Järve M, Saag L, Scheib CL, Pathak AK, Montinaro F, Pagani L, Flores R, Guellil M, Saag L, Tambets K, Kushniarevich A, Solnik A, Varul L, Zadnikov S, Petrauskas O, Avramenko M, Magomedov B, Didenko S, Toshev G, Bruyako I, et al. 2019. Shifts in the genetic landscape of the Western Eurasian Steppe associated with the beginning and end of the Scythian dominance. *Current Biology* **29**:2430–2441. DOI: https://doi.org/10.1016/j.cub.2019.06.019, PMID: 31303491

Jeong C, Balanovsky O, Lukianova E, Kahbatkyzy N, Flegontov P, Zaporozhchenko V, Immel A, Wang CC, Ixan O, Khussainova E, Bekmanov B, Zaibert V, Lavryashina M, Pocheshkhova E, Yusupov Y, Agdzhoyan A, Koshel S, Bukin A, Nymadawa P, Turdikulova S, et al. 2019. The genetic history of admixture across inner Eurasia. *Nature Ecology & Evolution* **3**:966–976. DOI: https://doi.org/10.1038/s41559-019-0878-2, PMID: 31036896

Jones ER, Gonzalez-Fortes G, Connell S, Siska V, Eriksson A, Martiniano R, McLaughlin RL, Gallego Llorente M, Cassidy LM, Gamba C, Meshveliani T, Bar-Yosef O, Müller W, Belfer-Cohen A, Matskevich Z, Jakeli N, Higham TFG, Currat M, Lordkipanidze D, Hofreiter M, et al. 2015. Upper Palaeolithic genomes reveal deep roots of modern Eurasians. *Nature Communications* **6**:8912. DOI: https://doi.org/10.1038/ncomms9912, PMID: 26567969

Jones ER, Zarina G, Moiseyev V, Lightfoot E, Nigst PR, Manica A, Pinhasi R, Bradley DG. 2017. The Neolithic transition in the baltic was not driven by admixture with early European farmers. *Current Biology* **27**:576–582. DOI: https://doi.org/10.1016/j.cub.2016.12.060, PMID: 28162894

Jónsson H, Ginolhac A, Schubert M, Johnson PLF, Orlando L. 2013. mapDamage2.0: fast approximate Bayesian estimates of ancient DNA damage parameters. *Bioinformatics* **29**:1682–1684. DOI: https://doi.org/10.1093/bioinformatics/btt193, PMID: 23613487

Kelleher J, Etheridge AM, McVean G. 2016. Efficient coalescent simulation and genealogical analysis for large sample sizes. *PLOS Computational Biology* **12**:e1004842. DOI: https://doi.org/10.1371/journal.pcbi.1004842, PMID: 27145223

Kelleher J, Thornton KR, Ashander J, Ralph PL. 2018. Efficient pedigree recording for fast population genetics simulation. *PLOS Computational Biology* **14**:e1006581. DOI: https://doi.org/10.1371/journal.pcbi.1006581, PMID: 30383757

Keller A, Graefen A, Ball M, Matzas M, Boisguerin V, Maixner F, Leidinger P, Backes C, Khairat R, Forster M, Stade B, Franke A, Mayer J, Spangler J, McLaughlin S, Shah M, Lee C, Harkins TT, Sartori A, Moreno-Estrada A, et al. 2012. New insights into the Tyrolean Iceman's origin and phenotype as inferred by whole-genome sequencing. *Nature Communications* **3**:698. DOI: https://doi.org/10.1038/ncomms1701, PMID: 22426219

Kılınç GM, Omrak A, Özer F, Günther T, Büyükkarakaya AM, Bıçakçı E, Baird D, Dönerttaş HM, Ghalichi A, Yaka R, Koptekin D, Açan SC, Parvizi P, Krzewińska M, Daskalaki EA, Yüncü E, Dağtaş ND, Fairbairn A, Pearson J, Mustafaoğlu G, et al. 2016. The demographic development of the first farmers in Anatolia. *Current Biology* **26**:2659–2666. DOI: https://doi.org/10.1016/j.cub.2016.07.057, PMID: 27498567

Korneliussen TS, Albrechtsen A, Nielsen R. 2014. ANGSD: analysis of next generation sequencing data. BMC Bioinformatics **15**:356. DOI: https://doi.org/10.1186/s12859-014-0356-4, PMID: 25420514

Kovacevic L, Tambets K, Ilumäe AM, Kushniarevich A, Yunusbayev B, Solnik A, Bego T, Primorac D, Skaro V, Leskovac A, Jakovski Z, Drobnic K, Tolk HV, Kovacevic S, Rudan P, Metspalu E, Marjanovic D. 2014. Standing at the gateway to Europe--the genetic structure of Western balkan populations based on autosomal and haploid markers. *PLOS ONE* **9**:e105090. DOI: https://doi.org/10.1371/journal.pone.0105090, PMID: 25148043

Krzewińska M, Kjellström A, Günther T, Hedenstierna-Jonson C, Zachrisson T, Omrak A, Yaka R, Kılınç GM, Somel M, Sobrado V, Evans J, Knipper C, Jakobsson M, Storå J, Götherström A. 2018a. Genomic and strontium isotope variation reveal immigration patterns in a Viking Age town. *Current Biology* **28**:2730–2738. DOI: https://doi.org/10.1016/j.cub.2018.06.053, PMID: 30146150

Krzewińska M, Kılınç GM, Juras A, Koptekin D, Chyleński M, Nikitin AG, Shcherbakov N, Shuteleva I, Leonova T, Kraeva L, Sungatov FA, Sultanova AN, Potekhina I, Łukasik S, Krenz-Niedbała M, Dalén L, Sinika V, Jakobsson M, Storå J, Götherström A. 2018b. Ancient genomes suggest the eastern Pontic-Caspian steppe as the source of western Iron Age nomads. *Science Advances* **4**:eaat4457. DOI: https://doi.org/10.1126/sciadv.aat4457, PMID: 30417088

Lamnidis TC, Majander K, Jeong C, Salmela E, Wessman A, Moiseyev V, Khartanovich V, Balanovsky O, Ongyerth M, Weihmann A, Sajantila A, Kelso J, Pääbo S, Onkamo P, Haak W, Krause J, Schiffels S. 2018. Ancient Fennoscandian genomes reveal origin and spread of Siberian ancestry in Europe. *Nature Communications* **9**:5018. DOI: https://doi.org/10.1038/s41467-018-07483-5, PMID: 30479341

Lazaridis I, Patterson N, Mittnik A, Renaud G, Mallick S, Kirsanow K, Sudmant PH, Schraiber JG, Castellano S, Lipson M, Berger B, Economou C, Bollongino R, Fu Q, Bos KI, Nordenfelt S, Li H, de Filippo C, Prüfer K, Sawyer S, et al. 2014. Ancient human genomes suggest three ancestral populations for present-day Europeans. *Nature* **513**:409–413. DOI: https://doi.org/10.1038/nature13673, PMID: 25230663

Lazaridis I, Nadel D, Rollefson G, Merrett DC, Rohland N, Mallick S, Fernandes D, Novak M, Gamarra B, Sirak K, Connell S, Stewardson K, Harney E, Fu Q, Gonzalez-Fortes G, Jones ER, Roodenberg SA, Lengyel G, Bocquentin F, Gasparian B, et al. 2016. Genomic insights into the origin of farming in the ancient Near East. *Nature* **536**:419–424. DOI: https://doi.org/10.1038/nature19310, PMID: 27459054

Lazaridis I, Mittnik A, Patterson N, Mallick S, Rohland N, Pfremgle S, Furtwängler A, Peltzer A, Posth C, Vasilakis A, McGeorge PJP, Konsolaki-Yannopoulou E, Korres G, Martlew H, Michalodimitrakis M, Özsait M, Özsait N, Papathanasiou A, Richards M, Roodenberg SA, et al. 2017. Genetic origins of the Minoans and Mycenaean. *Nature* **548**:214–218. DOI: https://doi.org/10.1038/nature23310

Li H, Durbin R. 2009. Fast and accurate short read alignment with Burrows-Wheeler transform. *Bioinformatics* **25**:1754–1760. DOI: https://doi.org/10.1093/bioinformatics/btp324, PMID: 19451168
Linderholm A, Kilinç GM, Szczepanek A, Włodarczak P, Jarosz P, Belka Z, Dopieralska J, Werens K, Górski J, Mazurek M, Hozer M, Rybicka M, Ostrowski M, Bagińska J, Koman W, Rodríguez-Varela R, Storá J, Götherström A, Krzewińska M. 2020. Corded Ware cultural complexity uncovered using genomic and isotopic analysis from south-eastern Poland. *Scientific Reports* **10**:6885. DOI: https://doi.org/10.1038/s41598-020-63138-w, PMID: 32303690

