Palaeogenomics and the Palaeolithic of Southwest Asia: Trends, Issues, and Future Directions


  • David L. G. Miedzianogora University College London




Palaeogenomics is the study of ancient subfossilised remains on a genome-wide level, and it has revolutionised the study and understanding of the deep past. This is also the case in Southwest Asia, where especially the Bronze and Iron Ages have seen substantial research. However, due to the poor preservation of DNA in the region, the Palaeolithic remains largely understudied despite the possibility of novel interpretations of this key period. Here, I review several ways that palaeogenomics has begun changing our understanding of the Palaeolithic of the region in three key areas: the dispersal of modern humans out of Africa, the interactions between Neanderthals and modern humans, and the formation of Southwest Asian population structures. Most of these interpretations are based on data from outside of Southwest Asia, and I argue that a closer integration between palaeogenomics, archaeology, and local stakeholders are necessary to begin solving the issues surrounding the poor preservation of DNA in the region. If this can be done, palaeogenomics holds many possibilities for future Palaeolithic research. 


Allentoft, Morten E. et al. 2012. “The Half-Life of DNA in Bone: Measuring Decay Kinetics in 158 Dated Fossils.” Proceedings. Biological Sciences 279 (1748): 4724–33. DOI: 10.1098/rspb.2012.1745.

Alpaslan-Roodenberg, Songül et al. 2021. “Ethics of DNA Research on Human Remains: Five Globally Applicable Guidelines.” Nature 599 (7883): 41–46. DOI: 10.1038/s41586-021-04008-x.

Ávila-Arcos, Maria C. et al. 2022. “Recommendations for Sustainable Ancient DNA Research in the Global South: Voices From a New Generation of Paleogenomicists.” Frontiers in Genetics 13.

Ávila-Arcos, Maria C., Raghavan, Maanasa, and Schlebusch, Carina. 2023. “Going Local with Ancient DNA: A Review of Human Histories from Regional Perspectives.” Science 382 (6666): 53-58. DOI: 10.1126/science.adh8140.

Bar-Yosef, Ofer. 2013. “Neanderthals and Modern Humans Across Eurasia.” In Dynamics of Learning in Neanderthals and Modern Humans Volume 1: Cultural Perspectives, edited by Takeru Akazawa et al., 7–20. Replacement of Neanderthals by Modern Humans Series. Tokyo: Springer Japan. DOI: 10.1007/978-4-431-54511-8_2.

Bergström, Anders et al. 2020. “Insights into Human Genetic Variation and Population History from 929 Diverse Genomes.” Science 367 (6484): eaay5012. DOI: 10.1126/science.aay5012.

Bergström, Anders et al. 2021. “Origins of Modern Human Ancestry.” Nature 590 (7845): 229–37. DOI: 10.1038/s41586-021-03244-5.

Beyer, Robert M. et al. 2021. “Climatic Windows for Human Migration out of Africa in the Past 300,000 Years.” Nature Communications 12 (1): 4889. DOI: 10.1038/s41467-021-24779-1.

Bird, Douglas W. et al. 2019. “Variability in the Organization and Size of Hunter-Gatherer Groups: Foragers Do Not Live in Small-Scale Societies.” Journal of Human Evolution 131 (June): 96–108. DOI: 10.1016/j.jhevol.2019.03.005.

Boaretto, Elisabetta et al. 2021. “The Absolute Chronology of Boker Tachtit (Israel) and Implications for the Middle to Upper Paleolithic Transition in the Levant.” Proceedings of the National Academy of Sciences 118 (25): e2014657118. DOI: 10.1073/pnas.2014657118.

Brandt, Guido et al. 2013. “Ancient DNA Reveals Key Stages in the Formation of Central European Mitochondrial Genetic Diversity.” Science 342 (6155): 257-261. DOI: 10.1126/science.1241844.

Broushaki, Farnaz et al. 2016. “Early Neolithic Genomes from the Eastern Fertile Crescent.” Science 353 (6298): 499–503. DOI: 10.1126/science.aaf7943.

Ceballos, Francisco C. et al. 2018. “Runs of Homozygosity: Windows into Population History and Trait Architecture.” Nature Reviews. Genetics 19 (4): 220–34. DOI: 10.1038/nrg.2017.109.

