts, including a bracelet, excavated in the cave at the same level were carbon dated to around 40,000 BP.

A team of scientists led by Johannes Krause and Swedish biologist Svante Pääbo from the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, sequenced mtDNA extracted from the fragment. Because of the cool climate in the location of the Denisova Cave, the discovery benefited from DNA's ability to survive for longer periods at lower temperatures. The average annual temperature of the cave remains at 0°C, which has contributed to the preservation of archaic DNA among the remains discovered. The analysis indicated that modern humans, Neanderthals, and the Denisova hominin last shared a common ancestor around 1 million years ago.

The mtDNA analysis further suggested this new hominin species was the result of an early migration out of Africa, distinct from the later out-of-Africa migrations associated with Neanderthals and modern humans, but also distinct from the earlier African exodus of Homo erectus. Pääbo noted the existence of this distant branch creates a much more complex picture of humankind during the Late Pleistocene.

Later in 2010, a second paper from the Svante Pääbo group reported the prior discovery, in 2000, of a third upper molar from a young adult, dating from about the same time (the finger was from level 11 in the cave sequence, the tooth from level 11.1). The tooth differed in several aspects from those of Neanderthals, while having archaic characteristics similar to the teeth of Homo erectus. They again performed mitochondrial DNA analysis on the tooth and found it to have a different but similar sequence to that of the finger bone, indicating a divergence time about 7,500 years before, and suggesting it belonged to a different individual from the same population.

In 2011, a toe bone was discovered in layer 11 of the cave, and hence was contemporary with the finger bone. Preliminary characterization of the bone's mitochondrial DNA suggests it belonged to a Neanderthal, not a Denisovan. The Altai cave also contains stone tools and bone artifacts made by modern humans, and Pääbo commented: "The one place where we are sure all three human forms have lived at one time or another is here in Denisova Cave."

Anatomy

Little is known of the precise anatomical features of the Denisovans since the only physical remains discovered thus far are the finger bone and two teeth from which genetic material has been gathered, and a toe bone. The single finger bone is unusually broad and robust, well outside the variation seen in modern people. Surprisingly, it belonged to a female, indicating the Denisovans were extremely robust, perhaps similar in build to the Neanderthals. The tooth that has been characterized shares no derived morphological features with Neanderthal or modern humans. An initial morphological characterization of the toe bone led to the suggestion that it may have belonged to a Neanderthal-Denisovan hybrid individual, although a critic suggested the morphology was inconclusive. This toe bone is currently undergoing DNA analysis by Pääbo.

Some older finds may or may not belong to the Denisovan line. These includes the skulls from Dali and Maba, and a number of more fragmentary remains from Asia. Asia is not well mapped with regard to human evolution, and the above finds may may represent a group of "Asian Neanderthals".

Mitochondrial DNA analysis

The mtDNA from the finger bone differs from that of modern humans by 385 bases (nucleotides) in the mtDNA strand out of approximately 16,500, whereas the difference between modern humans and Neanderthals is around 202 bases. In contrast, the difference between chimpanzees and modern humans is approximately 1,462 mtDNA base pairs. This suggested a divergence time around one million years ago. The mtDNA from a tooth bore a high similarity to that of the finger bone, indicating they belonged to the same population. From a second tooth, an mtDNA sequence was recovered that showed an unexpectedly large number of genetic differences compared to that found in the other tooth and the finger, suggesting a high degree of mtDNA diversity. These two individuals from the same cave showed more diversity than seen among sampled Neanderthals from all of Eurasia, and were as different as humans from different continents.

Nuclear genome analysis

In the same second 2010 paper, the authors reported the isolation and sequencing of nuclear DNA from the Denisova finger bone. This specimen showed an unusual degree of DNA preservation and low level of contamination. They were able to achieve near-complete genomic sequencing, allowing a detailed comparison with Neanderthal and modern humans. From this analysis, they concluded, in spite of the apparent divergence of their mitochondrial sequence, the Denisova population along with Neanderthal shared a common branch from the lineage leading to modern African humans. The estimated average time of divergence between Denisovan and Neanderthal sequences is 640,000 years ago, and the time between both of these and the sequences of modern Africans is 804,000 years ago. They suggest the divergence of the Denisova mtDNA results either from the persistence of a lineage purged from the other branches of humanity through genetic drift or else an introgression from an older hominin lineage. In 2013, the mtDNA sequence from the femur of a 400,000 year old Homo heidelbergensis from the Sima de los Huesos Cave in Spain was found to be most similar to that of Denisova.