Lipson M, Szécsényi-Nagy A, Mallick S, Pósa A, Stégmár B, Keerl V, Rohland N, Stewardson K, Ferry M, Michel M, Oppenheimer J, Broomandkhoshbacht N, Harney E, Nordenfelt S, Llamas B, Gusztáv Mende B, Köhler K, Oross K, Bondár M, Marton T, et al. 2017. Parallel palaeogenomic transects reveal complex genetic history of early European farmers. *Nature* **551**:368–372. DOI: https://doi.org/10.1038/nature24476, PMID: 29144465

Mallick S, Li H, Lipson M, Mathieson I, Gymrek M, Racimo F, Zhao M, Chennagiri N, Nordenfelt S, Tandon A, Skoglund P, Lazaridis I, Sankararaman S, Fu Q, Rohland N, Renaud G, Erlich Y, Willems T, Gallo C, Spence JP, et al. 2016. The simons genome diversity project: 300 genomes from 142 diverse populations. *Nature* **538**:201–206. DOI: https://doi.org/10.1038/nature18964, PMID: 27654912

Mallick S, Micco A, Mah M, Ringbauer H, Lazaridis I, Olalde I, Patterson N, Reich D. 2023. The Allen Ancient DNA Resource (AADR): a curated compendium of ancient human genomes. *bioRxiv*. DOI: https://doi.org/10.1101/2023.04.06.535797

Malmström H, Günther T, Svensson EM, Juras A, Fraser M, Munters AR, Pospieszny Ł, Törv M, Lindström J, Götherström A, Storå J, Jakobsson M. 2019. The genomic ancestry of the Scandinavian Battle Axe Culture people and their relation to the broader Corded Ware horizon. *Proceedings. Biological Sciences* **286**:20191528. DOI: https://doi.org/10.1098/rspb.2019.1528, PMID: 31594508

Margaryan A, Lawson DJ, Sikora M, Racimo F, Rasmussen S, Moltke I, Cassidy LM, Jørsboe E, Ingason A, Pedersen MW, Korneliussen T, Wilhelmson H, Buš MM, de Barros Damgaard P, Martiniano R, Renaud G, Bhérer C, Moreno-Mayar JV, Fotakis AK, Allen M, et al. 2020. Population genomics of the Viking world. *Nature* **585**:390–396. DOI: https://doi.org/10.1038/s41586-020-2688-8, PMID: 32939067

Martin M. 2011. Cutadapt removes adapter sequences from high-throughput sequencing reads. *EMBnet.Journal* **17**:10. DOI: https://doi.org/10.14806/ej.17.1.200

Martiniano R, Caffell A, Holst M, Hunter-Mann K, Montgomery J, Müldner G, McLaughlin RL, Teasdale MD, van Rheenen W, Veldink JH, van den Berg LH, Hardiman O, Carroll M, Roskams S, Oxley J, Morgan C, Thomas MG, Barnes I, McDonnell C, Collins MJ, et al. 2016. Genomic signals of migration and continuity in Britain before the Anglo-Saxons. *Nature Communications* **7**:10326. DOI: https://doi.org/10.1038/ ncomms10326, PMID: 26783717

Martiniano R, Cassidy LM, Ó'Maoldúin R, McLaughlin R, Silva NM, Manco L, Fidalgo D, Pereira T, Coelho MJ, Serra M, Burger J, Parreira R, Moran E, Valera AC, Porfirio E, Boaventura R, Silva AM, Bradley DG. 2017. The population genomics of archaeological transition in west Iberia: investigation of ancient substructure using imputation and haplotype-based methods. *PLOS Genetics* **13**:e1006852. DOI: https://doi.org/10.1371/journal. pgen.1006852, PMID: 28749934

Mathieson I, Lazaridis I, Rohland N, Mallick S, Patterson N, Roodenberg SA, Harney E, Stewardson K, Fernandes D, Novak M, Sirak K, Gamba C, Jones ER, Llamas B, Dryomov S, Pickrell J, Arsuaga JL, de Castro JMB, Carbonell E, Gerritsen F, et al. 2015. Genome-wide patterns of selection in 230 ancient Eurasians. *Nature* **528**:499–503. DOI: https://doi.org/10.1038/nature16152, PMID: 26595274

Mathieson I, Alpaslan-Roodenberg S, Posth C, Szécsényi-Nagy A, Rohland N, Mallick S, Olalde I, Broomandkhoshbacht N, Candilio F, Cheronet O, Fernandes D, Ferry M, Gamarra B, Fortes GG, Haak W, Harney E, Jones E, Keating D, Krause-Kyora B, Kucukkalipci I, et al. 2018. The genomic history of southeastern Europe. *Nature* **555**:197–203. DOI: https://doi.org/10.1038/nature25778, PMID: 29466330

Meyer M, Kircher M. 2010. Illumina sequencing library preparation for highly multiplexed target capture and sequencing. *Cold Spring Harbor Protocols* **2010**:pdb.prot5448. DOI: https://doi.org/10.1101/pdb.prot5448, PMID: 20516186

Mittnik A, Wang C-C, Pfrengle S, Daubaras M, Zariņa G, Hallgren F, Allmäe R, Khartanovich V, Moiseyev V, Törv M, Furtwängler A, Valtueña AA, Feldman M, Economou C, Oinonen M, Vasks A, Balanovska E, Reich D, Jankauskas R, Haak W, et al. 2018. Author correction: the genetic prehistory of the Baltic Sea region. *Nature Communications* **9**:1494. DOI: https://doi.org/10.1038/s41467-018-03872-y, PMID: 29643405

Mittnik A, Massy K, Knipper C, Wittenborn F, Friedrich R, Pfrengle S, Burri M, Carlichi-Witjes N, Deeg H, Furtwängler A, Harbeck M, von Heyking K, Kociumaka C, Kucukkalipci I, Lindauer S, Metz S, Staskiewicz A, Thiel A, Wahl J, Haak W, et al. 2019. Kinship-based social inequality in Bronze age Europe. *Science* **366**:731–734. DOI: https://doi.org/10.1126/science.aax6219, PMID: 31601705

Moots HM, Antonio M, Sawyer S, Spence JP, Oberreiter V, Weiß CL, Lucci M, Cherifi YMS, La Pastina F, Genchi F, Praxmeier E, Zagorc B, Cheronot O, Özdoğan KT, Demetz L, Amrani S, Candilio F, De Angelis D, Gasperetti G, Fernandes D, et al. 2022. A genetic history of continuity and mobility in the Iron age Central Mediterranean. *bioRxiv*. DOI: https://doi.org/10.1101/2022.03.13.483276

Morrison J, Knoblauch N, Marcus JH, Stephens M, He X. 2020. Publisher correction: Mendelian randomization accounting for correlated and uncorrelated pleiotropic effects using genome-wide summary statistics. *Nature Genetics* **52**:750. DOI: https://doi.org/10.1038/s41588-020-0655-9, PMID: 32472065

Narasimhan VM, Patterson N, Moorjani P, Rohland N, Bernardos R, Mallick S, Lazaridis I, Nakatsuka N, Olalde I, Lipson M, Kim AM, Olivieri LM, Coppa A, Vidale M, Mallory J, Moiseyev V, Kitov E, Monge J, Adamski N, Alex N, et al. 2019. The formation of human populations in South and Central Asia. *Science* **365**:eaat7487. DOI: https://doi.org/10.1126/science.aat7487, PMID: 31488661
Linderholm A, Kilinç GM, Szczepanek A, Włodarczak P, Jarosz P, Belka Z, Dopieralska J, Werens K, Górski J, Mazurek M, Hozer M, Rybicka M, Ostrowski M, Bagińska J, Koman W, Rodríguez-Varela R, Storá J, Götherström A, Krzewińska M. 2020. Corded Ware cultural complexity uncovered using genomic and isotopic analysis from south-eastern Poland. *Scientific Reports* **10**:6885. DOI: https://doi.org/10.1038/s41598-020-63138-w, PMID: 32303690

