Testing times: disentangling admixture histories in recent and complex demographies using ancient DNA
Language English Country United States Media print
Document type Journal Article
Grant support
R35GM146886
National Institute of Health
CZ.10.03.01/00/22_003/0000003
European Union
R35 GM146886
NIGMS NIH HHS - United States
CZ.10.03.01/00/22_003/0000003
European Union Operational Programme
61220
John Templeton Foundation
21-27624S
Czech Science Foundation
LL2103
Czech Ministry of Education, Youth and Sports
PubMed
39013011
PubMed Central
PMC11373510
DOI
10.1093/genetics/iyae110
PII: 7714968
Knihovny.cz E-resources
- Keywords
- f-statistics, aDNA, admixture, ancient DNA, archaeogenetics, paleogenomics, qpAdm,
- MeSH
- Demography MeSH
- Genomics * methods MeSH
- Models, Genetic MeSH
- Paleontology * methods MeSH
- Software MeSH
- Data Accuracy MeSH
- Publication type
- Journal Article MeSH
Our knowledge of human evolutionary history has been greatly advanced by paleogenomics. Since the 2020s, the study of ancient DNA has increasingly focused on reconstructing the recent past. However, the accuracy of paleogenomic methods in resolving questions of historical and archaeological importance amidst the increased demographic complexity and decreased genetic differentiation remains an open question. We evaluated the performance and behavior of two commonly used methods, qpAdm and the f3-statistic, on admixture inference under a diversity of demographic models and data conditions. We performed two complementary simulation approaches-firstly exploring a wide demographic parameter space under four simple demographic models of varying complexities and configurations using branch-length data from two chromosomes-and secondly, we analyzed a model of Eurasian history composed of 59 populations using whole-genome data modified with ancient DNA conditions such as SNP ascertainment, data missingness, and pseudohaploidization. We observe that population differentiation is the primary factor driving qpAdm performance. Notably, while complex gene flow histories influence which models are classified as plausible, they do not reduce overall performance. Under conditions reflective of the historical period, qpAdm most frequently identifies the true model as plausible among a small candidate set of closely related populations. To increase the utility for resolving fine-scaled hypotheses, we provide a heuristic for further distinguishing between candidate models that incorporates qpAdm model P-values and f3-statistics. Finally, we demonstrate a significant performance increase for qpAdm using whole-genome branch-length f2-statistics, highlighting the potential for improved demographic inference that could be achieved with future advancements in f-statistic estimations.
Department of Biology and Ecology University of Ostrava Ostrava 701 03 Czechia
Department of Biology Pennsylvania State University University Park PA 16802 USA
Department of Human Evolutionary Biology Harvard University Cambridge MA 02138 USA
See more in PubMed
Adrion JR, Cole CB, Dukler N, Galloway JG, Gladstein AL, Gower G, Kyriazis CC, Ragsdale AP, Tsambos G, Baumdicker F, et al. . 2020. A community-maintained standard library of population genetic models. eLife. 9:e54967. doi:10.7554/elife.54967. PubMed DOI PMC
Agranat-Tamir L, Waldman S, Martin MAS, Gokhman D, Mishol N, Eshel T, Cheronet O, Rohland N, Mallick S, Adamski N, et al. . 2020. The genomic history of the Bronze Age Southern Levant. Cell. 181(5):1146–1157.e11. doi:10.1016/j.cell.2020.04.024. PubMed DOI PMC