Chen, Lu et al. 2020. “Identifying and Interpreting Apparent Neanderthal Ancestry in African Individuals.” Cell 180 (4): 677-687.e16. DOI: 10.1016/j.cell.2020.01.012.

Dannemann, Michael and Fernando Racimo. 2018. “Something Old, Something Borrowed: Admixture and Adaptation in Human Evolution.” Current Opinion in Genetics & Development, Genetics of Human Origins, 53 (December): 1–8. DOI: 10.1016/j.gde.2018.05.009.

Dennell, Robin. 2020. From Arabia to the Pacific: How Our Species Colonised Asia. London and New York: Routledge.

Détroit, Florent et al. 2019. “A New Species of Homo from the Late Pleistocene of the Philippines.” Nature 568: 181-186. DOI: 10.1038/s41586-019-1067-9.

Ermini, Luca et al. 2015. “Major Transitions in Human Evolution Revisited: A Tribute to Ancient DNA.” Journal of Human Evolution, Special Issue: Ancient DNA and Human Evolution, 79 (February): 4–20. DOI: 10.1016/j.jhevol.2014.06.015.

Feldman, Michal et al. 2019. “Late Pleistocene Human Genome Suggests a Local Origin for the First Farmers of Central Anatolia.” Nature Communications 10 (1): 1218. DOI: 10.1038/s41467-019-09209-7.

Ferring, Reid et al. 2011. “Earliest Human Occupations at Dmanisi (Georgian Caucasus) Dated to 1.85–1.78 Ma.” Proceedings of the National Academy of Sciences 108 (26): 10432–36. DOI: 10.1073/pnas.1106638108.

Fregel, Rosa et al. 2018. “Ancient Genomes from North Africa Evidence Prehistoric Migrations to the Maghreb from Both the Levant and Europe.” Proceedings of the National Academy of Sciences 115 (26): 6774–79. DOI: 10.1073/pnas.1800851115.

Fu, Qiaomei et al. 2015. “An Early Modern Human from Romania with a Recent Neanderthal Ancestor.” Nature 524 (7564): 216–19. DOI: 10.1038/nature14558.

Fu, Qiaomei et al. 2014. “Genome Sequence of a 45,000-Year-Old Modern Human from Western Siberia.” Nature 514 (7523): 445–49. DOI: 10.1038/nature13810.

Fu, Qiaomei et al. 2016. “The Genetic History of Ice Age Europe.” Nature 534 (7606): 200–205. DOI: 10.1038/nature17993.

Furholt, Martin. 2018. “Massive Migrations? The Impact of Recent aDNA Studies on Our View of Third Millennium Europe.” European Journal of Archaeology 21 (2): 159–91. DOI: 10.1017/eaa.2017.43.

Gabunia, Leo et al. 2000. “Earliest Pleistocene Hominid Cranial Remains from Dmanisi, Republic of Georgia: Taxonomy, Geological Setting, and Age.” Science 288 (5468): 1019–25. DOI: 10.1126/science.288.5468.1019.

Gallego-Llorente, M. et al. 2016. “The Genetics of an Early Neolithic Pastoralist from the Zagros, Iran.” Scientific Reports 6 (1): 31326. DOI: 10.1038/srep31326.

Garcea, Elena A. A. 2012. “Successes and Failures of Human Dispersals from North Africa.” Quaternary International, Late Pleistocene lifeways, an African perspective: selected presentations, PAA-Safa 2010, 270 (August): 119–28. DOI: 10.1016/j.quaint.2011.06.034.

Garcea, Elena A. A. 2016. “Dispersals Out of Africa and Back to Africa: Modern Origins in North Africa.” Quaternary International, Special Issue: The Middle Palaeolithic in the Desert II, 408 (July): 79–89.

Gelabert, Pere et al. 2021. “Genome-Scale Sequencing and Analysis of Human, Wolf, and Bison DNA from 25,000-Year-Old Sediments”. Current Biology 31: 3564-3574. DOI: 10.1016/j.cub.2021.06.023.

Gittelman, Rachel M. et al. 2016. “Archaic Hominin Admixture Facilitated Adaptation to Out-of-Africa Environments.” Current Biology 26 (24): 3375–82. DOI: 10.1016/j.cub.2016.10.041.