Lipson M, Szécsényi-Nagy A, Mallick S, Pósa A, Stégmár B, Keerl V, Rohland N, Stewardson K, Ferry M, Michel M, Oppenheimer J, Broomandkhoshbacht N, Harney E, Nordenfelt S, Llamas B, Gusztáv Mende B, Köhler K, Oross K, Bondár M, Marton T, et al. 2017. Parallel palaeogenomic transects reveal complex genetic history of early European farmers. *Nature* **551**:368–372. DOI: https://doi.org/10.1038/nature24476, PMID: 29144465

Mallick S, Li H, Lipson M, Mathieson I, Gymrek M, Racimo F, Zhao M, Chennagiri N, Nordenfelt S, Tandon A, Skoglund P, Lazaridis I, Sankararaman S, Fu Q, Rohland N, Renaud G, Erlich Y, Willems T, Gallo C, Spence JP, et al. 2016. The simons genome diversity project: 300 genomes from 142 diverse populations. *Nature* **538**:201–206. DOI: https://doi.org/10.1038/nature18964, PMID: 27654912

Mallick S, Micco A, Mah M, Ringbauer H, Lazaridis I, Olalde I, Patterson N, Reich D. 2023. The Allen Ancient DNA Resource (AADR): a curated compendium of ancient human genomes. *bioRxiv*. DOI: https://doi.org/10.1101/2023.04.06.535797

Malmström H, Günther T, Svensson EM, Juras A, Fraser M, Munters AR, Pospieszny Ł, Törv M, Lindström J, Götherström A, Storå J, Jakobsson M. 2019. The genomic ancestry of the Scandinavian Battle Axe Culture people and their relation to the broader Corded Ware horizon. *Proceedings. Biological Sciences* **286**:20191528. DOI: https://doi.org/10.1098/rspb.2019.1528, PMID: 31594508

Margaryan A, Lawson DJ, Sikora M, Racimo F, Rasmussen S, Moltke I, Cassidy LM, Jørsboe E, Ingason A, Pedersen MW, Korneliussen T, Wilhelmson H, Buš MM, de Barros Damgaard P, Martiniano R, Renaud G, Bhérer C, Moreno-Mayar JV, Fotakis AK, Allen M, et al. 2020. Population genomics of the Viking world. *Nature* **585**:390–396. DOI: https://doi.org/10.1038/s41586-020-2688-8, PMID: 32939067

Martin M. 2011. Cutadapt removes adapter sequences from high-throughput sequencing reads. *EMBnet.Journal* **17**:10. DOI: https://doi.org/10.14806/ej.17.1.200

Martiniano R, Caffell A, Holst M, Hunter-Mann K, Montgomery J, Müldner G, McLaughlin RL, Teasdale MD, van Rheenen W, Veldink JH, van den Berg LH, Hardiman O, Carroll M, Roskams S, Oxley J, Morgan C, Thomas MG, Barnes I, McDonnell C, Collins MJ, et al. 2016. Genomic signals of migration and continuity in Britain before the Anglo-Saxons. *Nature Communications* **7**:10326. DOI: https://doi.org/10.1038/ ncomms10326, PMID: 26783717

Martiniano R, Cassidy LM, Ó'Maoldúin R, McLaughlin R, Silva NM, Manco L, Fidalgo D, Pereira T, Coelho MJ, Serra M, Burger J, Parreira R, Moran E, Valera AC, Porfirio E, Boaventura R, Silva AM, Bradley DG. 2017. The population genomics of archaeological transition in west Iberia: investigation of ancient substructure using imputation and haplotype-based methods. *PLOS Genetics* **13**:e1006852. DOI: https://doi.org/10.1371/journal. pgen.1006852, PMID: 28749934

Mathieson I, Lazaridis I, Rohland N, Mallick S, Patterson N, Roodenberg SA, Harney E, Stewardson K, Fernandes D, Novak M, Sirak K, Gamba C, Jones ER, Llamas B, Dryomov S, Pickrell J, Arsuaga JL, de Castro JMB, Carbonell E, Gerritsen F, et al. 2015. Genome-wide patterns of selection in 230 ancient Eurasians. *Nature* **528**:499–503. DOI: https://doi.org/10.1038/nature16152, PMID: 26595274

Mathieson I, Alpaslan-Roodenberg S, Posth C, Szécsényi-Nagy A, Rohland N, Mallick S, Olalde I, Broomandkhoshbacht N, Candilio F, Cheronet O, Fernandes D, Ferry M, Gamarra B, Fortes GG, Haak W, Harney E, Jones E, Keating D, Krause-Kyora B, Kucukkalipci I, et al. 2018. The genomic history of southeastern Europe. *Nature* **555**:197–203. DOI: https://doi.org/10.1038/nature25778, PMID: 29466330

Meyer M, Kircher M. 2010. Illumina sequencing library preparation for highly multiplexed target capture and sequencing. *Cold Spring Harbor Protocols* **2010**:pdb.prot5448. DOI: https://doi.org/10.1101/pdb.prot5448, PMID: 20516186

Mittnik A, Wang C-C, Pfrengle S, Daubaras M, Zariņa G, Hallgren F, Allmäe R, Khartanovich V, Moiseyev V, Törv M, Furtwängler A, Valtueña AA, Feldman M, Economou C, Oinonen M, Vasks A, Balanovska E, Reich D, Jankauskas R, Haak W, et al. 2018. Author correction: the genetic prehistory of the Baltic Sea region. *Nature Communications* **9**:1494. DOI: https://doi.org/10.1038/s41467-018-03872-y, PMID: 29643405

Mittnik A, Massy K, Knipper C, Wittenborn F, Friedrich R, Pfrengle S, Burri M, Carlichi-Witjes N, Deeg H, Furtwängler A, Harbeck M, von Heyking K, Kociumaka C, Kucukkalipci I, Lindauer S, Metz S, Staskiewicz A, Thiel A, Wahl J, Haak W, et al. 2019. Kinship-based social inequality in Bronze age Europe. *Science* **366**:731–734. DOI: https://doi.org/10.1126/science.aax6219, PMID: 31601705

Moots HM, Antonio M, Sawyer S, Spence JP, Oberreiter V, Weiß CL, Lucci M, Cherifi YMS, La Pastina F, Genchi F, Praxmeier E, Zagorc B, Cheronot O, Özdağ KT, Demetz L, Amrani S, Candilio F, De Angelis D, Gasperetti G, Fernandes D, et al. 2022. A genetic history of continuity and mobility in the Iron age Central Mediterranean. *bioRxiv*. DOI: https://doi.org/10.1101/2022.03.13.483276

Morrison J, Knoblauch N, Marcus JH, Stephens M, He X. 2020. Publisher correction: Mendelian randomization accounting for correlated and uncorrelated pleiotropic effects using genome-wide summary statistics. *Nature Genetics* **52**:750. DOI: https://doi.org/10.1038/s41588-020-0655-9, PMID: 32472065

Narasimhan VM, Patterson N, Moorjani P, Rohland N, Bernardos R, Mallick S, Lazaridis I, Nakatsuka N, Olalde I, Lipson M, Kim AM, Olivieri LM, Coppa A, Vidale M, Mallory J, Moiseyev V, Kitov E, Monge J, Adamski N, Alex N, et al. 2019. The formation of human populations in South and Central Asia. *Science* **365**:eaat7487. DOI: https://doi.org/10.1126/science.aat7487, PMID: 31488661
Nikitin AG, Stadler P, Kotova N, Teschler-Nicola M, Price TD, Hoover J, Kennett DJ, Lazaridis I, Rohland N, Lipson M, Reich D. 2019. Interactions between earliest Linearbandkeramik farmers and central European hunter gatherers at the dawn of European Neolithization. *Scientific Reports* **9**:19544. DOI: https://doi.org/10.1038/s41598-019-56029-2, PMID: 31863024

Novembre J, Johnson T, Bryc K, Kutalik Z, Boyko AR, Auton A, Indap A, King KS, Bergmann S, Nelson MR, Stephens M, Bustamante CD. 2008. Genes mirror geography within Europe. *Nature* **456**:98–101. DOI: https://doi.org/10.1038/nature07331, PMID: 18758442

Olalde I, Allentoft ME, Sánchez-Quinto F, Santpere G, Chiang CWK, DeGiorgio M, Prado-Martinez J, Rodríguez JA, Rasmussen S, Quilez J, Ramírez O, Marigorta UM, Fernández-Callejo M, Prada ME, Encinas JMV, Nielsen R, Netea MG, Novembre J, Sturm RA, Sabeti P, et al. 2014. Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European. *Nature* **507**:225–228. DOI: https://doi.org/10.1038/nature12960, PMID: 24463515