Alexander DH, Novembre J, Lange K. 2009. Fast model-based estimation of ancestry in unrelated individuals. Genome Res. 19(9):1655–1664. doi:10.1101/gr.094052.109. PubMed DOI PMC
Antonio ML, Gao Z, Moots HM, Lucci M, Candilio F, Sawyer S, Oberreiter V, Calderon D, Devitofranceschi K, Aikens RC, et al. . 2019. Ancient Rome: a genetic crossroads of Europe and the Mediterranean. Science. 366(6466):708–714. doi:10.1126/science.aay6826. PubMed DOI PMC
Arning N, Wilson DJ. 2020. The past, present and future of ancient bacterial DNA. Microb Genom. 6(7):mgen000384. doi:10.1099/mgen.0.000384. PubMed DOI PMC
Ávila-Arcos MC, Raghavan M, Schlebusch C. 2023. Going local with ancient DNA: a review of human histories from regional perspectives. Science. 382(6666):53–58. doi:10.1126/science.adh8140. PubMed DOI
Bartash V. 2020. The early dynastic near East. In: Radner K, Moeller N, Potts DT, editors. The Oxford History of the Ancient Near East. From the Beginnings to Old Kingdom Egypt and the Dynasty of Akkad. New York: Oxford Academic. p. 531–6111. doi:10.1093/oso/9780190687854.003.0009. DOI
Baumdicker F, Bisschop G, Goldstein D, Gower G, Ragsdale AP, Tsambos G, Zhu S, Eldon B, Ellerman EC, Galloway JG, et al. . 2022. Efficient ancestry and mutation simulation with msprime 1.0. Genetics. 220(3):iyab229. doi:10.1093/genetics/iyab229. PubMed DOI PMC
Bergström A, McCarthy SA, Hui R, Almarri MA, Ayub Q, Danecek P, Chen Y, Felkel S, Hallast P, Kamm J, et al. . 2020. Insights into human genetic variation and population history from 929 diverse genomes. Science. 367(6484):eaay5012. doi:10.1126/science.aay5012. PubMed DOI PMC
Bergström A, Stanton DWG, Taron UH, Frantz L, Sinding M-HS, Ersmark E, Pfrengle S, Cassatt-Johnstone M, Lebrasseur O, Girdland-Flink L, et al. . 2022. Grey wolf genomic history reveals a dual ancestry of dogs. Nature. 607(7918):313–320. doi:10.1038/s41586-022-04824-9. PubMed DOI PMC
Broushaki F, Thomas MG, Link V, López S, van Dorp L, Kirsanow K, Hofmanová Z, Diekmann Y, Cassidy LM, Díez-Del-Molino D, et al. . 2016. Early Neolithic genomes from the eastern Fertile Crescent. Science. 353(6298):499–503. doi:10.1126/science.aaf7943. PubMed DOI PMC
Brunson K, Reich D. 2019. The promise of paleogenomics beyond our own species. Trends Genet. 35(5):319–329. doi:10.1016/j.tig.2019.02.006. PubMed DOI
Carlhoff S, Duli A, Nägele K, Nur M, Skov L, Sumantri I, Oktaviana AA, Hakim B, Burhan B, Syahdar FA, et al. . 2021. Genome of a middle Holocene hunter-gatherer from Wallacea. Nature. 596(7873):543–547. doi:10.1038/s41586-021-03823-6. PubMed DOI PMC
Changmai P, Jaisamut K, Kampuansai J, Kutanan W, Altınışık NE, Flegontova O, Inta A, Yüncü E, Boonthai W, Pamjav H, et al. . 2022a. Indian genetic heritage in Southeast Asian populations. PLoS Genet. 18(2):e1010036. doi:10.1371/journal.pgen.1010036. PubMed DOI PMC
Changmai P, Pinhasi R, Pietrusewsky M, Stark MT, Ikehara-Quebral RM, Reich D, Flegontov P. 2022b. Ancient DNA from protohistoric period Cambodia indicates that South Asians admixed with local populations as early as 1st–3rd centuries CE. Sci Rep. 12(1):22507. doi:10.1038/s41598-022-26799-3. PubMed DOI PMC
Clemente F, Unterländer M, Dolgova O, Amorim CEG, Coroado-Santos F, Neuenschwander S, Ganiatsou E, Cruz Dávalos DI, Anchieri L, Michaud F, et al. . 2021. The genomic history of the Aegean palatial civilizations. Cell. 184(10):2565–2586.e21. doi:10.1016/j.cell.2021.03.039. PubMed DOI PMC