Gokcumen, Omer and Michael Frachetti. 2020. “The Impact of Ancient Genome Studies in Archaeology.” Annual Review of Anthropology 49 (1). DOI: 10.1146/annurev-anthro-010220-074353.

Gómez-Robles, Aida. “Dental Evolutionary Rates and Its Implications for the Neanderthal-Modern Human Divergence”. Science Advances 5 (5): eeaw1268. DOI: 10.1126/sciadv.aaw1268.

Goring-Morris, Nigel and Anna Belfer-Cohen. 2020. “Noisy Beginnings: The Initial Upper Palaeolithic in Southwest Asia.” Quaternary International 551: 40–46.

Green, Richard E. et al. 2010. “A Draft Sequence of the Neandertal Genome.” Science 328 (5979): 710–22. DOI: 10.1126/science.1188021.

Greenbaum, Gili et al. 2019. “Was Inter-Population Connectivity of Neanderthals and Modern Humans the Driver of the Upper Paleolithic Transition Rather than Its Product?” Quaternary Science Reviews, Neanderthals: Ecology and Evolution, 217 (August): 316–29. DOI: 10.1016/j.quascirev.2018.12.011.

Groucutt, Huw S. et al. 2021. “Multiple Hominin Dispersals into Southwest Asia over the Past 400,000 Years.” Nature 597 (7876): 376–80. DOI: 10.1038/s41586-021-03863-y.

Haak, Wolfgang et al. 2010. “Ancient DNA from European Early Neolithic Farmers Reveals Their Near Eastern Affinities.” PLOS Biology 8 (11): e1000536. DOI: 10.1371/journal.pbio.1000536.

Hagelberg, Erika et al. 2015. “Ancient DNA: The First Three Decades.” Philosophical Transactions of the Royal Society B: Biological Sciences 370 (1660): 20130371. DOI: 10.1098/rstb.2013.0371.

Haile, James et al. 2007. “Ancient DNA Chronology within Sediment Deposits: Are Paleobiological Reconstructions Possible and Is DNA Leaching a Factor?”. Molecular Biology and Evolution 24(4): 092-989. DOI: 10.1093/molbev/msm016.

Hajdinjak, Mateja et al. 2018. “Reconstructing the Genetic History of Late Neanderthals.” Nature 555 (7698): 652–56. DOI: 10.1038/nature26151.

Hajdinjak, Mateja et al. 2021. “Initial Upper Palaeolithic Humans in Europe Had Recent Neanderthal Ancestry.” Nature 592 (7853): 253–57. DOI: 10.1038/s41586-021-03335-3.

Harvati, Katerina et al. 2019. “Apidima Cave Fossils Provide Earliest Evidence of Homo Sapiens in Eurasia.” Nature 571 (7766): 500–504. DOI: 10.1038/s41586-019-1376-z.

Hayden, Brian. 2012. “Neandertal Social Structure?” Oxford Journal of Archaeology 31 (1): 1–26. DOI: 10.1111/j.1468-0092.2011.00376.x.

Henn, Brenna M. et al. 2012. “Genomic Ancestry of North Africans Supports Back-to-Africa Migrations.” PLOS Genetics 8 (1): e1002397. DOI: 10.1371/journal.pgen.1002397.

Hershkovitz, Israel et al. 2021. “A Middle Pleistocene Homo from Nesher Ramla, Israel.” Science 372 (6549): 1424–28. DOI: 10.1126/science.abh3169.

Hershkovitz, Israel et al. 2018. “The Earliest Modern Humans Outside Africa.” Science 359 (6374): 456–59. DOI: 10.1126/science.aap8369.

Hervella, M. et al. 2016. “The Mitogenome of a 35,000-Year-Old Homo Sapiens from Europe Supports a Palaeolithic Back-Migration to Africa.” Scientific Reports 6 (1): 25501. DOI: 10.1038/srep25501.

Heyd, Volker. 2017. “Kossinna’s Smile.” Antiquity 91 (356): 348–59. DOI: 10.15184/aqy.2017.21.

Higham, Thomas et al. 2014. “The Timing and Spatiotemporal Patterning of Neanderthal Disappearance”. Nature 512: 306-309. DOI: 10.1038/nature13621.