Olalde I, Schroeder H, Sandoval-Velasco M, Vinner L, Lobón I, Ramirez O, Civit S, García Borja P, Salazar-García DC, Talamo S, María Fullola J, Xavier Oms F, Pedro M, Martínez P, Sanz M, Daura J, Zilhão J, Marquès-Bonet T, Gilbert MTP, Lalueza-Fox C. 2015. A common genetic origin for early farmers from Mediterranean Cardial and Central European LBK cultures. *Molecular Biology and Evolution* **32**:3132–3142. DOI: https://doi.org/10.1093/molbev/msv181, PMID: 26337550

Olalde I, Brace S, Allentoft ME, Armit I, Kristiansen K, Booth T, Rohland N, Mallick S, Szécsényi-Nagy A, Mittnik A, Altena E, Lipson M, Lazaridis I, Harper TK, Patterson N, Broomandkhoshbacht N, Diekmann Y, Faltyskova Z, Fernandes D, Ferry M, et al. 2018. Erratum: the beaker phenomenon and the genomic transformation of northwest Europe. *Nature* **555**:190–196. DOI: https://doi.org/10.1038/nature26164, PMID: 29565364

Olalde I, Mallick S, Patterson N, Rohland N, Villalba-Mouco V, Silva M, Dulias K, Edwards CJ, Gandini F, Pala M, Soares P, Ferrando-Bernal M, Adamski N, Broomandkhoshbacht N, Cheronet O, Culleton BJ, Fernandes D, Lawson AM, Mah M, Oppenheimer J, et al. 2019. The genomic history of the Iberian Peninsula over the past 8000 years. *Science* **363**:1230–1234. DOI: https://doi.org/10.1126/science.aav4040, PMID: 30872528

Oleson JP. 2009. *The Oxford Handbook of Engineering and Technology in the Classical World* Oxford University Press. DOI: https://doi.org/10.1093/oxfordhb/9780199734856.001.0001

Omrak A, Günther T, Valdiosera C, Svensson EM, Malmström H, Kiesewetter H, Aylward W, Storå J, Jakobsson M, Götherström A. 2016. Genomic evidence establishes anatolia as the source of the European Neolithic gene pool. *Current Biology* **26**:270–275. DOI: https://doi.org/10.1016/j.cub.2015.12.019, PMID: 26748850

O'Sullivan N, Posth C, Coia V, Schuenemann VJ, Price TD, Wahl J, Pinhasi R, Zink A, Krause J, Maixner F. 2018. Ancient genome-wide analyses infer kinship structure in an Early Medieval Alemannic graveyard. *Science Advances* **4**:eaaao1262. DOI: https://doi.org/10.1126/sciadv.aao1262, PMID: 30191172

Pagani L, Lawson DJ, Jagoda E, Mörseburg A, Eriksson A, Mitt M, Clemente F, Hudjashov G, DeGiorgio M, Saag L, Wall JD, Cardona A, Mägi R, Wilson Sayres MA, Kaewert S, Inchley C, Scheib CL, Järve M, Karmin M, Jacobs GS, et al. 2016. Genomic analyses inform on migration events during the peopling of Eurasia. *Nature* **538**:238–242. DOI: https://doi.org/10.1038/nature19792, PMID: 27654910

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**:1065–1093. DOI: https://doi.org/10.1534/genetics.112.145037, PMID: 22960212

Pinhasi R, Fernandes D, Sirak K, Novak M, Connell S, Alpaslan-Roodenberg S, Gerritsen F, Moiseyev V, Gromov A, Raczky P, Anders A, Pietrusewsky M, Rollefson G, Jovanovic M, Trinhoang H, Bar-Oz G, Oxenham M, Matsumura H, Hofreiter M. 2015. Optimal ancient DNA yields from the inner ear part of the human petrous bone. *PLOS ONE* **10**:e0129102. DOI: https://doi.org/10.1371/journal.pone.0129102, PMID: 26086078

Pinhasi R, Fernandes DM, Sirak K, Cheronet O. 2019. Isolating the human cochlea to generate bone powder for ancient DNA analysis. *Nature Protocols* **14**:1194–1205. DOI: https://doi.org/10.1038/s41596-019-0137-7, PMID: 30842617

Posth C, Zaro V, Spyrou MA, Vai S, Gneчи-Ruscone GA, Modi A, Peltzer A, Mötsch A, Nägele K, Vågene ÅJ, Nelson EA, Radzevičiūtė R, Freund C, Bondioli LM, Cappuccini L, Frenzel H, Pacciani E, Boschin F, Capecchi G, Martini I, et al. 2021. The origin and legacy of the Etruscans through a 2000-year archeogenomic time transect. *Science Advances* **7**:eabi7673. DOI: https://doi.org/10.1126/sciadv.abi7673, PMID: 34559560

Prüfer K, de Filippo C, Grote S, Mafessoni F, Korlević P, Hajdinjak M, Vernot B, Skov L, Hsieh P, Peyrégne S, Reher D, Hopfe C, Nagel S, Maricic T, Fu Q, Theunert C, Rogers R, Skoglund P, Chintalapati M, Dannemann M, et al. 2017. A high-coverage Neandertal genome from Vindija Cave in Croatia. *Science* **358**:655–658. DOI: https://doi.org/10.1126/science.aao1887, PMID: 28982794

Renaud G, Slon V, Duggan AT, Kelso J. 2015. Schmutzi: estimation of contamination and endogenous mitochondrial consensus calling for ancient DNA. *Genome Biology* **16**:224. DOI: https://doi.org/10.1186/s13059-015-0776-0, PMID: 26458810

Rivollat M, Jeong C, Schiffels S, Küçükkalıpçı İ, Pemonge MH, Rohrlach AB, Alt KW, Binder D, Friederich S, Ghesquière E, Gronenborn D, Laporte L, Lefranc P, Meller H, Réveillas H, Rosenstock E, Rottier S, Scarre C, Soler L, Wahl J, et al. 2020. Ancient genome-wide DNA from France highlights the complexity of interactions between Mesolithic hunter-gatherers and Neolithic farmers. *Science Advances* **6**:eaaaz5344. DOI: https://doi.org/10.1126/sciadv.aaz5344, PMID: 32523989
Nikitin AG, Stadler P, Kotova N, Teschler-Nicola M, Price TD, Hoover J, Kennett DJ, Lazaridis I, Rohland N, Lipson M, Reich D. 2019. Interactions between earliest Linearbandkeramik farmers and central European hunter gatherers at the dawn of European Neolithization. *Scientific Reports* **9**:19544. DOI: https://doi.org/10.1038/s41598-019-56029-2, PMID: 31863024

Novembre J, Johnson T, Bryc K, Kutalik Z, Boyko AR, Auton A, Indap A, King KS, Bergmann S, Nelson MR, Stephens M, Bustamante CD. 2008. Genes mirror geography within Europe. *Nature* **456**:98–101. DOI: https://doi.org/10.1038/nature07331, PMID: 18758442

Olalde I, Allentoft ME, Sánchez-Quinto F, Santpere G, Chiang CWK, DeGiorgio M, Prado-Martinez J, Rodríguez JA, Rasmussen S, Quilez J, Ramírez O, Marigorta UM, Fernández-Callejo M, Prada ME, Encinas JMV, Nielsen R, Netea MG, Novembre J, Sturm RA, Sabeti P, et al. 2014. Derived immune and ancestral pigmentation alleles in a 7,000-year-old Mesolithic European. *Nature* **507**:225–228. DOI: https://doi.org/10.1038/nature12960, PMID: 24463515

Olalde I, Schroeder H, Sandoval-Velasco M, Vinner L, Lobón I, Ramirez O, Civit S, García Borja P, Salazar-García DC, Talamo S, María Fullola J, Xavier Oms F, Pedro M, Martínez P, Sanz M, Daura J, Zilhão J, Marquès-Bonet T, Gilbert MTP, Lalueza-Fox C. 2015. A common genetic origin for early farmers from Mediterranean Cardial and Central European LBK cultures. *Molecular Biology and Evolution* **32**:3132–3142. DOI: https://doi.org/10.1093/molbev/msv181, PMID: 26337550

Olalde I, Brace S, Allentoft ME, Armit I, Kristiansen K, Booth T, Rohland N, Mallick S, Szécsényi-Nagy A, Mittnik A, Altena E, Lipson M, Lazaridis I, Harper TK, Patterson N, Broomandkhoshbacht N, Diekmann Y, Faltyskova Z, Fernandes D, Ferry M, et al. 2018. Erratum: the beaker phenomenon and the genomic transformation of northwest Europe. *Nature* **555**:190–196. DOI: https://doi.org/10.1038/nature26164, PMID: 29565364