de Damgaard PB, Martiniano R, Kamm J, Moreno-Mayar JV, Kroonen G, Peyrot M, Barjamovic G, Rasmussen S, Zacho C, Baimukhanov N, et al. . 2018. The first horse herders and the impact of early Bronze Age steppe expansions into Asia. Science. 360(6396):eaar7711. doi:10.1126/science.aar7711. PubMed DOI PMC
Durand EY, Patterson N, Reich D, Slatkin M. 2011. Testing for ancient admixture between closely related populations. Mol Biol Evol. 28(8):2239–2252. doi:10.1093/molbev/msr048. PubMed DOI PMC
Feldman M, Master DM, Bianco RA, Burri M, Stockhammer PW, Mittnik A, Aja AJ, Jeong C, Krause J. 2019. Ancient DNA sheds light on the genetic origins of early Iron Age Philistines. Sci Adv. 5(7). doi:10.1126/sciadv.aax0061. PubMed DOI PMC
Fernandes DM, Mittnik A, Olalde I, Lazaridis I, Cheronet O, Rohland N, Mallick S, Bernardos R, Broomandkhoshbacht N, Carlsson J, et al. . 2020. The spread of steppe and Iranian-related ancestry in the islands of the Western Mediterranean. Nat Ecol Evol. 4(3):334–345. doi:10.1038/s41559-020-1102-0. PubMed DOI PMC
Flegontov P, Işıldak U, Maier R, Yüncü E, Changmai P, Reich D. 2023. Modeling of African population history using f-statistics is biased when applying all previously proposed SNP ascertainment schemes. PLOS Genet. 19(9):e1010931. doi:10.1371/journal.pgen.1010931. PubMed DOI PMC
Fregel R, Méndez FL, Bokbot Y, Martín-Socas D, Camalich-Massieu MD, Santana J, Morales J, Ávila-Arcos MC, Underhill PA, Shapiro B, et al. . 2018. Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe. Proc Natl Acad Sci. 115(26):6774–6779. doi:10.1073/pnas.1800851115. PubMed DOI PMC
Fu Q, Posth C, Hajdinjak M, Petr M, Mallick S, Fernandes D, Furtwängler A, Haak W, Meyer M, Mittnik A, et al. . 2016. The genetic history of Ice Age Europe. Nature. 534(7606):200–205. doi:10.1038/nature17993. PubMed DOI PMC
Gower G, Ragsdale AP, Bisschop G, Gutenkunst RN, Hartfield M, Noskova E, Schiffels S, Struck TJ, Kelleher J, Thornton KR, et al. . 2022. Demes: a standard format for demographic models. Genetics. 222(3):iyac131. doi:10.1093/genetics/iyac131. PubMed DOI PMC
Green RE, Krause J, Briggs AW, Maricic T, Stenzel U, Kircher M, Patterson N, Li H, Zhai W, Fritz MH-Y, et al. . 2010. A draft sequence of the neandertal genome. Science. 328(5979):710–722. doi:10.1126/science.1188021. PubMed DOI PMC
Haak W, Lazaridis I, Patterson N, Rohland N, Mallick S, Llamas B, Brandt G, Nordenfelt S, Harney E, Stewardson K, et al. . 2015. Massive migration from the steppe was a source for Indo-European languages in Europe. Nature. 522(7555):207–211. doi:10.1038/nature14317. PubMed DOI PMC
Haber M, Doumet-Serhal C, Scheib C, Xue Y, Danecek P, Mezzavilla M, Youhanna S, Martiniano R, Prado-Martinez J, Szpak M, et al. . 2017. Continuity and admixture in the last five millennia of Levantine history from ancient Canaanite and present-day Lebanese genome sequences. Am J Hum Genet. 101(2):274–282. doi:10.1016/j.ajhg.2017.06.013. PubMed DOI PMC
Haber M, Nassar J, Almarri MA, Saupe T, Saag L, Griffith SJ, Doumet-Serhal C, Chanteau J, Saghieh-Beydoun M, Xue Y, et al. . 2020. A genetic history of the near east from an aDNA time course sampling eight points in the past 4,000 years. Am J Hum Genet. 107(1):149–157. doi:10.1016/j.ajhg.2020.05.008. PubMed DOI PMC
