Anatoliy P. Derevianko
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Previous Organizations / Institutes
The organizations / institutes from the scholar's other articles in the past ten years
Research Subjects
Anatoliy P. Derevianko has a total of 768 co-authors. Here are the top co-authors by number of shared publications:
- Maxim B. Kozlikin
Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences
20 shared publications - Michael V. Shunkov
Institute of Archaeology and Ethnography, Siberian Branch, Russian Academy of Sciences
20 shared publications - Janet Kelso
Max Planck Institute for Evolutionary Anthropology, Leipzig, Germany
16 shared publications - Alexander Viktorovich Kandyba
Institute of Archaeology and Ethnography, Siberian Branch of the Russian Academy of Sciences
15 shared publications - Bence Viola
Department of Anthropology, University of Toronto, Toronto, Canada
15 shared publications - Kseniya Anatolievna Kolobova
Institute of Archaeology and Ethnography SB RAS
15 shared publications
Latest Publications
- Open Access
- 4 October 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 54 (3) , 3-27
The Origin of Anatomically Modern Humans ( Homo sapiens ) in Africa
The emergence of anatomically modern humans was a pivotal event in evolution. It has been dealt with in hundreds of books and an infi nite number of articles. I have been tackling this problem for more than half a century, especially in the last 15–20 years, after the results of sequencing Neanderthal and Denisovan DNA had become available. My conclusions will be outlined in two articles. The fi rst one focuses on the evolution of H. erectus in Africa, culminating in the emergence of early Homo sapiens , while in the second one the emergence of H. s. neanderthalensis in Europe and H. s. denisovensis in Central Asia, and the convergent evolution of local H. erectus populations toward H. sapiens in East and Southeast Asia will be examined. Homo sapiens sapiens originated in the process of the dispersal of anatomically modern humans from Africa 60–30 thousand years ago and their admixture with H. s. neanderthalensis in Europe, Denisovans (the informal term is Homo s. denisovensis ) in Central Asia, and early modern humans, informally termed Homo s. orienthalensis , who had originated in China and Southeast Asia.
- 4 October 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 54 (3) , 3-27
Open AccessThe Origin of Anatomically Modern Humans ( Homo sapiens ) in Africa
The emergence of anatomically modern humans was a pivotal event in evolution. It has been dealt with in hundreds of books and an infi nite number of articles. I have been tackling this problem for more than half a century, especially in the last 15–20 years, after the results of sequencing Neanderthal and Denisovan DNA had become available. My conclusions will be outlined in two articles. The fi rst one focuses on the evolution of H. erectus in Africa, culminating in the emergence of early Homo sapiens , while in the second one the emergence of H. s. neanderthalensis in Europe and H. s. denisovensis in Central Asia, and the convergent evolution of local H. erectus populations toward H. sapiens in East and Southeast Asia will be examined. Homo sapiens sapiens originated in the process of the dispersal of anatomically modern humans from Africa 60–30 thousand years ago and their admixture with H. s. neanderthalensis in Europe, Denisovans (the informal term is Homo s. denisovensis ) in Central Asia, and early modern humans, informally termed Homo s. orienthalensis , who had originated in China and Southeast Asia.
- Open Access
A Method to Analyze Low-Quality Archaic Human Genomes and its Application to the Teshik-Tash 1 Neandertal
The Teshik-Tash 1 child whose remains were found in Uzbekistan represents the southeastern-most extent of the known Neandertal range, providing an important link with the better studied Caucasus and Altai Mountain ranges. However, due to poor DNA preservation, studying the genetics of Teshik Tash 1 has remained elusive. Here we present analyses of the nuclear DNA from the Teshik-Tash 1, from extracts that are highly contaminated with present-day human DNA. To achieve this, we developed a new computational method, admixslug , that jointly models contamination and population relationships, in order to infer the relationship of a target individual from which only low-quality nuclear DNA is available, to high-quality archaic human genomes. After validating admixslug , we show that Teshik-Tash 1 is genetically more similar to later Neandertals from Western Eurasia than to older Neandertals from the Altai Mountains. We estimate that Teshik-Tash 1 split from the Western Eurasian lineage between 80,000 and 100,000 years ago. Despite the geographical proximity of Teshik-Tash 1 to the Denisovan range, we find no evidence for Denisovan ancestry in his genome. Our results demonstrate that admixslug enables the study of archaic human specimens in cases where DNA preservation was previously considered too poor for population genetic analyses.