Olalde I, Mallick S, Patterson N, Rohland N, Villalba-Mouco V, Silva M, Dulias K, Edwards CJ, Gandini F, Pala M, Soares P, Ferrando-Bernal M, Adamski N, Broomandkhoshbacht N, Cheronet O, Culleton BJ, Fernandes D, Lawson AM, Mah M, Oppenheimer J, et al. 2019. The genomic history of the Iberian Peninsula over the past 8000 years. *Science* **363**:1230–1234. DOI: https://doi.org/10.1126/science.aav4040, PMID: 30872528

Oleson JP. 2009. *The Oxford Handbook of Engineering and Technology in the Classical World* Oxford University Press. DOI: https://doi.org/10.1093/oxfordhb/9780199734856.001.0001

Omrak A, Günther T, Valdiosera C, Svensson EM, Malmström H, Kiesewetter H, Aylward W, Storå J, Jakobsson M, Götherström A. 2016. Genomic evidence establishes anatolia as the source of the European Neolithic gene pool. *Current Biology* **26**:270–275. DOI: https://doi.org/10.1016/j.cub.2015.12.019, PMID: 26748850

O'Sullivan N, Posth C, Coia V, Schuenemann VJ, Price TD, Wahl J, Pinhasi R, Zink A, Krause J, Maixner F. 2018. Ancient genome-wide analyses infer kinship structure in an Early Medieval Alemannic graveyard. *Science Advances* **4**:eaaao1262. DOI: https://doi.org/10.1126/sciadv.aao1262, PMID: 30191172

Pagani L, Lawson DJ, Jagoda E, Mörseburg A, Eriksson A, Mitt M, Clemente F, Hudjashov G, DeGiorgio M, Saag L, Wall JD, Cardona A, Mägi R, Wilson Sayres MA, Kaewert S, Inchley C, Scheib CL, Järve M, Karmin M, Jacobs GS, et al. 2016. Genomic analyses inform on migration events during the peopling of Eurasia. *Nature* **538**:238–242. DOI: https://doi.org/10.1038/nature19792, PMID: 27654910

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**:1065–1093. DOI: https://doi.org/10.1534/genetics.112.145037, PMID: 22960212

Pinhasi R, Fernandes D, Sirak K, Novak M, Connell S, Alpaslan-Roodenberg S, Gerritsen F, Moiseyev V, Gromov A, Raczky P, Anders A, Pietrusewsky M, Rollefson G, Jovanovic M, Trinhoang H, Bar-Oz G, Oxenham M, Matsumura H, Hofreiter M. 2015. Optimal ancient DNA yields from the inner ear part of the human petrous bone. *PLOS ONE* **10**:e0129102. DOI: https://doi.org/10.1371/journal.pone.0129102, PMID: 26086078

Pinhasi R, Fernandes DM, Sirak K, Cheronet O. 2019. Isolating the human cochlea to generate bone powder for ancient DNA analysis. *Nature Protocols* **14**:1194–1205. DOI: https://doi.org/10.1038/s41596-019-0137-7, PMID: 30842617

Posth C, Zaro V, Spyrou MA, Vai S, Gneчи-Ruscone GA, Modi A, Peltzer A, Mötsch A, Nägele K, Vågene ÅJ, Nelson EA, Radzevičiūtė R, Freund C, Bondioli LM, Cappuccini L, Frenzel H, Pacciani E, Boschin F, Capecchi G, Martini I, et al. 2021. The origin and legacy of the Etruscans through a 2000-year archeogenomic time transect. *Science Advances* **7**:eabi7673. DOI: https://doi.org/10.1126/sciadv.abi7673, PMID: 34559560

Prüfer K, de Filippo C, Grote S, Mafessoni F, Korlević P, Hajdinjak M, Vernot B, Skov L, Hsieh P, Peyrégne S, Reher D, Hopfe C, Nagel S, Maricic T, Fu Q, Theunert C, Rogers R, Skoglund P, Chintalapati M, Dannemann M, et al. 2017. A high-coverage Neandertal genome from Vindija Cave in Croatia. *Science* **358**:655–658. DOI: https://doi.org/10.1126/science.aao1887, PMID: 28982794

Renaud G, Slon V, Duggan AT, Kelso J. 2015. Schmutzi: estimation of contamination and endogenous mitochondrial consensus calling for ancient DNA. *Genome Biology* **16**:224. DOI: https://doi.org/10.1186/s13059-015-0776-0, PMID: 26458810

Rivollat M, Jeong C, Schiffels S, Küçükkalıpçı İ, Pemonge MH, Rohrlach AB, Alt KW, Binder D, Friederich S, Ghesquière E, Gronenborn D, Laporte L, Lefranc P, Meller H, Réveillas H, Rosenstock E, Rottier S, Scarre C, Soler L, Wahl J, et al. 2020. Ancient genome-wide DNA from France highlights the complexity of interactions between Mesolithic hunter-gatherers and Neolithic farmers. *Science Advances* **6**:eaaaz5344. DOI: https://doi.org/10.1126/sciadv.aaz5344, PMID: 32523989
Rodríguez-Varela R, Günther T, Krzewińska M, Storå J, Gillingwater TH, MacCallum M, Arsuaga JL, Dobney K, Valdiosera C, Jakobsson M, Göhtherström A, Girdland-Flink L. 2017. Genomic analyses of pre-European conquest human remains from the Canary Islands Reveal close affinity to modern North Africans. *Current Biology* **27**:3396–3402. DOI: https://doi.org/10.1016/j.cub.2017.09.059, PMID: 29107554

Rohland N, Hofreiter M. 2007. Ancient DNA extraction from bones and teeth. *Nature Protocols* **2**:1756–1762. DOI: https://doi.org/10.1038/nprot.2007.247, PMID: 17641642

Rohland N, Harney E, Mallick S, Nordenfelt S, Reich D. 2015. Partial uracil-DNA-glycosylase treatment for screening of ancient DNA. *Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences* **370**:20130624. DOI: https://doi.org/10.1098/rstb.2013.0624, PMID: 25487342

Roymans N, Derks T, Heeren S. 2020. Roman imperialism and the transformation of rural society in a frontier Province: diversifying the narrative. *Britannia* **51**:265–294. DOI: https://doi.org/10.1017/S0068113X20000148

Saag L, Varul L, Scheib CL, Stenderup J, Allentoft ME, Saag L, Pagani L, Reidla M, Tambets K, Metspalu E, Kriiska A, Willerslev E, Kivisild T, Metspalu M. 2017. Extensive farming in estonia started through a sex-biased migration from the Steppe. *Current Biology* **27**:2185–2193. DOI: https://doi.org/10.1016/j.cub.2017.06.022, PMID: 28712569

Saag L, Laneman M, Varul L, Malve M, Valk H, Razzak MA, Shirobokov IG, Khartanovich VI, Mikhaylova ER, Kushniarevich A, Scheib CL, Solnik A, Reisberg T, Parik J, Saag L, Metspalu E, Rootsi S, Montinaro F, Remm M, Mägi R, et al. 2019. The arrival of Siberian ancestry connecting the Eastern Baltic to Uralic speakers further east. *Current Biology* **29**:1701–1711. DOI: https://doi.org/10.1016/j.cub.2019.04.026, PMID: 31080083

Sánchez-Quinto F, Malmström H, Fraser M, Girdland-Flink L, Svensson EM, Simões LG, George R, Hollfelder N, Burenhult G, Noble G, Britton K, Talamo S, Curtis N, Brzobohata H, Sumberova R, Göhtherström A, Storå J, Jakobsson M. 2019. Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society. *PNAS* **116**:9469–9474. DOI: https://doi.org/10.1073/pnas.1818037116, PMID: 30988179

Saupe T, Montinaro F, Scaggion C, Carrara N, Kivisild T, D'Atanasio E, Hui R, Solnik A, Lebrasseur O, Larson G, Alessandri L, Arienzo I, De Angelis F, Rolfo MF, Skeates R, Silvestri L, Beckett J, Talamo S, Dolfini A, Miari M, et al. 2021. Ancient genomes reveal structural shifts after the arrival of Steppe-related ancestry in the Italian Peninsula. *Current Biology* **31**:2576–2591. DOI: https://doi.org/10.1016/j.cub.2021.04.022, PMID: 33974848