Haber M, Mezzavilla M, Xue Y, Tyler-Smith C. 2016. Ancient DNA and the rewriting of human history: be sparing with Occam's razor. Genome Biol. 17(1):1. doi:10.1186/s13059-015-0866-z. PubMed DOI PMC
Harney É, May H, Shalem D, Rohland N, Mallick S, Lazaridis I, Sarig R, Stewardson K, Nordenfelt S, Patterson N, et al. . 2018. Ancient DNA from Chalcolithic Israel reveals the role of population mixture in cultural transformation. Nat Commun. 9(1):3336. doi:10.1038/s41467-018-05649-9. PubMed DOI PMC
Harney É, Patterson N, Reich D, Wakeley J. 2021. Assessing the performance of qpAdm: a statistical tool for studying population admixture. Genetics. 217(4):iyaa045. doi:10.1093/genetics/iyaa045. PubMed DOI PMC
Harris AM, DeGiorgio M. 2017. Admixture and ancestry inference from ancient and modern samples through measures of population genetic drift. Hum Biol. 89(1):21–46. doi:10.13110/humanbiology.89.1.02. PubMed DOI
Hollfelder N, Breton G, Sjödin P, Jakobsson M. 2021. The deep population history in Africa. Hum Mol Genet. 30(R1):R2–R10. doi:10.1093/hmg/ddab005. PubMed DOI PMC
Jónsson H, Sulem P, Kehr B, Kristmundsdottir S, Zink F, Hjartarson E, Hardarson MT, Hjorleifsson KE, Eggertsson HP, Gudjonsson SA, et al. . 2017. Parental influence on human germline de novo mutations in 1,548 trios from Iceland. Nature. 549(7673):519–522. doi:10.1038/nature24018. PubMed DOI
Jukes TH, Cantor CR. 1969. Mammalian protein metabolism. In: Munro HN, editor. Part IV: Protein Metab. Evol. Dev. Mamm. Cambridge (MA): Academic Press. p. 21–132. doi:10.1016/b978-1-4832-3211-9.50009-7III. DOI
Kamm J, Terhorst J, Durbin R, Song YS. 2020. Efficiently inferring the demographic history of many populations with allele count data. J Am Stat Assoc. 115(531):1472–1487. doi:10.1080/01621459.2019.1635482. PubMed DOI PMC
Kelleher J, Etheridge AM, McVean G. 2016. Efficient coalescent simulation and genealogical analysis for large sample sizes. PLoS Comput Biol. 12(5):e1004842. doi:10.1371/journal.pcbi.1004842. PubMed DOI PMC
Knappett C, Kiriatzi E. 2016. Technological mobilities: perspectives from the Eastern Mediterranean—an Introduction. In: Kiriatzi E, Knappett C, editors. Human Mobility and Technological Transfer in the Prehistoric Mediterranean. British School at Athens Studies in Greek Antiquity: Cambridge University Press, British School at Athens Studies in Greek Antiquity. p. 1–17.
Koptekin D, Yüncü E, Rodríguez-Varela R, Altınışık NE, Psonis N, Kashuba N, Yorulmaz S, George R, Kazancı DD, Kaptan D, et al. . 2023. Spatial and temporal heterogeneity in human mobility patterns in Holocene Southwest Asia and the East Mediterranean. Curr Biol. 33(1):41–57.e15. doi:10.1016/j.cub.2022.11.034. PubMed DOI PMC
Kristiansen K. 2016. 10 - interpreting Bronze Age trade and migration. In: Kiriatzi E, Knappett C, editors. Human Mobility and Technological Transfer in the Prehistoric Mediterranean. British School at Athens Studies in Greek Antiquity: Cambridge University Press. p. 154–180. doi:10.1017/9781316536063.011. DOI
Lauterbur ME, Cavassim MIA, Gladstein AL, Gower G, Pope NS, Tsambos G, Adrion J, Belsare S, Biddanda A, Caudill V, et al. . 2023. Expanding the stdpopsim species catalog, and lessons learned for realistic genome simulations. eLife. 12:RP84874. doi:10.7554/elife.84874. PubMed DOI PMC
Lazaridis I, Alpaslan-Roodenberg S, Acar A, Açıkkol A, Agelarakis A, Aghikyan L, Akyüz U, Andreeva D, Andrijašević G, Antonović D, et al. . 2022a. A genetic probe into the ancient and medieval history of Southern Europe and West Asia. Science. 377(6609):940–951. doi:10.1126/science.abq0755. PubMed DOI PMC