Open AccessA Method to Analyze Low-Quality Archaic Human Genomes and its Application to the Teshik-Tash 1 Neandertal
The Teshik-Tash 1 child whose remains were found in Uzbekistan represents the southeastern-most extent of the known Neandertal range, providing an important link with the better studied Caucasus and Altai Mountain ranges. However, due to poor DNA preservation, studying the genetics of Teshik Tash 1 has remained elusive. Here we present analyses of the nuclear DNA from the Teshik-Tash 1, from extracts that are highly contaminated with present-day human DNA. To achieve this, we developed a new computational method, admixslug , that jointly models contamination and population relationships, in order to infer the relationship of a target individual from which only low-quality nuclear DNA is available, to high-quality archaic human genomes. After validating admixslug , we show that Teshik-Tash 1 is genetically more similar to later Neandertals from Western Eurasia than to older Neandertals from the Altai Mountains. We estimate that Teshik-Tash 1 split from the Western Eurasian lineage between 80,000 and 100,000 years ago. Despite the geographical proximity of Teshik-Tash 1 to the Denisovan range, we find no evidence for Denisovan ancestry in his genome. Our results demonstrate that admixslug enables the study of archaic human specimens in cases where DNA preservation was previously considered too poor for population genetic analyses.
- Open Access
- 23 March 2026
- Published by National Academy of Sciences in Proceedings of the National Academy of Sciences
- Vol. 123 (13)
A high-coverage Neandertal genome from the Altai Mountains reveals population structure among Neandertals
We present a genome sequenced to ~37-fold genomic coverage from an approximately 110,000-y-old male Neandertal from Denisova Cave in the Altai Mountains and analyze it together with previously published Neandertal genomes of high quality. We show that he belonged to a population more closely related to a ~120,000-y-old Neandertal from Denisova Cave than to Neandertals in Europe or to a ~80,000-y-old Neandertal from Chagyrskaya Cave in the Altai Mountains. Both Neandertals from Denisova Cave show evidence of gene flow from Denisovans, a pattern not seen in later Neandertals from the Altai region or from Western Europe. The extent of chromosomal regions of homozygosity in Neandertals from the Altai region between 120,000 and 80,000 y ago indicates that they lived in smaller and more isolated groups than later Neandertals in Europe (54,000 to 40,000 y ago). We estimate the extent of allele frequency differentiation among Neandertal populations and find that the older Eastern Neandertals in the Altai region and younger Western Neandertals in Europe were as differentiated as the most differentiated present-day human populations worldwide.
- 23 March 2026
- Published by National Academy of Sciences in Proceedings of the National Academy of Sciences
- Vol. 123 (13)
Open AccessA high-coverage Neandertal genome from the Altai Mountains reveals population structure among Neandertals
We present a genome sequenced to ~37-fold genomic coverage from an approximately 110,000-y-old male Neandertal from Denisova Cave in the Altai Mountains and analyze it together with previously published Neandertal genomes of high quality. We show that he belonged to a population more closely related to a ~120,000-y-old Neandertal from Denisova Cave than to Neandertals in Europe or to a ~80,000-y-old Neandertal from Chagyrskaya Cave in the Altai Mountains. Both Neandertals from Denisova Cave show evidence of gene flow from Denisovans, a pattern not seen in later Neandertals from the Altai region or from Western Europe. The extent of chromosomal regions of homozygosity in Neandertals from the Altai region between 120,000 and 80,000 y ago indicates that they lived in smaller and more isolated groups than later Neandertals in Europe (54,000 to 40,000 y ago). We estimate the extent of allele frequency differentiation among Neandertal populations and find that the older Eastern Neandertals in the Altai region and younger Western Neandertals in Europe were as differentiated as the most differentiated present-day human populations worldwide.