Hill, Kim R. et al. 2011. “Co-Residence Patterns in Hunter-Gatherer Societies Show Unique Human Social Structure.” Science 331 (6022): 1286–89. DOI: 10.1126/science.1199071.

Horsburgh, K. Ann. 2015. “Molecular Anthropology: The Judicial Use of Genetic Data in Archaeology.” Journal of Archaeological Science, Scoping the Future of Archaeological Science: Papers in Honour of Richard Klein, 56 (April): 141–45. DOI: 10.1016/j.jas.2015.02.033.

Hsieh, PingHsun et al. 2016. “Model-Based Analyses of Whole-Genome Data Reveal a Complex Evolutionary History Involving Archaic Introgression in Central African Pygmies.” Genome Research 26 (3): 291–300. DOI: 10.1101/gr.196634.115.

Jobling, Mark A. et al. 2004. Human Evolutionary Genetics: Origins, Peoples & Disease. New

York: Garland Science.

Jones, Elizabeth D. and Elsbeth Bösl. 2021. “Ancient Human DNA: A History of Hype (Then and Now).” Journal of Social Archaeology 21 (2): 236–55. DOI: 10.1177/1469605321990115.

Jones, Eppie R. et al. 2015. “Upper Palaeolithic Genomes Reveal Deep Roots of Modern Eurasians.” Nature Communications 6 (1): 8912. DOI: 10.1038/ncomms9912.

Karger, D.N. et al. 2017. “Climatologies at High Resolution for the Earth Land Surface Areas”. Scientific Data 4: 170122. DOI: 10.1038/sdata.2017.122.

Karger, D.N. et al. 2018. “Data From: Climatologies at High Resolution for the Earth’s Land Surface Areas.” Dryad Digital Repository. DOI: 10.5061/dryad.kd1d4.

Koller, Dora et al. 2022. “Denisovan and Neanderthal Archaic Introgression Differentially Impacted the Genetics of Complex Traits in Modern Populations.” BMC Biology 20 (1): 249. DOI: 10.1186/s12915-022-01449-2.

Kristiansen, Kristian. 2014. “Towards a New Paradigm? The Third Science Revolution and Its Possible Consequences in Archaeology.” Current Swedish Archaeology 22 (1): 11–34. DOI: 10.37718/CSA.2014.01.

Kuhlwilm, Martin et al. 2016. “Ancient Gene Flow from Early Modern Humans into Eastern Neanderthals.” Nature 530 (7591): 429–33. DOI: 10.1038/nature16544.

Kuhn, Steven L. 2003. “In What Sense Is the Levantine Initial Upper Paleolithic a ‘Transitional’ Industry?” In The Chronology of the Aurignacian and of the Transitional Technocomplexes. Dating, Stratigraphies, Cultural Implications, edited by João Zilhão and F. d’Errico, 61–70. Trabalhos de Arqueologia 33. Lisbon: Instituto Português de Arqueologia.

Lalueza-Fox, Carles et al. 2011. “Genetic Evidence for Patrilocal Mating Behavior among Neandertal Groups.” Proceedings of the National Academy of Sciences 108 (1): 250–53. DOI: 10.1073/pnas.1011553108.

Lazaridis, Iosif et al. 2018. “Paleolithic DNA from the Caucasus Reveals Core of West Eurasian Ancestry.” bioRxiv. DOI: 10.1101/423079.

Lazaridis, Iosif et al. 2016. “Genomic Insights into the Origin of Farming in the Ancient Near East.” Nature 536 (7617): 419–24. DOI: 10.1038/nature19310.

Lazaridis, Iosif et al. 2014. “Ancient Human Genomes Suggest Three Ancestral Populations for Present-Day Europeans.” Nature 513 (7518): 409–13. DOI: 10.1038/nature13673.

Liu, Yichen et al. 2021. “Insights into Human History from the First Decade of Ancient Human Genomics.” Science 373 (6562): 1479–84. DOI: 10.1126/science.abi8202.