Scheidel W, Morris I, Saller RP. 2007. *The Cambridge Economic History of the Greco-Roman World* Cambridge: Cambridge University Press. DOI: https://doi.org/10.1017/CHOL9780521780537

Scheidel W. 2015. Orbis: the stanford geospatial network model of the Roman world. *SSRN Electronic Journal* **1**:2609654. DOI: https://doi.org/10.2139/ssrn.2609654

Scheidel W. 2019. *Escape from Rome Princeton University Press*. DOI: https://doi.org/10.1515/9780691198835

Schiffels S, Haak W, Paajanen P, Llamas B, Popescu E, Loe L, Clarke R, Lyons A, Mortimer R, Sayer D, Tyler-Smith C, Cooper A, Durbin R. 2016. Iron age and Anglo-Saxon genomes from East England reveal British migration history. *Nature Communications* **7**:10408. DOI: https://doi.org/10.1038/ncomms10408, PMID: 26783965

Schroeder H, Margaryan A, Szmyt M, Theulot B, Włodarczak P, Rasmussen S, Gopalakrishnan S, Szczepanek A, Konopka T, Jensen TZT, Witkowska B, Wilk S, Przybyła MM, Pospieszny Ł, Sjögren KG, Belka Z, Olsen J, Kristiansen K, Willerslev E, Frei KM, et al. 2019. Unraveling ancestry, kinship, and violence in a Late Neolithic mass grave. *PNAS* **116**:10705–10710. DOI: https://doi.org/10.1073/pnas.1820210116, PMID: 31061125

Schuenemann VJ, Peltzer A, Welte B, van Pelt WP, Molak M, Wang CC, Furtwängler A, Urban C, Reiter E, Nieselt K, Teßmann B, Francken M, Harvati K, Haak W, Schiffels S, Krause J. 2017. Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods. *Nature Communications* **8**:15694. DOI: https://doi.org/10.1038/ncomms15694, PMID: 28556824

Sikora M, Seguin-Orlando A, Sousa VC, Albrechtsen A, Korneliussen T, Ko A, Rasmussen S, Dupanloup I, Nigst PR, Bosch MD, Renaud G, Allentoft ME, Margaryan A, Vasilyev SV, Veselovskaya EV, Borutskaya SB, Deviese T, Comeskey D, Higham T, Manica A, et al. 2017. Ancient genomes show social and reproductive behavior of early upper Paleolithic foragers. *Science* **358**:659–662. DOI: https://doi.org/10.1126/science.aao1807, PMID: 28982795

Skoglund P, Malmström H, Raghavan M, Storå J, Hall P, Willerslev E, Gilbert MTP, Göhtherström A, Jakobsson M. 2012. Origins and genetic legacy of Neolithic farmers and hunter-gatherers in Europe. *Science* **336**:466–469. DOI: https://doi.org/10.1126/science.1216304, PMID: 22539720

Skoglund P, Malmström H, Omrak A, Raghavan M, Valdiosera C, Günther T, Hall P, Tambets K, Parik J, Sjögren KG, Apel J, Willerslev E, Storå J, Göhtherström A, Jakobsson M. 2014. Genomic diversity and admixture differs for Stone-Age Scandinavian foragers and farmers. *Science* **344**:747–750. DOI: https://doi.org/10.1126/science.1253448, PMID: 24762536

Skourtanioti E, Erdal YS, Frangipane M, Balossi Restelli F, Yener KA, Pinnock F, Matthiae P, Özbal R, Schoop U-D, Guliyev F, Akhundov T, Lyonnet B, Hammer EL, Nugent SE, Burri M, Neumann GU, Penske S, Ingman T, Akar M, Shafiq R, et al. 2020. Genomic history of Neolithic to Bronze Age Anatolia, Northern Levant, and Southern Caucasus. *Cell* **181**:1158–1175. DOI: https://doi.org/10.1016/j.cell.2020.04.044, PMID: 32470401

Symonds M. 2017. *Protecting the Roman Empire: Fortlets, Frontiers, and the Quest for Post-Conquest Security* Cambridge: Cambridge University Press. DOI: https://doi.org/10.1017/9781108377287

Tacoma L. 2016. Approaching migration in the early Roman EmpireMigration and mobility in the Early Roman Empire. *Brill* **1**:1–22. DOI: https://doi.org/10.1163/9789004307377

Unterländer M, Palstra F, Lazaridis I, Pilipenko A, Hofmanová Z, Groß M, Sell C, Blöcher J, Kirsanow K, Rohland N, Rieger B, Kaiser E, Schier W, Pozdniakov D, Khokhlov A, Georges M, Wilde S, Powell A, Heyer E,
Rodríguez-Varela R, Günther T, Krzewińska M, Storå J, Gillingwater TH, MacCallum M, Arsuaga JL, Dobney K, Valdiosera C, Jakobsson M, Göhtherström A, Girdland-Flink L. 2017. Genomic analyses of pre-European conquest human remains from the Canary Islands Reveal close affinity to modern North Africans. *Current Biology* **27**:3396–3402. DOI: https://doi.org/10.1016/j.cub.2017.09.059, PMID: 29107554

Rohland N, Hofreiter M. 2007. Ancient DNA extraction from bones and teeth. *Nature Protocols* **2**:1756–1762. DOI: https://doi.org/10.1038/nprot.2007.247, PMID: 17641642

Rohland N, Harney E, Mallick S, Nordenfelt S, Reich D. 2015. Partial uracil-DNA-glycosylase treatment for screening of ancient DNA. *Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences* **370**:20130624. DOI: https://doi.org/10.1098/rstb.2013.0624, PMID: 25487342

Roymans N, Derks T, Heeren S. 2020. Roman imperialism and the transformation of rural society in a frontier Province: diversifying the narrative. *Britannia* **51**:265–294. DOI: https://doi.org/10.1017/S0068113X20000148

Saag L, Varul L, Scheib CL, Stenderup J, Allentoft ME, Saag L, Pagani L, Reidla M, Tambets K, Metspalu E, Kriiska A, Willerslev E, Kivisild T, Metspalu M. 2017. Extensive farming in estonia started through a sex-biased migration from the Steppe. *Current Biology* **27**:2185–2193. DOI: https://doi.org/10.1016/j.cub.2017.06.022, PMID: 28712569

Saag L, Laneman M, Varul L, Malve M, Valk H, Razzak MA, Shirobokov IG, Khartanovich VI, Mikhaylova ER, Kushniarevich A, Scheib CL, Solnik A, Reisberg T, Parik J, Saag L, Metspalu E, Rootsi S, Montinaro F, Remm M, Mägi R, et al. 2019. The arrival of Siberian ancestry connecting the Eastern Baltic to Uralic speakers further east. *Current Biology* **29**:1701–1711. DOI: https://doi.org/10.1016/j.cub.2019.04.026, PMID: 31080083

Sánchez-Quinto F, Malmström H, Fraser M, Girdland-Flink L, Svensson EM, Simões LG, George R, Hollfelder N, Burenhult G, Noble G, Britton K, Talamo S, Curtis N, Brzobohata H, Sumberova R, Göhtherström A, Storå J, Jakobsson M. 2019. Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society. *PNAS* **116**:9469–9474. DOI: https://doi.org/10.1073/pnas.1818037116, PMID: 30988179

Saupe T, Montinaro F, Scaggion C, Carrara N, Kivisild T, D'Atanasio E, Hui R, Solnik A, Lebrasseur O, Larson G, Alessandri L, Arienzo I, De Angelis F, Rolfo MF, Skeates R, Silvestri L, Beckett J, Talamo S, Dolfini A, Miari M, et al. 2021. Ancient genomes reveal structural shifts after the arrival of Steppe-related ancestry in the Italian Peninsula. *Current Biology* **31**:2576–2591. DOI: https://doi.org/10.1016/j.cub.2021.04.022, PMID: 33974848

Scheidel W, Morris I, Saller RP. 2007. *The Cambridge Economic History of the Greco-Roman World* Cambridge: Cambridge University Press. DOI: https://doi.org/10.1017/CHOL9780521780537

Scheidel W. 2015. Orbis: the stanford geospatial network model of the Roman world. *SSRN Electronic Journal* **1**:2609654. DOI: https://doi.org/10.2139/ssrn.2609654

Scheidel W. 2019. *Escape from Rome Princeton University Press*. DOI: https://doi.org/10.1515/9780691198835