- Open Access
- 13 January 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 53 (4) , 03-27
The Dispersal of Denisovans and Neanderthals in China
The initial peopling of China by H. erectus occurred ~1.7 Ma BP, maybe even earlier. Despite climatic changes in the Pleistocene, ecological conditions for the permanent occupation of this territory by members of that taxon remained favorable, causing their continuous morphological and genetic evolution in situ along the line leading to H. sapiens . As a result, between 120–40 ka BP, they evolved convergently into early anatomically modern humans in this territory. A large part of the morphological variation of Late Middle to Early Upper Pleistocene hominins in China is attributable not only to the divergence of separate groups of H. erectus descendants, who convergently, under different ecological conditions, evolved into anatomically modern humans using various adaptive strategies, but also to the immigration of Denisovans and their assimilation of native groups. Contacts between the latter and Neanderthals in the second half of the Upper Pleistocene cannot be ruled out too.
- 13 January 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 53 (4) , 03-27
Open AccessThe Dispersal of Denisovans and Neanderthals in China
The initial peopling of China by H. erectus occurred ~1.7 Ma BP, maybe even earlier. Despite climatic changes in the Pleistocene, ecological conditions for the permanent occupation of this territory by members of that taxon remained favorable, causing their continuous morphological and genetic evolution in situ along the line leading to H. sapiens . As a result, between 120–40 ka BP, they evolved convergently into early anatomically modern humans in this territory. A large part of the morphological variation of Late Middle to Early Upper Pleistocene hominins in China is attributable not only to the divergence of separate groups of H. erectus descendants, who convergently, under different ecological conditions, evolved into anatomically modern humans using various adaptive strategies, but also to the immigration of Denisovans and their assimilation of native groups. Contacts between the latter and Neanderthals in the second half of the Upper Pleistocene cannot be ruled out too.
- Open Access
- 13 January 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 53 (4) , 41-51
Biface Trimming Elements as Indicators of Tool Rejuvenation Intensity: The Case of Sibiryachikha, Altai
Biface trimming elements represent an important diagnostic source of information for studying various stages of bifacial tool production. In Middle Paleolithic assemblages, they often constitute a significant portion of the sample, making them particularly useful for identifying specific technological operations such as primary reduction and shaping of bifacial preforms, edge formation and retouching, working edge rejuvenation, and modification of bifacial tools. Here, we present the findings of a systematic typological analysis of biface trimming elements. Through a comprehensive metric and morphological analysis of experimental collections and Middle Paleolithic assemblages representing the Sibiryachikha industries of the Altai, we have developed a refined typology of technical spalls with regard to local archaeological specificity. Our analysis revealed no statistically significant differences either between parameters of type 0 and type 1 spalls or between those of type 2 and type 3, suggesting that these groups correspond to specific technological operations. In contrast, type 4 and type 5 technical spalls, which indicate onsite tool modification, show significant metric differences. The results demonstrate functional and technological distinctions between the assemblages. Materials from Chagyrskaya Cave are especially indicative of primary edge retouching and tool shaping, while the Okladnikov Cave industry mostly evidences maintenance and modification of bifacial tools.
- 13 January 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 53 (4) , 41-51
Open AccessBiface Trimming Elements as Indicators of Tool Rejuvenation Intensity: The Case of Sibiryachikha, Altai
Biface trimming elements represent an important diagnostic source of information for studying various stages of bifacial tool production. In Middle Paleolithic assemblages, they often constitute a significant portion of the sample, making them particularly useful for identifying specific technological operations such as primary reduction and shaping of bifacial preforms, edge formation and retouching, working edge rejuvenation, and modification of bifacial tools. Here, we present the findings of a systematic typological analysis of biface trimming elements. Through a comprehensive metric and morphological analysis of experimental collections and Middle Paleolithic assemblages representing the Sibiryachikha industries of the Altai, we have developed a refined typology of technical spalls with regard to local archaeological specificity. Our analysis revealed no statistically significant differences either between parameters of type 0 and type 1 spalls or between those of type 2 and type 3, suggesting that these groups correspond to specific technological operations. In contrast, type 4 and type 5 technical spalls, which indicate onsite tool modification, show significant metric differences. The results demonstrate functional and technological distinctions between the assemblages. Materials from Chagyrskaya Cave are especially indicative of primary edge retouching and tool shaping, while the Okladnikov Cave industry mostly evidences maintenance and modification of bifacial tools.