Lohse, Konrad and Laurent A. F. Frantz. 2014. “Neandertal Admixture in Eurasia Confirmed by Maximum-Likelihood Analysis of Three Genomes.” Genetics 196 (4): 1241–51. DOI: 10.1534/genetics.114.162396.

van de Loosdrecht, Marieke et al. 2018. “Pleistocene North African Genomes Link Near Eastern and Sub-Saharan African Human Populations.” Science 360 (6388): 548–52. DOI: 10.1126/science.aar8380.

Lorente-Galdos et al. 2019. “Whole-Genome Sequence Analysis of a Pan African Set of Samples Reveals Archaic Gene Flow from an Extinct Basal Population of Modern Humans into Sub-Saharan Populations.” Genome Biology 20 (1): 77. DOI: 10.1186/s13059-019-1684-5.

de Lumley, Marie-Antoinette et al. 2020. “Apidima 1 and Apidima 2: Two Anteneandertal Skulls in the Peloponnese, Greece.” L’Anthropologie, APIDIMA, Péloponnèse, Grèce, 124 (1): 102743.DOI: 10.1016/j.anthro.2019.102743.

Malaspinas, Anna-Sapfo et al. 2016. “A Genomic History of Aboriginal Australia.” Nature 538 (7624): 207–14. DOI: 10.1038/nature18299.

Mallick, Swapan et al. 2016. “The Simons Genome Diversity Project: 300 Genomes from 142 Diverse Populations.” Nature 538 (7624): 201–6. DOI: 10.1038/nature18964.

Mallick, Swapan et al. 2023. “The Allen Ancient DNA Resource (AADR): A Curated Compendium of Ancient Human Genomes.” bioRxiv. DOI: 10.1101/2023.04.06.535797.

Mallick, Swapan and David Reich. 2023. “The Allen Ancient DNA Resource (AADR): A Curated Compendium of Ancient Human Genomes.” Harvard Dataverse, V8. DOI: 10.7910/DVN/FFIDCW.

Massilani, Diyendo et al. 2021. ”Microstratigraphic Preservation of Ancient Faunal and Hominin DNA in Pleistocene Cave Sediments”. Proceedings of the National Academy of Sciences 119(1): e2113666118. DOI: 10.1073/pnas.2113666118.

McArthur, Evonne et al. 2021. “Quantifying the Contribution of Neanderthal Introgression to the Heritability of Complex Traits.” Nature Communications 12 (1): 4481. DOI: 10.1038/s41467-021-24582-y.

Meignen, L. 2012. “Levantine Perspectives on the Middle to Upper Paleolithic ‘Transition.’” Archaeology, Ethnology and Anthropology of Eurasia 40 (3): 12–21. DOI: 10.1016/j.aeae.2012.11.003.

Narasimhan, Vagheesh M. et al. 2019. “The formation of human populations in South and Central Asia.” Science 365 (6457): eaat7487. DOI: 10.1126/science.aat7487.

Nielsen, Rasmus et al. 2017. “Tracing the Peopling of the World through Genomics.” Nature 541 (7637): 302–10. DOI: 10.1038/nature21347.

Olalde, Iñigo and Cosimo Posth. 2020. “Latest Trends in Archaeogenetic Research of West Eurasians.” Current Opinion in Genetics & Development, Genetics of Human Origin, 62 (June): 36–43. DOI: 10.1016/j.gde.2020.05.021.

Olszewski, Deborah I. 2017. “The Initial Upper Palaeolithic in the Levant.” In Quaternary of the Levant: Environments, Climate Change, and Humans, edited by Yehouda Enzel and Ofer Bar-Yosef, 621–26. Cambridge: Cambridge University Press. DOI: 10.1017/9781316106754.

Omrak, Ayça et al. 2016. “Genomic Evidence Establishes Anatolia as the Source of the European Neolithic Gene Pool.” Current Biology 26 (2): 270-275. DOI: 10.1016/j.cub.2015.12.019.

Orlando, Ludovic. 2020. “Filling Important Gaps in the Genomic History of Southwest Asia.” Cell 181 (5): 966–68. DOI: 10.1016/j.cell.2020.05.010.

Orlando, Ludovic et al. 2021. “Ancient DNA Analysis.” Nature Reviews Methods Primers 1 (1): 1–26. DOI: 10.1038/s43586-020-00011-0.