Schiffels S, Haak W, Paajanen P, Llamas B, Popescu E, Loe L, Clarke R, Lyons A, Mortimer R, Sayer D, Tyler-Smith C, Cooper A, Durbin R. 2016. Iron age and Anglo-Saxon genomes from East England reveal British migration history. *Nature Communications* **7**:10408. DOI: https://doi.org/10.1038/ncomms10408, PMID: 26783965

Schroeder H, Margaryan A, Szmyt M, Theulot B, Włodarczak P, Rasmussen S, Gopalakrishnan S, Szczepanek A, Konopka T, Jensen TZT, Witkowska B, Wilk S, Przybyła MM, Pospieszny Ł, Sjögren KG, Belka Z, Olsen J, Kristiansen K, Willerslev E, Frei KM, et al. 2019. Unraveling ancestry, kinship, and violence in a Late Neolithic mass grave. *PNAS* **116**:10705–10710. DOI: https://doi.org/10.1073/pnas.1820210116, PMID: 31061125

Schuenemann VJ, Peltzer A, Welte B, van Pelt WP, Molak M, Wang CC, Furtwängler A, Urban C, Reiter E, Nieselt K, Teßmann B, Francken M, Harvati K, Haak W, Schiffels S, Krause J. 2017. Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods. *Nature Communications* **8**:15694. DOI: https://doi.org/10.1038/ncomms15694, PMID: 28556824

Sikora M, Seguin-Orlando A, Sousa VC, Albrechtsen A, Korneliussen T, Ko A, Rasmussen S, Dupanloup I, Nigst PR, Bosch MD, Renaud G, Allentoft ME, Margaryan A, Vasilyev SV, Veselovskaya EV, Borutskaya SB, Deviese T, Comeskey D, Higham T, Manica A, et al. 2017. Ancient genomes show social and reproductive behavior of early upper Paleolithic foragers. *Science* **358**:659–662. DOI: https://doi.org/10.1126/science.aao1807, PMID: 28982795

Skoglund P, Malmström H, Raghavan M, Storå J, Hall P, Willerslev E, Gilbert MTP, Göhtherström A, Jakobsson M. 2012. Origins and genetic legacy of Neolithic farmers and hunter-gatherers in Europe. *Science* **336**:466–469. DOI: https://doi.org/10.1126/science.1216304, PMID: 22539720

Skoglund P, Malmström H, Omrak A, Raghavan M, Valdiosera C, Günther T, Hall P, Tambets K, Parik J, Sjögren KG, Apel J, Willerslev E, Storå J, Göhtherström A, Jakobsson M. 2014. Genomic diversity and admixture differs for Stone-Age Scandinavian foragers and farmers. *Science* **344**:747–750. DOI: https://doi.org/10.1126/science.1253448, PMID: 24762536

Skourtanioti E, Erdal YS, Frangipane M, Balossi Restelli F, Yener KA, Pinnock F, Matthiae P, Özbal R, Schoop U-D, Guliyev F, Akhundov T, Lyonnet B, Hammer EL, Nugent SE, Burri M, Neumann GU, Penske S, Ingman T, Akar M, Shafiq R, et al. 2020. Genomic history of Neolithic to Bronze Age Anatolia, Northern Levant, and Southern Caucasus. *Cell* **181**:1158–1175. DOI: https://doi.org/10.1016/j.cell.2020.04.044, PMID: 32470401

Symonds M. 2017. *Protecting the Roman Empire: Fortlets, Frontiers, and the Quest for Post-Conquest Security* Cambridge: Cambridge University Press. DOI: https://doi.org/10.1017/9781108377287

Tacoma L. 2016. Approaching migration in the early Roman EmpireMigration and mobility in the Early Roman Empire. *Brill* **1**:1–22. DOI: https://doi.org/10.1163/9789004307377

Unterländer M, Palstra F, Lazaridis I, Pilipenko A, Hofmanová Z, Groß M, Sell C, Blöcher J, Kirsanow K, Rohland N, Rieger B, Kaiser E, Schier W, Pozdniakov D, Khokhlov A, Georges M, Wilde S, Powell A, Heyer E,
Currat M, et al. 2017. Ancestry and demography and descendants of Iron Age nomads of the Eurasian Steppe. *Nature Communications* **8**:14615. DOI: https://doi.org/10.1038/ncomms14615, PMID: 28256537

Valdiosera C, Günther T, Vera-Rodríguez JC, Ureña I, Iriarte E, Rodríguez-Varela R, Simões LG, Martínez-Sánchez RM, Svensson EM, Malmström H, Rodríguez L, Bermúdez de Castro JM, Carbonell E, Alday A, Hernández Vera JA, Göttherström A, Carretero JM, Arsuaga JL, Smith CI, Jakobsson M. 2018. Four millennia of Iberian biomolecular prehistory illustrate the impact of prehistoric migrations at the far end of Eurasia. *PNAS* **115**:3428–3433. DOI: https://doi.org/10.1073/pnas.1717762115, PMID: 29531053

van den Brink ECM, Beeri R, Kirzner D, Bron E, Cohen-Weinberger A, Kamaisky E, Gonen T, Gershuny L, Nagar Y, Ben-Tor D, Sukenik N, Shamir O, Maher EF, Reich D. 2017. A late bronze age II clay coffin from Tel shaddud in the Central Jezreel Valley, Israel: context and historical implications. *Levant* **49**:105–135. DOI: https://doi.org/10.1080/00758914.2017.1368204

Veeramah KR, Rott A, Groß M, van Dorp L, López S, Kirsanow K, Sell C, Blöcher J, Wegmann D, Link V, Hofmanová Z, Peters J, Trautmann B, Gairhos A, Haberstroh J, Päffgen B, Hellenthal G, Haas-Gebhard B, Harbeck M, Burger J. 2018. Population genomic analysis of elongated skulls reveals extensive female-biased immigration in early Medieval Bavaria. *PNAS* **115**:3494–3499. DOI: https://doi.org/10.1073/pnas.1719880115, PMID: 29531040

Verhagen P, Joyce J, Groenhuijzen MR. 2019. *Current Trends in Roman Demography and Empirical Approaches to the Dynamics of the Limes Populations Finding the Limits of the Limes* Cham: Springer. DOI: https://doi.org/10.1007/978-3-030-04576-0

Villalba-Mouco V, van de Loosdrecht MS, Posth C, Mora R, Martínez-Moreno J, Rojo-Guerra M, Salazar-García DC, Royo-Guillén JI, Kunst M, Rougier H, Crevecoeur I, Arcusa-Magallón H, Tejedor-Rodríguez C, García-Martínez de Lagrán I, Garrido-Pena R, Alt KW, Jeong C, Schiffels S, Utrilla P, Krause J, et al. 2019. Survival of late pleistocene hunter-gatherer ancestry in the Iberian Peninsula. *Current Biology* **29**:1169–1177. DOI: https://doi.org/10.1016/j.cub.2019.02.006, PMID: 30880015

Wang CC, Reinhold S, Kalmykov A, Wissgott A, Brandt G, Jeong C, Cheronet O, Ferry M, Harney E, Keating D, Mallick S, Rohland N, Stewardson K, Kantorovich AR, Maslov VE, Petrenko VG, Erlikh VR, Atabiev BC, Magomedov RG, Kohl PL, et al. 2019. Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions. *Nature Communications* **10**:590. DOI: https://doi.org/10.1038/s41467-018-08220-8, PMID: 30713341

Zalloua P, Collins CJ, Gosling A, Biagini SA, Costa B, Kardailsky O, Nigro L, Khalil W, Calafell F, Matisoo-Smith E. 2018. Ancient DNA of phoenician remains indicates discontinuity in the settlement history of Ibiza. *Scientific Reports* **8**:17567. DOI: https://doi.org/10.1038/s41598-018-35667-y, PMID: 30514893

Žegarac A, Winkelbach L, Blöcher J, Diekmann Y, Krečković Gavrilović M, Porčić M, Stojković B, Milašinović L, Schreiber M, Wegmann D, Veeramah KR, Stefanović S, Burger J. 2021. Ancient genomes provide insights into family structure and the heredity of social status in the early Bronze age of southeastern Europe. *Scientific Reports* **11**:10072. DOI: https://doi.org/10.1038/s41598-021-89090-x, PMID: 33980902
Currat M, et al. 2017. Ancestry and demography and descendants of Iron Age nomads of the Eurasian Steppe. *Nature Communications* **8**:14615. DOI: https://doi.org/10.1038/ncomms14615, PMID: 28256537