- 1 January 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia (Russian-language)
- Vol. 54 (3) , 3-27
The Origin of Anatomically Modern Humans (Homo sapiens) in Africa
Проблема происхождения человека современного типа была и остается одной из сложнейших в науке. Она рассматривалась в книгах и бесчисленных статьях. Меня эта проблема интересует более полувека, особенно последние 15-20 лет, когда были получены данные секвенирования ДНК неандертальцев и денисовцев. Проблема будет рассмотрена в двух статьях. Первая статья посвящена эволюции H. erectus по сапиентной линии в Африке и формированию раннего человека современного типа (Homo sapiens), вторая — проблеме формирования в Европе H. s. neanderthalensis и в Центральной Азии H. s. denisovensis, а также конвергентного формирования в Восточной и Юго-Восточной Азии ранних людей современного типа в процессе сапиентного эволюционного развития H. erectus на этой территории. Формирование Homo sapiens sapiens произошло в процессе расселения из Африки 60-30 тыс. л.н. ранних людей современного типа в Евразии и ассимиляции их с H. s. neanderthalensis в Европе, с денисовцами (неформальное название Homo s. denisovensis) в Центральной Азии и с ранними людьми современного типа (неформальное название Homo s. orienthalensis), которые сформировались в Китае и Юго-Восточной Азии.
- 1 January 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia (Russian-language)
- Vol. 54 (3) , 3-27
The Origin of Anatomically Modern Humans (Homo sapiens) in Africa
Проблема происхождения человека современного типа была и остается одной из сложнейших в науке. Она рассматривалась в книгах и бесчисленных статьях. Меня эта проблема интересует более полувека, особенно последние 15-20 лет, когда были получены данные секвенирования ДНК неандертальцев и денисовцев. Проблема будет рассмотрена в двух статьях. Первая статья посвящена эволюции H. erectus по сапиентной линии в Африке и формированию раннего человека современного типа (Homo sapiens), вторая — проблеме формирования в Европе H. s. neanderthalensis и в Центральной Азии H. s. denisovensis, а также конвергентного формирования в Восточной и Юго-Восточной Азии ранних людей современного типа в процессе сапиентного эволюционного развития H. erectus на этой территории. Формирование Homo sapiens sapiens произошло в процессе расселения из Африки 60-30 тыс. л.н. ранних людей современного типа в Евразии и ассимиляции их с H. s. neanderthalensis в Европе, с денисовцами (неформальное название Homo s. denisovensis) в Центральной Азии и с ранними людьми современного типа (неформальное название Homo s. orienthalensis), которые сформировались в Китае и Юго-Восточной Азии.
- 1 January 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia (Russian-language)
- Vol. 54 (2) , 3-26
Could African Eve Have Admired the Pacific Sunrise?
Одной из фундаментальных задач современной науки является решение проблемы происхождения человека современного типа и его расселения по планете. Результаты антропологических, археологических и генетических исследований позволяют утверждать, что ранний человек современного типа сформировался в Африке не позднее 200-150 тыс. л.н., а может быть, и ранее, если учитывать окаменелости из Джебель Ирхуд в Марокко, древность которых 300 тыс. лет. Покинув Африку, H.sapiens начал расселяться ок. 70-60 тыс. л.н. в Евразии. Процесс расселения этого таксона сопровождался, по мнению одних исследователей, замещением коренного населения, по мнению других — частичной гибридизацией. В третьем номере журнала за 2025 г. опубликована статья, в которой рассматривался вопрос о формировании человека современного типа в Китае. Данная работа является ее продолжением. В ней обсуждается проблема возможного формирования на предковой основе H. erectus H. sapiens на сопредельной с Китаем территории — в Юго-Восточной Азии.