Pagani, Luca et al. 2016. “Genomic Analyses Inform on Migration Events during the Peopling of Eurasia.” Nature 538 (7624): 238–42. DOI: 10.1038/nature19792.

Pedersen, Mikkel Winther et al. 2015. “Ancient and Modern Environmental DNA.” Philosophical Transactions of the Royal Society B: Biological Sciences 370 (1660): 20130383. DOI: 10.1098/rstb.2013.0383.

Peyrégne, Stéphane, Viviane Slon, and Janet Kelso. 2024. “More Than a Decade of Genetic Research on the Denisovans.” Nature Reviews Genetics 25: 83-103. DOI: 10.1038/s41576-023-00643-4.

Pedersen, Mikkel Winther et al. 2015. “Ancient and Modern Environmental DNA”. Philosophical Transactions of the Royal Society B: Biological Sciences 370(1660): 20130383. DOI: 10.1098/rstb.2013.0383.

Petr, Martin et al. 2020. “The Evolutionary History of Neanderthal and Denisovan Y Chromosomes.” Science 369 (6511): 1653–56. DOI: 10.1126/science.abb6460.

Posth, Cosimo et al. 2017. “Deeply Divergent Archaic Mitochondrial Genome Provides Lower Time Boundary for African Gene Flow into Neanderthals.” Nature Communications 8 (1): 16046. DOI: 10.1038/ncomms16046.

Posth, Cosimo et al. 2023. “Palaeogenomics of Upper Palaeolithic to Neolithic European Hunter-Gatherers.” Nature 615: 117-126. DOI: 10.1038/s41586-023-05726-0.

Prüfer, Kay et al. 2017. “A High-Coverage Neandertal Genome from Vindija Cave in Croatia.” Science 358 (6363): 655–58. DOI: 10.1126/science.aao1887.

Prüfer, Kay et al. 2021. “A Genome Sequence from a Modern Human Skull over 45,000 Years Old from Zlatý Kůň in Czechia.” Nature Ecology & Evolution 5 (6): 820–25. DOI: 10.1038/s41559-021-01443-x.

Prüfer, Kay et al. 2014. “The Complete Genome Sequence of a Neanderthal from the Altai Mountains.” Nature 505 (7481): 43–49. DOI: 10.1038/nature12886.

Quilodrán, Claudio S. et al. 2023. “Past Human Expansions Shaped the Spatial Pattern of Neanderthal Ancestry”. Science Advances 9(42): eadg9817. DOI: 10.1126/sciadv.adg9817.

R Core Team 2023. R: A language and environment for statistical computing. R Foundation for Statistical Computing: Vienna, Austria.

Rabett, Ryan J. 2018. “The Success of Failed Homo Sapiens Dispersals out of Africa and into Asia.” Nature Ecology & Evolution 2 (2): 212–19. DOI: 10.1038/s41559-017-0436-8.

Reilly, Patrick F. et al. 2022. “The Contribution of Neanderthal Introgression to Modern Human Traits.” Current Biology 32 (18): R970–83. DOI: 10.1016/j.cub.2022.08.027.

Richter, Tobias et al. 2011. “Interaction before Agriculture: Exchanging Material and Sharing Knowledge in the Final Pleistocene Levant.” Cambridge Archaeological Journal 21 (1): 95–114. DOI: 10.1017/S0959774311000060.

Roberts, Benjamin W. and Marc Vander Linden. 2011. “Investigating Archaeological Cultures: Material Culture, Variability, and Transmission.” In Investigating Archaeological Cultures: Material Culture, Variability, and Transmission, edited by Benjamin W. Roberts and Marc Vander Linden, 1–21. New York, NY: Springer. DOI: 10.1007/978-1-4419-6970-5_1.

Ronen, A. 1991. “The Yiron-Gravel Lithic Assemblage: Artifacts Older than 2.4 My in Israel.” Archäeologisches Korrespondenzblatt 21 (2): 159–65.

Rosas, Antionio and Markus Bastir. 2020. “An Assessment of the Late Middle Pleistocene Occipital from Apidima 1 Skull (Greece).” L’Anthropologie, APIDIMA, Péloponnèse, Grèce, 124 (1): 102745. DOI: 10.1016/j.anthro.2020.102745.