Valdiosera C, Günther T, Vera-Rodríguez JC, Ureña I, Iriarte E, Rodríguez-Varela R, Simões LG, Martínez-Sánchez RM, Svensson EM, Malmström H, Rodríguez L, Bermúdez de Castro JM, Carbonell E, Alday A, Hernández Vera JA, Göttherström A, Carretero JM, Arsuaga JL, Smith CI, Jakobsson M. 2018. Four millennia of Iberian biomolecular prehistory illustrate the impact of prehistoric migrations at the far end of Eurasia. *PNAS* **115**:3428–3433. DOI: https://doi.org/10.1073/pnas.1717762115, PMID: 29531053

van den Brink ECM, Beeri R, Kirzner D, Bron E, Cohen-Weinberger A, Kamaisky E, Gonen T, Gershuny L, Nagar Y, Ben-Tor D, Sukenik N, Shamir O, Maher EF, Reich D. 2017. A late bronze age II clay coffin from Tel shaddud in the Central Jezreel Valley, Israel: context and historical implications. *Levant* **49**:105–135. DOI: https://doi.org/10.1080/00758914.2017.1368204

Veeramah KR, Rott A, Groß M, van Dorp L, López S, Kirsanow K, Sell C, Blöcher J, Wegmann D, Link V, Hofmanová Z, Peters J, Trautmann B, Gairhos A, Haberstroh J, Päffgen B, Hellenthal G, Haas-Gebhard B, Harbeck M, Burger J. 2018. Population genomic analysis of elongated skulls reveals extensive female-biased immigration in early Medieval Bavaria. *PNAS* **115**:3494–3499. DOI: https://doi.org/10.1073/pnas.1719880115, PMID: 29531040

Verhagen P, Joyce J, Groenhuijzen MR. 2019. *Current Trends in Roman Demography and Empirical Approaches to the Dynamics of the Limes Populations Finding the Limits of the Limes* Cham: Springer. DOI: https://doi.org/10.1007/978-3-030-04576-0

Villalba-Mouco V, van de Loosdrecht MS, Posth C, Mora R, Martínez-Moreno J, Rojo-Guerra M, Salazar-García DC, Royo-Guillén JI, Kunst M, Rougier H, Crevecoeur I, Arcusa-Magallón H, Tejedor-Rodríguez C, García-Martínez de Lagrán I, Garrido-Pena R, Alt KW, Jeong C, Schiffels S, Utrilla P, Krause J, et al. 2019. Survival of late pleistocene hunter-gatherer ancestry in the Iberian Peninsula. *Current Biology* **29**:1169–1177. DOI: https://doi.org/10.1016/j.cub.2019.02.006, PMID: 30880015

Wang CC, Reinhold S, Kalmykov A, Wissgott A, Brandt G, Jeong C, Cheronet O, Ferry M, Harney E, Keating D, Mallick S, Rohland N, Stewardson K, Kantorovich AR, Maslov VE, Petrenko VG, Erlikh VR, Atabiev BC, Magomedov RG, Kohl PL, et al. 2019. Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions. *Nature Communications* **10**:590. DOI: https://doi.org/10.1038/s41467-018-08220-8, PMID: 30713341

Zalloua P, Collins CJ, Gosling A, Biagini SA, Costa B, Kardailsky O, Nigro L, Khalil W, Calafell F, Matisoo-Smith E. 2018. Ancient DNA of phoenician remains indicates discontinuity in the settlement history of Ibiza. *Scientific Reports* **8**:17567. DOI: https://doi.org/10.1038/s41598-018-35667-y, PMID: 30514893

Žegarac A, Winkelbach L, Blöcher J, Diekmann Y, Krečković Gavrilović M, Porčić M, Stojković B, Milašinović L, Schreiber M, Wegmann D, Veeramah KR, Stefanović S, Burger J. 2021. Ancient genomes provide insights into family structure and the heredity of social status in the early Bronze age of southeastern Europe. *Scientific Reports* **11**:10072. DOI: https://doi.org/10.1038/s41598-021-89090-x, PMID: 33980902
**Appendix 1**

For each region, we show the results from pairwise individual-based qpAdm analysis in the form of heatmaps, annotated by dendrograms from the hierarchical clustering analysis. The heatmap color scheme is chosen to signify non-rejected hypotheses (above 0.05) in shades of red and orange, and rejected hypotheses in shades of gray and yellow.

Columns of the heatmaps are annotated by time period: Copper Age (CA), Bronze Age (BA), Iron Age (IA), Imperial Rome & Late Antiquity (IRLA), Middle Ages & Early Modern (MAEM).

Rows of the heatmaps are annotated by cluster assignment based on the hierarchical clustering (see Methods). Clusters identified as outliers through our pipeline downstream are also annotated on the rows. Please note that outlier detection was only performed for individuals from historical periods (IA, IRLA, MAEM).

<figure><img src="image_13.png" /><figcaption>Appendix 1—figure 1. Armenia.</figcaption></figure>
# Appendix 1

For each region, we show the results from pairwise individual-based qpAdm analysis in the form of heatmaps, annotated by dendrograms from the hierarchical clustering analysis. The heatmap color scheme is chosen to signify non-rejected hypotheses (above 0.05) in shades of red and orange, and rejected hypotheses in shades of gray and yellow.

Columns of the heatmaps are annotated by time period: Copper Age (CA), Bronze Age (BA), Iron Age (IA), Imperial Rome & Late Antiquity (IRLA), Middle Ages & Early Modern (MAEM).

Rows of the heatmaps are annotated by cluster assignment based on the hierarchical clustering (see Methods). Clusters identified as outliers through our pipeline downstream are also annotated on the rows. Please note that outlier detection was only performed for individuals from historical periods (IA, IRLA, MAEM).

<figure><img src="image_12.png" /><figcaption>**Appendix 1—figure 1.** Armenia.</figcaption></figure>
<figure><img src="image_14.png" /><figcaption>Appendix 1—figure 2. Mainland Italy.</figcaption></figure>
<figure><img src="image_13.png" /><figcaption>Appendix 1—figure 2. Mainland Italy.</figcaption></figure>
<figure><img src="image_15.png" /><figcaption>Appendix 1—figure 3. Sardinia.</figcaption></figure>
<figure><img src="image_14.png" /><figcaption>Appendix 1—figure 3. Sardinia.</figcaption></figure>
<figure><img src="image_16.png" /><figcaption>**Appendix 1—figure 4.** Levant & Egypt.</figcaption></figure>
<figure><img src="image_15.png" /><figcaption>**Appendix 1—figure 4.** Levant & Egypt.</figcaption></figure>
<figure><img src="image_17.png" /><figcaption>**Appendix 1—figure 5.** Southeastern Europe.</figcaption></figure>
<figure><img src="image_16.png" /><figcaption>**Appendix 1—figure 5.** Southeastern Europe.</figcaption></figure>
<figure><img src="image_18.png" /><figcaption>**Appendix 1—figure 6.** Southwestern Europe.</figcaption></figure>
<figure><img src="image_17.png" /><figcaption>**Appendix 1—figure 6.** Southwestern Europe.</figcaption></figure>
<figure><img src="image_19.png" /><figcaption>**Appendix 1—figure 7.** Western Europe.</figcaption></figure>
<figure><img src="image_18.png" /><figcaption>**Appendix 1—figure 7.** Western Europe.</figcaption></figure>
<figure><img src="image_20.png" /><figcaption>**Appendix 1—figure 8.** Northern Europe.</figcaption></figure>
<figure><img src="image_19.png" /><figcaption>**Appendix 1—figure 8.** Northern Europe.</figcaption></figure>
<figure><img src="image_21.png" /><figcaption>**Appendix 1—figure 9.** Great Britain & Ireland.</figcaption></figure>
<figure><img src="image_20.png" /><figcaption>**Appendix 1—figure 9.** Great Britain & Ireland.</figcaption></figure>
<figure><img src="image_22.png" /><figcaption>**Appendix 1—figure 10.** Eastern Central Europe.</figcaption></figure>
<figure><img src="image_21.png" /><figcaption>**Appendix 1—figure 10.** Eastern Central Europe.</figcaption></figure>
<figure><img src="image_23.png" /><figcaption>**Appendix 1—figure 11.** Eastern Europe & Steppe.</figcaption></figure>
<figure><img src="image_22.png" /><figcaption>**Appendix 1—figure 11.** Eastern Europe & Steppe.</figcaption></figure>
<figure><img src="image_24.png" /><figcaption>Appendix 1—figure 12. North Africa.</figcaption></figure>
<figure><img src="image_23.png" /><figcaption>Appendix 1—figure 12. North Africa.</figcaption></figure>