Interbreeding

A detailed comparison of the Denisovan, Neanderthal, and human genomes has revealed evidence for a complex web of interbreeding among the lineages. Through such interbreeding, 17% of the Denisova genome represents DNA from the local Neanderthal population, while evidence was also found of a contribution to the nuclear genome from an ancient hominin lineage yet to be identified, perhaps the source of the anomalously ancient mtDNA.

Analysis of genomes of modern humans show that they mated with at least two groups of ancient humans: Neanderthals (more similar to those found in the Caucasus than those from the Altai region) and Denisovans. Approximately 4% of the DNA of non-African modern humans is shared with Neanderthals, suggesting interbreeding. Tests comparing the Denisova hominin genome with those of six modern humans – a ǃKung from South Africa, a Nigerian, a Frenchman, a Papua New Guinean, a Bougainville Islander and a Han Chinese – showed that between 4% and 6% of the genome of Melanesians (represented by the Papua New Guinean and Bougainville Islander) derives from a Denisovan population. This DNA was possibly introduced during the early migration to Melanesia. These findings are in concordance with the results of other comparison tests which show a relative increase in allele sharing between the Denisovan and the Aboriginal Australian genome, compared to other Eurasians and African populations, however it has been observed that Papuans, the population of Papua New Guinea, have more allele sharing than Aboriginal Australians.

Melanesians may not be the only modern-day descendants of Denisovans. David Reich of Harvard University, in collaboration with Mark Stoneking of the Planck Institute team, found genetic evidence that Denisovan ancestry is shared by Melanesians, Australian Aborigines, and smaller scattered groups of people in Southeast Asia, such as the Mamanwa, a Negrito people in the Philippines. However, not all Negritos were found to possess Denisovan genes; Onge Andaman Islanders and Malaysian Jehai, for example, were found to have no significant Denisovan inheritance. These data place the interbreeding event in mainland Southeast Asia, and suggest that Denisovans once ranged widely over eastern Asia. Based on the modern distribution of Denisova DNA, Denisovans may have crossed the Wallace Line, with Wallacea serving as their last refugium.

The immune system's HLA alleles have drawn particular attention in the attempt to identify genes that may derive from archaic human populations. Although not present in the sequenced Denisova genome, the distribution pattern and divergence of HLA-B*73 from other HLA alleles has led to the suggestion that it introgressed from Denisovans into humans in west Asia. Indeed, half of the HLA alleles of modern Eurasians represent archaic HLA haplotypes, and have been inferred to be of Denisovan or Neanderthal origin. The apparent over-representation of these alleles suggests a positive selective pressure for their retention in the human population.

Further reading

• Barras, Colin (13 August 2011), Stone Age toe could redraw human family tree .
• Fox, Maggie (24 March 2010), Possible new human ancestor found in Siberia .
• Ghosh, Pallab (22 December 2010), Ancient humans, dubbed 'Denisovans', interbred with us, BBC News online 
• Hawks, John (22 December 2010), The Denisova genome FAQ .
• Rincon, Paul (25 March 2010), DNA identifies new ancient human dubbed 'X-woman', BBC News online .
• Shreeve, Jamie (July 2013), "The Case of the Missing Ancestor", National Geographic .
• Wade, Nicholas (24 March 2010), "Bone May Reveal a New Human Group", The New York Times .
• Zimmer, Carl (24 March 2010), Hybrid speculation (blog), Discover .
• Zimmer, Carl (22 December 2010), Meet the Denisovans, the newest members of the human tree of life (blog), Discover .

External links

• The Denisova Consortium's raw sequence data and alignments

Part of the series on Human evolution

Hominini

Sahelanthropus tchadensis • Orrorin tugenensis • Ardipithecus

Australopithecines

Australopithecus: A. anamensis • A. afarensis • A. bahrelghazali • A. africanus • A. garhi

Paranthropus: P. aethiopicus • P. boisei • P. robustus

Humans and Proto-humans

Kenyanthropus platyops

Homo: H. habilis • H. rudolfensis • H. georgicus • H. ergaster • H. erectus (H. e. lantianensis • H. e. palaeojavanicus • H. e. pekinensis • H. e. soloensis) • H. cepranensis • H. antecessor • H. heidelbergensis • H. neanderthalensis • H. rhodesiensis • H. floresiensis • Homo sapiens (H. s. idaltu • H. s. sapiens)

Topics: Timeline of human evolution • List of human fossils • Human evolutionary genetics

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