- 1 January 2026
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia (Russian-language)
- Vol. 54 (2) , 3-26
Could African Eve Have Admired the Pacific Sunrise?
Одной из фундаментальных задач современной науки является решение проблемы происхождения человека современного типа и его расселения по планете. Результаты антропологических, археологических и генетических исследований позволяют утверждать, что ранний человек современного типа сформировался в Африке не позднее 200-150 тыс. л.н., а может быть, и ранее, если учитывать окаменелости из Джебель Ирхуд в Марокко, древность которых 300 тыс. лет. Покинув Африку, H.sapiens начал расселяться ок. 70-60 тыс. л.н. в Евразии. Процесс расселения этого таксона сопровождался, по мнению одних исследователей, замещением коренного населения, по мнению других — частичной гибридизацией. В третьем номере журнала за 2025 г. опубликована статья, в которой рассматривался вопрос о формировании человека современного типа в Китае. Данная работа является ее продолжением. В ней обсуждается проблема возможного формирования на предковой основе H. erectus H. sapiens на сопредельной с Китаем территории — в Юго-Восточной Азии.
- Open Access
- 9 October 2025
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 53 (3) , 107-130
The Origin of Anatomically Modern Humans in China
Genetic, skeletal, and archaeological studies alike demonstrate that anatomically modern humans originated ~200–150 ka BP in Africa, and this finding cannot be questioned. Over the last 4–5 decades, however, human fossils with undoubtedly modern skeletal markers, dating to 120–40 ka BP, have been discovered in China and Southeast Asia, suggesting that those territories, too, may have been part of the region where H. s. sapiens arose convergently on the basis of H. erectus. An important fact supporting this hypothesis is that a new taxon, Denisovan (informally H. s. denisovan), was established on the basis of the genetic analysis of a bone sample from a phalanx of the fifth finger of a 9–12-year-old girl from Denisova Cave. Also, genetic studies show that all three taxa, African anatomically modern humans, European Neanderthals, and Central Asian Denisovans, had interbred and produced fertile offspring, implying that all these taxa formed an open genetic system with processes such as admixture and assimilation, suggestive of a single species. Based on this conclusion, the following scenario of the origin of H. sapiens sapiens can be proposed: 80–60 ka BP anatomically modern migrants engaged in such processes with H. s. neanderthalensis in Europe, Denisovans in Central Asia, and early anatomically modern descendants of H. erectus (informally H. s. orientalensis) in East and Southeast Asia. The key role in the origin of H. s. sapiens, then, was played by migrants from Africa (informally H. s. africanensis), who were engaged in interbreeding and assimilation with H. s. neanderthalensis, H. s. orientalensis, and H. s. denisovan.
- 9 October 2025
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 53 (3) , 107-130
Open AccessThe Origin of Anatomically Modern Humans in China
Genetic, skeletal, and archaeological studies alike demonstrate that anatomically modern humans originated ~200–150 ka BP in Africa, and this finding cannot be questioned. Over the last 4–5 decades, however, human fossils with undoubtedly modern skeletal markers, dating to 120–40 ka BP, have been discovered in China and Southeast Asia, suggesting that those territories, too, may have been part of the region where H. s. sapiens arose convergently on the basis of H. erectus. An important fact supporting this hypothesis is that a new taxon, Denisovan (informally H. s. denisovan), was established on the basis of the genetic analysis of a bone sample from a phalanx of the fifth finger of a 9–12-year-old girl from Denisova Cave. Also, genetic studies show that all three taxa, African anatomically modern humans, European Neanderthals, and Central Asian Denisovans, had interbred and produced fertile offspring, implying that all these taxa formed an open genetic system with processes such as admixture and assimilation, suggestive of a single species. Based on this conclusion, the following scenario of the origin of H. sapiens sapiens can be proposed: 80–60 ka BP anatomically modern migrants engaged in such processes with H. s. neanderthalensis in Europe, Denisovans in Central Asia, and early anatomically modern descendants of H. erectus (informally H. s. orientalensis) in East and Southeast Asia. The key role in the origin of H. s. sapiens, then, was played by migrants from Africa (informally H. s. africanensis), who were engaged in interbreeding and assimilation with H. s. neanderthalensis, H. s. orientalensis, and H. s. denisovan.