Lazaridis I, Alpaslan-Roodenberg S, Acar A, Açıkkol A, Agelarakis A, Aghikyan L, Akyüz U, Andreeva D, Andrijašević G, Antonović D, et al. . 2022b. The genetic history of the Southern Arc: a bridge between West Asia and Europe. Science. 377(6609):eabm4247. doi:10.1126/science.abm4247. PubMed DOI PMC
Lazaridis I, Belfer-Cohen A, Mallick S, Patterson N, Cheronet O, Rohland N, Bar-Oz G, Bar-Yosef O, Jakeli N, Kvavadze E, et al. . 2018. Paleolithic DNA from the Caucasus reveals core of West Eurasian ancestry. bioRxiv. 10.1101/423079, preprint: not peer reviewed. DOI
Lazaridis I, Mittnik A, Patterson N, Mallick S, Rohland N, Pfrengle S, Furtwängler A, Peltzer A, Posth C, Vasilakis A, et al. . 2017. Genetic origins of the Minoans and Mycenaeans. Nature. 548(7666):214–218. doi:10.1038/nature23310. PubMed DOI PMC
Lazaridis I, Nadel D, Rollefson G, Merrett DC, Rohland N, Mallick S, Fernandes D, Novak M, Gamarra B, Sirak K, et al. . 2016. Genomic insights into the origin of farming in the ancient Near East. Nature. 536(7617):419–424. doi:10.1038/nature19310. PubMed DOI PMC
Lee J, Miller BK, Bayarsaikhan J, Johannesson E, Miller AV, Warinner C, Jeong C. 2023. Genetic population structure of the Xiongnu Empire at imperial and local scales. Sci Adv. 9(15):eadf3904. doi:10.1126/sciadv.adf3904. PubMed DOI PMC
Librado P, Khan N, Fages A, Kusliy MA, Suchan T, Tonasso-Calvière L, Schiavinato S, Alioglu D, Fromentier A, Perdereau A, et al. . 2021. The origins and spread of domestic horses from the Western Eurasian steppes. Nature. 598(7882):634–640. doi:10.1038/s41586-021-04018-9. PubMed DOI PMC
Lipson M, Loh P-R, Levin A, Reich D, Patterson N, Berger B. 2013. Efficient moment-based inference of admixture parameters and sources of gene flow. Mol Biol Evol. 30(8):1788–1802. doi:10.1093/molbev/mst099. PubMed DOI PMC
Lipson M, Loh P-R, Patterson N, Moorjani P, Ko Y-C, Stoneking M, Berger B, Reich D. 2014. Reconstructing Austronesian population history in Island Southeast Asia. Nat Commun. 5(1):4689. doi:10.1038/ncomms5689. PubMed DOI PMC
Liu Y, Mao X, Krause J, Fu Q. 2021. Insights into human history from the first decade of ancient human genomics. Science. 373(6562):1479–1484. doi:10.1126/science.abi8202. PubMed DOI
Llamas B, Willerslev E, Orlando L. 2017. Human evolution: a tale from ancient genomes. Philos Trans R Soc B Biol Sci. 372(1713):20150484. doi:10.1098/rstb.2015.0484. PubMed DOI PMC
Maier R, Flegontov P, Flegontova O, Işıldak U, Changmai P, Reich D. 2023. On the limits of fitting complex models of population history to f-statistics. eLife. 12:e85492. doi:10.7554/elife.85492. PubMed DOI PMC
Mallick S, Micco A, Mah M, Ringbauer H, Lazaridis I, Olalde I, Patterson N, Reich D. 2024. The Allen Ancient DNA Resource (AADR) a curated compendium of ancient human genomes. Sci Data. 11(1):182. doi:10.1038/s41597-024-03031-7. PubMed DOI PMC
Marchi N, Winkelbach L, Schulz I, Brami M, Hofmanová Z, Blöcher J, Reyna-Blanco CS, Diekmann Y, Thiéry A, Kapopoulou A, et al. . 2022. The genomic origins of the world's first farmers. Cell. 185(11):1842–1859.e18. doi:10.1016/j.cell.2022.04.008. PubMed DOI PMC
Marcus JH, Posth C, Ringbauer H, Lai L, Skeates R, Sidore C, Beckett J, Furtwängler A, Olivieri A, Chiang CWK, et al. . 2020. Genetic history from the Middle Neolithic to present on the Mediterranean island of Sardinia. Nat Commun. 11(1):939. doi:10.1038/s41467-020-14523-6. PubMed DOI PMC
