Multiple Deeply Divergent Denisovan Ancestries in Papuans
Open accessAbstract
Genome sequences are known for two archaic hominins-Neanderthals and Denisovans-which interbred with anatomically modern humans as they dispersed out of Africa. We identified high-confidence archaic haplotypes in 161 new genomes spanning 14 island groups in Island Southeast Asia and New Guinea and found large stretches of DNA that are inconsistent with a single introgressing Denisovan origin. Instead, modern Papuans carry hundreds of gene variants from two deeply divergent Denisovan lineages that separated over 350 thousand years ago. Spatial and temporal structure among these lineages suggest that introgression from one of these Denisovan groups predominantly took place east of the Wallace line and continued until near the end of the Pleistocene. A third Denisovan lineage occurs in modern East Asians. This regional mosaic suggests considerable complexity in archaic contact, with modern humans interbreeding with multiple Denisovan groups that were geographically isolated from each other over deep evolutionary time.
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Recent studies have shown the existence of genetic structure characterising the Indonesian archipelago, with Papuan-and thus, Denisovan-ancestry showing a marked west to east cline.
“Recent studies have shown the existence of genetic structure characterising the Indonesian archipelago, with Papuan—and thus, Denisovan—ancestry showing a marked west to east cline [6, 53].”
From Denisovan introgression has shaped the immune system of present-day Papuans (2022) · cites this paper as [6]
Denisovan-like ancestry is the result of multiple introgression events from different source populations.
“We find support for the hypothesis that Denisovan-like ancestry is the result of multiple introgression events from different source populations (9, 10).”
From An ancestral recombination graph of human, Neanderthal, and Denisovan genomes (2021) · cites this paper as [10]
Prior studies have suggested that Denisovan-like haplotype blocks in humans have two or three distinct sources with different levels of divergence to the Denisovan genome, with the best-matching haplotype blocks in East Asia.
“Prior studies have suggested that Denisovan-like haplotype blocks in humans have two or three distinct sources with different levels of divergence to the Denisovan genome, with the best-matching haplotype blocks in East Asia (9, 10).”
From An ancestral recombination graph of human, Neanderthal, and Denisovan genomes (2021) · cites this paper as [10]
Denisovan components in Papuans exhibit deep divergence from the reference Altai Denisovan genome, providing evidence for multiple Denisovan introgression events across time and space.
“these components exhibit a deep divergence from the reference Altai Denisovan genome, providing strong evidence for the occurrence of multiple Denisovan introgression events across time and space [6, 7].”
From Denisovan introgression has shaped the immune system of present-day Papuans (2022) · cites this paper as [6]
The filtering criteria ensure that only variants with a high likelihood of being truly introgressed are considered.
“While our analyses focus on a small subset of introgressed variants identified by [6], our filtering criteria ensure that we only consider variants with a high likelihood of being truly introgressed.”
From Denisovan introgression has shaped the immune system of present-day Papuans (2022) · cites this paper as [6]
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Zilhão, J., d'Errico, F., Banks, W., & Teyssandier, N. (2024) · Quaternary Environments and Humans · doi:10.1016/j.qeh.2024.100037 · Cited by 28 · see citing works
Rong, S., Neil, C. R., Welch, A., Duan, C., Maguire, S., Meremikwu, I. C., Meyerson, M., Evans, B. J., & Fairbrother, W. G. (2023) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2218308120 · Cited by 24 · see citing works
Langdon, Q. K., Groh, J. S., Aguillon, S. M., Powell, D. L., Gunn, T. R., Payne, C. Y., Baczenas, J. J., Donny, A. E., Dodge, T. O., Du, K., et al. (2024) · PLoS Biology · doi:10.1371/journal.pbio.3002742 · Cited by 24 · see citing works
Predicting Archaic Hominin Phenotypes from Genomic Data
Brand, C. M., Colbran, L. L., & Capra, J. A. (2022) · Annual Review of Genomics and Human Genetics · doi:10.1146/annurev-genom-111521-121903 · Cited by 23 · see citing works
Apportioning archaic variants among modern populations
Witt, K. E., Villanea, F. A., Loughran, E., Zhang, X., & Huerta‐Sánchez, E. (2022) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2020.0411 · Cited by 24 · see citing works
Positive selection in the genomes of two Papua New Guinean populations at distinct altitude levels
André, M., Brucato, N., Hudjashov, G., Pankratov, V., Yermakovich, D., Montinaro, F., Kreevan, R., Kariwiga, J., Muke, J., Boland, A., et al. (2024) · Nature Communications · doi:10.1038/s41467-024-47735-1 · Cited by 25 · see citing works
Self-citationBlinkhorn, J., Achyuthan, H., Durcan, J. A., Roberts, P., & Ilgner, J. (2021) · Scientific Reports · doi:10.1038/s41598-021-98897-7 · Cited by 21 · see citing works
Tracing the Evolution of Human Immunity Through Ancient DNA
Patin, É., & Quintana‐Murci, L. (2024) · Annual Review of Immunology · doi:10.1146/annurev-immunol-082323-024638 · Cited by 20 · see citing works
The MUC19 gene: An evolutionary history of recurrent introgression and natural selection
Villanea, F. A., Peede, D., Kaufman, E. J., Añorve-Garibay, V., Chevy, E. T., Villa-Islas, V., Witt, K. E., Zeloni, R., Marnetto, D., Moorjani, P., et al. (2025) · Science · doi:10.1126/science.adl0882 · Cited by 21 · see citing works
Flemming, N. C. (2020) · The Journal of Island and Coastal Archaeology · doi:10.1080/15564894.2020.1781712 · Cited by 20 · see citing works
Archaic mitochondrial DNA inserts in modern day nuclear genomes
Bücking, R., Cox, M. P., Hudjashov, G., Saag, L., Sudoyo, H., & Stoneking, M. (2019) · BMC Genomics · doi:10.1186/s12864-019-6392-8 · Cited by 19 · see citing works
Self-citationReconstruction of the Austronesian Diaspora in the Era of Genomics
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Kaifu, Y., Kurniawan, I., Yurnaldi, D., Setiawan, R., Setiyabudi, E., Insani, H., Takai, M., Nishioka, Y., Takahashi, A., Aziz, F., et al. (2022) · Journal of Human Evolution · doi:10.1016/j.jhevol.2021.103122 · Cited by 19 · see citing works
Rogers, N. M., Zammit, N. W., Nguyen-Ngo, D., Souilmi, Y., Minhas, N., Meijles, D. N., Self, E., Walters, S. N., Warren, J. A., Cultrone, D., et al. (2023) · Kidney International · doi:10.1016/j.kint.2023.02.030 · Cited by 19 · see citing works
sstar: A Python Package for Detecting Archaic Introgression from Population Genetic Data with S *
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East and Southeast Asian hominin dispersal and evolution: A review
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Skeletal remains of a Pleistocene modern human (Homo sapiens) from Sulawesi
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Archaic Introgression Shaped Human Circadian Traits
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“The Eurasian environments where Neanderthals and Denisovans lived for several hundred thousand years are located at higher latitudes with more variable photoperiods than the landscape where AMH evolved before leaving Africa.”