Sánchez-Quinto et al. 2012. “North African Populations Carry the Signature of Admixture with Neandertals.” PLOS ONE 7 (10): e47765. DOI: 10.1371/journal.pone.0047765.

Sankararaman, Sriram et al. 2012. “The Date of Interbreeding between Neandertals and Modern Humans.” PLOS Genetics 8 (10): e1002947. DOI: 10.1371/journal.pgen.1002947.

Scardia, Giancarlo et al. 2019. “Chronological Constraints on Hominin Dispersal Outside Africa Since 2.48 Ma from the Zarqa Valley, Jordan.” Quaternary Science Reviews 219: 1-19. DOI: 10.1016/j.quascirev.2019.06.007.

Scardia, Giancarlo et al. 2021. “What Kind of Hominin First Left Africa?” Evolutionary Anthropology: Issues, News, and Reviews 30(2): 122-127. DOI: 10.1002/evan.21863.

Scarre, Christopher. 2009. The Human Past: World Prehistory and the Development of Human Societies. 2. ed. London: Thames & Hudson.

Sharp, Warren D. and James B. Paces. 2018. “Comment on ‘The Earliest Modern Humans Outside Africa.’” Science 362 (6413): eaat6598. DOI: 10.1126/science.aat6598.

Shea, John J. 2001. “The Middle Paleolithic: Early Modern Humans and Neandertals in the Levant.” Near Eastern Archaeology 64 (1–2): 38–64. DOI: 10.2307/3210819.

Shea, John J. 2003. “Neandertals, Competition, and the Origin of Modern Human Behavior in the Levant.” Evolutionary Anthropology: Issues, News, and Reviews 12 (4): 173–87. DOI: 10.1002/evan.10101.

Shea, John J. 2008. “Transitions or Turnovers? Climatically-Forced Extinctions of Homo Sapiens and Neanderthals in the East Mediterranean Levant.” Quaternary Science Reviews, The Coastal Shelf of the Mediterranean and Beyond: Corridor and Refugium for Human Populations in the Pleistocene, 27 (23): 2253–70. DOI: 10.1016/j.quascirev.2008.08.015.

Singh, Manvir and Luke Glowacki. 2022. “Human Social Organization during the Late Pleistocene: Beyond the Nomadic-Egalitarian Model.” Evolution and Human Behavior 43 (5): 418–31. DOI: 10.1016/j.evolhumbehav.2022.07.003.

Skourtanioti, Eirini et al. 2020. “Genomic History of Neolithic to Bronze Age Anatolia, Northern Levant, and Southern Caucasus.” Cell 181 (5): 1158-1175.e28. DOI: 10.1016/j.cell.2020.04.044.

Skov, Laurits et al. 2022. “Genetic Insights into the Social Organization of Neanderthals.” Nature 610 (7932): 519–25. DOI: 10.1038/s41586-022-05283-y.

Slimak, Ludovic et al. 2023. “A Late Neanderthal Reveals Genetic Isolation in their Populations Before Extinction”. bioRxiv. DOI: 10.1101/2023.04.10.536015.

Slon, Viviane et al. 2017. “Neanderthal and Denisovan DNA from Pleistocene Sediments”. Science 356(6338): 605-608. DOI: 10.1126/science.aam9695.

Slon, Viviane et al. 2018. “The Genome of the Offspring of a Neanderthal Mother and a Denisovan Father.” Nature 561 (7721): 113–16. DOI: 10.1038/s41586-018-0455-x.

Smith, Colin I. et al. 2003. “The Thermal History of Human Fossils and the Likelihood of Successful DNA Amplification.” Journal of Human Evolution 45 (3): 203–17. DOI: 10.1016/S0047-2484(03)00106-4.

Soares, Pedro et al. 2012. “The Expansion of mtDNA Haplogroup L3 within and out of Africa.” Molecular Biology and Evolution 29 (3): 915–27. DOI: 10.1093/molbev/msr245.

Somel, Mehmet et al. 2021. “Collaborate Equitably in Ancient DNA Research and Beyond.” Nature 600 (7887): 37–37. DOI: 10.1038/d41586-021-03541-z.