Maróti Z, Neparáczki E, Schütz O, Maár K, Varga GIB, Kovács B, Kalmár T, Nyerki E, Nagy I, Latinovics D, et al. . 2022. The genetic origin of Huns, Avars, and conquering Hungarians. Curr Biol. 32(13):2858–2870.e7. doi:10.1016/j.cub.2022.04.093. PubMed DOI
Martin SH, Davey JW, Jiggins CD. 2015. Evaluating the use of ABBA–BABA statistics to locate introgressed loci. Mol Biol Evol. 32(1):244–257. doi:10.1093/molbev/msu269. PubMed DOI PMC
Mathieson I, Alpaslan-Roodenberg S, Posth C, Szécsényi-Nagy A, Rohland N, Mallick S, Olalde I, Broomandkhoshbacht N, Candilio F, Cheronet O, et al. . 2018. The genomic history of Southeastern Europe. Nature. 555(7695):197–203. doi:10.1038/nature25778. PubMed DOI PMC
McVean G. 2009. A genealogical interpretation of principal components analysis. PLoS Genet. 5(10):e1000686. doi:10.1371/journal.pgen.1000686. PubMed DOI PMC
Mitchell KJ, Rawlence NJ. 2021. Examining natural history through the lens of palaeogenomics. Trends Ecol Evol. 36(3):258–267. doi:10.1016/j.tree.2020.10.005. PubMed DOI
Moots HM, Antonio M, Sawyer S, Spence JP, Oberreiter V, Weiß CL, Lucci M, Cherifi YMS, La Pastina F, Genchi F, et al. . 2023. A genetic history of continuity and mobility in the Iron Age Central Mediterranean. Nat Ecol Evol. 7:1515–1524. doi:10.1038/s41559-023-02143-4. PubMed DOI
Narasimhan VM, Patterson N, Moorjani P, Rohland N, Bernardos R, Mallick S, Lazaridis I, Nakatsuka N, Olalde I, Lipson M, et al. . 2019. The formation of human populations in South and Central Asia. Science. 365(6457), eaat7487. doi:10.1126/science.aat7487. PubMed DOI PMC
Nelson D, Kelleher J, Ragsdale AP, Moreau C, McVean G, Gravel S. 2020. Accounting for long-range correlations in genome-wide simulations of large cohorts. PLoS Genet. 16(5):e1008619. doi:10.1371/journal.pgen.1008619. PubMed DOI PMC
Nielsen SV, Vaughn AH, Leppälä K, Landis MJ, Mailund T, Nielsen R. 2023. Bayesian inference of admixture graphs on Native American and Arctic populations. PLoS Genet. 19(2):e1010410. doi:10.1371/journal.pgen.1010410. PubMed DOI PMC
Ning C, Li T, Wang K, Zhang F, Li T, Wu X, Gao S, Zhang Q, Zhang H, Hudson MJ, et al. . 2020. Ancient genomes from northern China suggest links between subsistence changes and human migration. Nat Commun. 11(1):2700. doi:10.1038/s41467-020-16557-2. PubMed DOI PMC
Olalde I, Mallick S, Patterson N, Rohland N, Villalba-Mouco V, Silva M, Dulias K, Edwards CJ, Gandini F, Pala M, et al. . 2019. The genomic history of the Iberian Peninsula over the past 8000 years. Science. 363(6432):1230–1234. doi:10.1126/science.aav4040. PubMed DOI PMC
Oliveira S, Nägele K, Carlhoff S, Pugach I, Koesbardiati T, Hübner A, Meyer M, Oktaviana AA, Takenaka M, Katagiri C, et al. . 2022. Ancient genomes from the last three millennia support multiple human dispersals into Wallacea. Nat Ecol Evol. 6(7):1024–1034. doi:10.1038/s41559-022-01775-2. PubMed DOI PMC
Papac L, Ernée M, Dobeš M, Langová M, Rohrlach AB, Aron F, Neumann GU, Spyrou MA, Rohland N, Velemínský P, et al. . 2021. Dynamic changes in genomic and social structures in third millennium BCE central Europe. Sci Adv. 7(35):eabi6941. doi:10.1126/sciadv.abi6941. PubMed DOI PMC
Patterson N, Isakov M, Booth T, Büster L, Fischer C-E, Olalde I, Ringbauer H, Akbari A, Cheronet O, Bleasdale M, et al. . 2022. Large-scale migration into Britain during the middle to late Bronze Age. Nature. 601(7894):588–594. doi:10.1038/s41586-021-04287-4. PubMed DOI PMC