BackgroundInfluentialIntrogressive Hybridization and Hypoxia Adaptation in High-Altitude Vertebrates
Storz, J. F., & Signore, A. V. (2021) · Frontiers in Genetics · doi:10.3389/fgene.2021.696484 · Cited by 15 · see citing works
An Extended Admixture Pulse Model Reveals the Limitations to Human–Neandertal Introgression Dating
Iasi, L. N. M., Ringbauer, H., & Peter, B. M. (2021) · Molecular Biology and Evolution · doi:10.1093/molbev/msab210 · Cited by 15 · see citing works
Genomic evidence supports the “long chronology” for the peopling of Sahul
Gandini, F., Almeida, M., Foody, M. G. B., Nagle, N., Bergström, A., Olivieri, A., Rodrigues, S., Fichera, A., Oteo‐García, G., Torroni, A., et al. (2025) · Science Advances · doi:10.1126/sciadv.ady9493 · Cited by 13 · see citing works
- Authors
- Jacobs, G. S., Hudjashov, G., Saag, L., Kusuma, P., Darusallam, C. C., Lawson, D. J., Mondal, M., Pagani, L., Ricaut, F.-X., Stoneking, M., Metspalu, M., Sudoyo, H., Lansing, J. S., & Cox, M. P.
- Affiliations
- Nanyang Technological University (SG)
- Massey University (NZ)
- University of Tartu (EE)
- Eijkman Institute for Molecular Biology (ID)
- University of Bristol (GB)
- University of Padua (IT)
- Centre National de la Recherche Scientifique (FR)
- Max Planck Institute for Evolutionary Anthropology (DE)
- The University of Sydney (AU)
- Santa Fe Institute (US)
- Corresponding author
- Hudjashov, Georgi
- Cox, Murray P.
- ORCID
- Venue
- Cell
- Publisher
- Elsevier BV
- Published
- 2019-05
- Volume
- 177
- Issue
- 4
- Pages
- 1010-1021.e32
- ISSN
- 0092-8674 (ISSN-L)
- OpenAlex
- W2923745223
- Cited by
- 346
- References
- 95
- Citations by year
- 2026: 27 · 2025: 31 · 2024: 104 · 2023: 22 · 2022: 37 · 2021: 63 · 2020: 43 · 2019: 18 · 2014: 1
- FWCI
- 185.42
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- Cell (publishedVersion)
- PubMed (publishedVersion)
- Explore Bristol Research (acceptedVersion, cc-by-nc-nd)
- HAL (submittedVersion)
- MPG.PuRe (Max Planck Society) (submittedVersion)
- Bristol Research (University of Bristol) (submittedVersion, cc-by-nc-nd)
- Apollo (University of Cambridge) (submittedVersion, cc-by-nc-nd)
- Padua Research Archive (University of Padova) (submittedVersion, other-oa)
- Apollo
- Funders
- National Science Foundation (SES 0725470)
- Singapore Ministry of Education Tier II (MOE2015-T2-1-127)
- NTU Presidential Postdoctoral Fellowship
- Complexity Institute Individual Fellowship
- French ANR grant (ANR-14-CE31-0013-01)
- European Union grant through the European Regional Development Fund (2014-2020.4.01.15-0012)
- Royal Society of New Zealand Marsden (17-MAU-040)
- German Alexander von Humboldt Foundation fellowship
- Microsoft research grant for Azure cloud computing
- High Performance Computing Center, University of Tartu, Estonia
- Topics
- Social Sciences › Anthropology › Pleistocene-Era Hominins and Archaeology
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Forensic and Genetic Research
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Genetic diversity and population structure
- Keywords
- Denisovans, Papuans, archaic introgression, Wallace's Line, Island Southeast Asia, New Guinea, Pleistocene, human dispersal out of Africa
- MeSH terms
- Genetic Introgression, Animals, Biological Evolution, Haplotypes, Humans, Indonesia, Genetic Variation, Hominidae, Genome, Human, Oceania, Asian People, Gene Flow, Neanderthals
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
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