Stewart, J. R. and Chris Stringer. 2012. “Human Evolution Out of Africa: The Role of Refugia and Climate Change.” Science 335 (6074): 1317–21. DOI: 10.1126/science.1215627.

Stringer, Chris. 2002. “Modern Human Origins: Progress and Prospects.” Philosophical Transactions of the Royal Society B: Biological Sciences 357 (1420): 563–79. DOI: 10.1098/rstb.2001.1057.

Stringer, Chris and Lucile Crété. 2022. “Mapping Interactions of H. Neanderthalensis and Homo Sapiens from the Fossil and Genetic Records.” PaleoAnthropology 2022 (2). DOI: 10.48738/2022.iss2.130.

Sutikna, Thomas et al. 2016. “Revised Stratigraphy and Chronology for Homo floresiensis at Liang Bua in Indonesia.” Nature 532: 366-369. DOI: 10.1038/nature17179.

Tierney, Jessica E. et al. 2017. “A Climatic Context for the Out-of-Africa Migration.” Geology 45 (11): 1023–26. DOI: 10.1130/G39457.1.

Tsosie, Krystal S. et al. 2021. “Ancient-DNA Researchers Write Their Own Rules.” Nature 600 (7887): 37–37. DOI: 10.1038/d41586-021-03542-y.

Underhill, Peter A. and Toomas Kivisild. 2007. “Use of Y Chromosome and Mitochondrial DNA Population Structure in Tracing Human Migrations.” Annual Review of Genetics 41 (1): 539–64. DOI: 10.1146/annurev.genet.41.110306.130407.

Veeramah, Krishna R. 2018. “The Importance of Fine-Scale Studies for Integrating Paleogenomics and Archaeology.” Current Opinion in Genetics & Development, Genetics of Human Origins, 53 (December): 83–89. DOI: 10.1016/j.gde.2018.07.007.

Vernot, Benjamin et al. 2021. “Unearthing Neanderthal Population History Using Nuclear and Mitochondrial DNA from Cave Sediments”. Science 372(6542): eabf1667. DOI: 10.1126/science.abf1667.

Villanea, Fernando A. and Joshua G. Schraiber. 2019. “Multiple Episodes of Interbreeding between Neanderthal and Modern Humans.” Nature Ecology & Evolution 3 (1): 39–44. DOI: 10.1038/s41559-018-0735-8.

Wagner, Jennifer K. et al. 2020. “Fostering Responsible Research on Ancient DNA.” The American Journal of Human Genetics 107 (2): 183–95. DOI: 10.1016/j.ajhg.2020.06.017.

Wickham et al. 2019. “Welcome to the tidyverse.” Journal of Open Source Software 4 (43): 1686. DOI: 10.21105/joss.01686.

Williams, Sloan R. 2019. “The Neanderthal and Denisovan Genomes.” In A Companion to Anthropological Genetics, edited by Dennis H. O’Rourke. 69–88. John Wiley & Sons, Ltd. DOI: 10.1002/9781118768853.ch6.

Zavala, Elena I. et al. 2021. “Pleistocene Sediment DNA Reveals Hominin and Faunal Turnovers at Denisova Cave”. Nature 595(7867): 399-403. DOI: 10.1038/s41586-021-03675-0.

Zhang, Dongju et al. 2021. “Denisovan DNA in Late Pleistocene Sediments from Baishiya Karst Cave on the Tibetan Plateau”. Science 370(6516): 584-587. DOI: 10.1126/science.abb6320.

Zhu, Zhaoyu et al. 2018. “Hominin Occupation of the Chinese Loess Plateau Since About 2.1 Million Years Ago.” Nature 559(7715): 608-612. DOI: 10.1038/s41586-018-0299-4.

Zilhão, João. 2014. “The Neanderthals: Evolution, Palaeoecology, and Extinction.” In The Oxford Handbook of the Archaeology and Anthropology of Hunter-Gatherers, edited by Vicki Cummings et al. Oxford University Press. DOI: 10.1093/oxfordhb/9780199551224.013.054.


Additional Files



How to Cite

L. G. Miedzianogora, D. (2024) “Palaeogenomics and the Palaeolithic of Southwest Asia: Trends, Issues, and Future Directions”, Chronolog, 2(2), pp. 6–30. Available at: (Accessed: 22 July 2024).