Patterson N, Moorjani P, Luo Y, Mallick S, Rohland N, Zhan Y, Genschoreck T, Webster T, Reich D. 2012. Ancient admixture in human history. Genetics. 192(3):1065–1093. doi:10.1534/genetics.112.145037. PubMed DOI PMC
Patterson N, Price AL, Reich D. 2006. Population structure and eigenanalysis. PLoS Genet. 2(12):e190. doi:10.1371/journal.pgen.0020190. PubMed DOI PMC
Peter BM. 2016. Admixture, population structure, and F-statistics. Genetics. 202(4):1485–1501. doi:10.1534/genetics.115.183913. PubMed DOI PMC
Peter BM. 2022. A geometric relationship of F2, F3 and F4-statistics with principal component analysis. Philos Trans R Soc B. 377(1852):20200413. doi:10.1098/rstb.2020.0413. PubMed DOI PMC
Pickrell JK, Pritchard JK. 2012. Inference of population splits and mixtures from genome-wide allele frequency data. PLoS Genet. 8(11):e1002967. doi:10.1371/journal.pgen.1002967. PubMed DOI PMC
Reich D, Patterson N, Campbell D, Tandon A, Mazieres S, Ray N, Parra MV, Rojas W, Duque C, Mesa N, et al. . 2012. Reconstructing Native American population history. Nature. 488(7411):370–374. doi:10.1038/nature11258. PubMed DOI PMC
Reich D, Price AL, Patterson N. 2008. Principal component analysis of genetic data. Nat Genet. 40(5):491–492. doi:10.1038/ng0508-491. PubMed DOI
Reich D, Thangaraj K, Patterson N, Price AL, Singh L. 2009. Reconstructing Indian population history. Nature. 461(7263):489–494. doi:10.1038/nature08365. PubMed DOI PMC
Renfrew C, Boyle KV. 2000. Archaeogenetics: DNA and the Population Prehistory of Europe, Editors. McDonald Institute monographs. Cambridge: McDonald Institute for Archaeological Research.
Salazar L, Burger R, Forst J, Barquera R, Nesbitt J, Calero J, Washburn E, Verano J, Zhu K, Sop K, et al. . 2023. Insights into the genetic histories and lifeways of Machu Picchu's occupants. Sci Adv. 9(30):eadg3377. doi:10.1126/sciadv.adg3377. PubMed DOI PMC
Schepper SD, Ray JL, Skaar KS, Sadatzki H, Ijaz UZ, Stein R, Larsen A. 2019. The potential of sedimentary ancient DNA for reconstructing past sea ice evolution. ISME J. 13(10):2566–2577. doi:10.1038/s41396-019-0457-1. PubMed DOI PMC
Schmid C, Schiffels S. 2023. Estimating human mobility in Holocene Western Eurasia with large-scale ancient genomic data. Proc Natl Acad Sci. 120(9):e2218375120. doi:10.1073/pnas.2218375120. PubMed DOI PMC
Schuenemann VJ, Peltzer A, Welte B, van Pelt WP, Molak M, Wang C-C, Furtwängler A, Urban C, Reiter E, Nieselt K, et al. . 2017. Ancient Egyptian mummy genomes suggest an increase of sub-Saharan African ancestry in post-Roman periods. Nat Commun. 8(1):15694. doi:10.1038/ncomms15694. PubMed DOI PMC
Sirak KA, Fernandes DM, Lipson M, Mallick S, Mah M, Olalde I, Ringbauer H, Rohland N, Hadden CS, Harney É, et al. . 2021. Social stratification without genetic differentiation at the site of Kulubnarti in Christian Period Nubia. Nat Commun. 12(1):7283. doi:10.1038/s41467-021-27356-8. PubMed DOI PMC
Skoglund P, Thompson JC, Prendergast ME, Mittnik A, Sirak K, Hajdinjak M, Salie T, Rohland N, Mallick S, Peltzer A, et al. . 2017. Reconstructing prehistoric African population structure. Cell. 171(1):59–71.e21. doi:10.1016/j.cell.2017.08.049. PubMed DOI PMC
Skourtanioti E, Erdal YS, Frangipane M, Restelli FB, Yener KA, Pinnock F, Matthiae P, Özbal R, Schoop U-D, Guliyev F, 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. PubMed DOI