- Open Access
- 15 September 2025
- Published by Mongolian Journals Online in Studia Archaeologica
- p. 6-22
The significance of Tsagaan Agui in Mongolian Paleolithic Archaeology
Tsagaan Agui (White Cave; Цагаан Агуй), located in the Gobi Altai Mountains of southern Mongolia, represents one of the few stratified and well-dated Pleistocene archaeological sites now known in the Gobi Desert. Archaeological studies undertaken at Tsagaan Agui since 1995 have revealed that the cave’s sediments contain cultural remains ranging from the Middle and, possibly, Lower Paleolithic, to the later historic period. Analyses of these deposits suggest environmental conditions favorable for intermittent human occupation existed throughout most of the Pleistocene and early to middle Holocene, the latter likely during periods of larger-scale climatic and environmental degradation.
- 15 September 2025
- Published by Mongolian Journals Online in Studia Archaeologica
- p. 6-22
Open AccessThe significance of Tsagaan Agui in Mongolian Paleolithic Archaeology
Tsagaan Agui (White Cave; Цагаан Агуй), located in the Gobi Altai Mountains of southern Mongolia, represents one of the few stratified and well-dated Pleistocene archaeological sites now known in the Gobi Desert. Archaeological studies undertaken at Tsagaan Agui since 1995 have revealed that the cave’s sediments contain cultural remains ranging from the Middle and, possibly, Lower Paleolithic, to the later historic period. Analyses of these deposits suggest environmental conditions favorable for intermittent human occupation existed throughout most of the Pleistocene and early to middle Holocene, the latter likely during periods of larger-scale climatic and environmental degradation.
- Open Access
- 4 July 2025
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 53 (2) , 3-24
The Peopling of Mongolia by Homo sapiens denisovan
The article continues a series of publications in this journal, addressing the peopling of Iran, Tajikistan, and Uzbekistan by H. s. denisovan. Around 400 ka BP, two large populations branched off from the ancestral taxon H. heidelbergensis in the Levant. One of them migrated to Europe, where its hybridization with natives gave rise to a new taxon H. s. neanderthalensis 200–150 ka BP. The other, having passed through the Iranian Highland, settled in Central Asia, where its mixture with local Central Asiatic H. erectus, natural and sexual selection, and adaptation to various environments in the vast Central Asiatic region 200–150 ka BP resulted in the emergence of a morphologically and genetically distinct taxon, whose informal name is H. s. denisovan.
- 4 July 2025
- Published by Institute of Archaeology and Ethnography SB RAS in Archaeology, Ethnology and Anthropology of Eurasia
- Vol. 53 (2) , 3-24
Open AccessThe Peopling of Mongolia by Homo sapiens denisovan
The article continues a series of publications in this journal, addressing the peopling of Iran, Tajikistan, and Uzbekistan by H. s. denisovan. Around 400 ka BP, two large populations branched off from the ancestral taxon H. heidelbergensis in the Levant. One of them migrated to Europe, where its hybridization with natives gave rise to a new taxon H. s. neanderthalensis 200–150 ka BP. The other, having passed through the Iranian Highland, settled in Central Asia, where its mixture with local Central Asiatic H. erectus, natural and sexual selection, and adaptation to various environments in the vast Central Asiatic region 200–150 ka BP resulted in the emergence of a morphologically and genetically distinct taxon, whose informal name is H. s. denisovan.