Skourtanioti E, Ringbauer H, Ruscone GAG, Bianco RA, Burri M, Freund C, Furtwängler A, Gomes Martins NF, Knolle F, Neumann GU, et al. . 2023. Ancient DNA reveals admixture history and endogamy in the prehistoric Aegean. Nat Ecol Evol. 7(2):290–303. doi:10.1038/s41559-022-01952-3. PubMed DOI PMC
Slatkin M, Racimo F. 2016. Ancient DNA and human history. Proc Natl Acad Sci. 113(23):6380–6387. doi:10.1073/pnas.1524306113. PubMed DOI PMC
Soraggi S, Wiuf C. 2019. General theory for stochastic admixture graphs and F-statistics. Theor Popul Biol. 125:56–66. doi:10.1016/j.tpb.2018.12.002. PubMed DOI
Spyrou MA, Bos KI, Herbig A, Krause J. 2019. Ancient pathogen genomics as an emerging tool for infectious disease research. Nat Rev Genet. 20(6):323–340. doi:10.1038/s41576-019-0119-1. PubMed DOI PMC
Taylor WTT, Librado P, Horse CJA, Gover CSC, Arterberry J, Luta Wiη A, Nujipi A, Omniya T, Gonzalez M, Means B, et al. . 2023. Early dispersal of domestic horses into the Great Plains and northern Rockies. Science. 379(6639):1316–1323. doi:10.1126/science.adc9691. PubMed DOI
Tricou T, Tannier E, de Vienne DM. 2022. Ghost lineages highly influence the interpretation of introgression tests. Syst Biol. 71(5):1147–1158. doi:10.1093/sysbio/syac011. PubMed DOI PMC
van de Loosdrecht M, Bouzouggar A, Humphrey L, Posth C, Barton N, Aximu-Petri A, Nickel B, Nagel S, Talbi EH, Hajraoui MAE, et al. . 2018. Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations. Science. 360(6388):548–552. doi:10.1126/science.aar8380. PubMed DOI
Wang K, Mathieson I, O’Connell J, Schiffels S. 2020. Tracking human population structure through time from whole genome sequences. PLoS Genet. 16(3):e1008552. doi:10.1371/journal.pgen.1008552. PubMed DOI PMC
Wang C-C, Reinhold S, Kalmykov A, Wissgott A, Brandt G, Jeong C, Cheronet O, Ferry M, Harney E, Keating D, et al. . 2019. Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions. Nat Commun. 10(1):590. doi:10.1038/s41467-018-08220-8. PubMed DOI PMC
Wang C-C, Yeh H-Y, Popov AN, Zhang H-Q, Matsumura H, Sirak K, Cheronet O, Kovalev A, Rohland N, Kim AM, et al. . 2021. Genomic insights into the formation of human populations in East Asia. Nature. 591(7850):413–419. doi:10.1038/s41586-021-03336-2. PubMed DOI PMC
Wibowo MC, Yang Z, Borry M, Hübner A, Huang KD, Tierney BT, Zimmerman S, Barajas-Olmos F, Contreras-Cubas C, García-Ortiz H, et al. . 2021. Reconstruction of ancient microbial genomes from the human gut. Nature. 594(7862):234–239. doi:10.1038/s41586-021-03532-0. PubMed DOI PMC
Williams M, Teixeira J. 2020. A genetic perspective on human origins. Biochemist. 42(1):6–10. doi:10.1042/bio04201006. PubMed DOI
Wood SN. 2004. Stable and efficient multiple smoothing parameter estimation for generalized additive models. J Am. Stat Assoc. 99(467):673–686. doi:10.1198/016214504000000980. DOI
Yang MA, Fan X, Sun B, Chen C, Lang J, Ko Y-C, Tsang C-H, Chiu H, Wang T, Bao Q, et al. . 2020. Ancient DNA indicates human population shifts and admixture in northern and southern China. Science. 369(6501):282–288. doi:10.1126/science.aba0909. PubMed DOI
Yüncü E, Işıldak U, Williams MP, Huber CD, Flegontova O, Vyazov LA, Changmai P, Flegontov P. 2023. False discovery rates of qpAdm-based screens for genetic admixture. bioRxiv. 10.1101/2023.04.25.538339, preprint: not peer reviewed. DOI
Zheng Y, Janke A. 2018. Gene flow analysis method, the D-statistic, is robust in a wide parameter space. BMC Bioinform. 19(1):10. 10.1186/s12859-017-2002-4. PubMed DOI PMC