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A genetic history of continuity and mobility in the Iron Age central Mediterranean
- Hannah M. Moots ORCID:orcid.org/0000-0002-6637-63211,2 nAff22,
- Margaret Antonio3 na1,
- Susanna Sawyer4,5 na1,
- Jeffrey P. Spence ORCID:orcid.org/0000-0002-3199-14476 na1,
- Victoria Oberreiter4 na1 nAff23,
- Clemens L. Weiß6 na1,
- Michaela Lucci7 na1,
- Yahia Mehdi Seddik Cherifi ORCID:orcid.org/0000-0003-4533-57044,8,9 na1,
- Francesco La Pastina7,
- Francesco Genchi7,10,
- Elisa Praxmeier4,
- Brina Zagorc ORCID:orcid.org/0000-0002-7685-19584 nAff23,
- Olivia Cheronet4 nAff23,
- Kadir T. Özdoğan11,
- Lea Demetz4,
- Selma Amrani12,
- Francesca Candilio ORCID:orcid.org/0000-0002-4668-136113,
- Daniela De Angelis14,
- Gabriella Gasperetti15,
- Daniel Fernandes ORCID:orcid.org/0000-0002-7434-65524,16,
- Ziyue Gao ORCID:orcid.org/0000-0001-9244-023817 na2,
- Mounir Fantar18 na2,
- Alfredo Coppa19,20 na2,
- Jonathan K. Pritchard ORCID:orcid.org/0000-0002-8828-52366,21 na2 &
- …
- Ron Pinhasi ORCID:orcid.org/0000-0003-1629-813122 na2 nAff4
Nature Ecology & Evolutionvolume 7, pages1515–1524 (2023)Cite this article
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Abstract
The Iron Age was a dynamic period in central Mediterranean history, with the expansion of Greek and Phoenician colonies and the growth of Carthage into the dominant maritime power of the Mediterranean. These events were facilitated by the ease of long-distance travel following major advances in seafaring. We know from the archaeological record that trade goods and materials were moving across great distances in unprecedented quantities, but it is unclear how these patterns correlate with human mobility. Here, to investigate population mobility and interactions directly, we sequenced the genomes of 30 ancient individuals from coastal cities around the central Mediterranean, in Tunisia, Sardinia and central Italy. We observe a meaningful contribution of autochthonous populations, as well as highly heterogeneous ancestry including many individuals with non-local ancestries from other parts of the Mediterranean region. These results highlight both the role of local populations and the extreme interconnectedness of populations in the Iron Age Mediterranean. By studying these trans-Mediterranean neighbours together, we explore the complex interplay between local continuity and mobility that shaped the Iron Age societies of the central Mediterranean.
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Data availability
All tools and data needed to reproduce and evaluate the conclusions in this paper are presented in the main text and theSupplementary Information. Alignment files for the DNA sequences for all newly reported individual genomes are available at the European Nucleotide Archive (ENA) database under project accession numberPRJEB49419.
References
López-Ruiz, C. & Doak, B. R.The Oxford Handbook of the Phoenician and Punic Mediterranean (Oxford Univ. Press, 2019).
Quinn, J. C. & Vella, N. C.The Punic Mediterranean: Identities and Identification from Phoenician Settlement to Roman Rule (Cambridge Univ. Press, 2014).
Turfa, J. M.The Etruscan World (Routledge, 2014).
Benelli, E. inItaly and the West: Comparative Issues in Romanization (eds Keay, S. J. & Terrenato, N.) 7–16 (Oxford: Oxbow Books, 2001).
Broodbank, C.The Making of the Middle Sea: A History of the Mediterranean from the Beginning to the Emergence of the Classical World (Oxford Univ. Press, 2013).
Fernandes, D. M. et al. The spread of steppe and Iranian-related ancestry in the islands of the western Mediterranean.Nat. Ecol. Evol.4, 334–345 (2020).
Marcus, J. H. et al. Genetic history from the Middle Neolithic to present on the Mediterranean island of Sardinia.Nat. Commun.11, 939 (2020).
Antonio, M. L. et al. Ancient Rome: a genetic crossroads of Europe and the Mediterranean.Science366, 708–714 (2019).
Posth, C. et al. The origin and legacy of the Etruscans through a 2000-year archeogenomic time transect.Sci. Adv.7, eabi7673 (2021).
Reitsema, L. J. et al. The diverse genetic origins of a Classical period Greek army.Proc. Natl Acad. Sci. USA119, e2205272119 (2022).
Matisoo-Smith, E. A. et al. A European mitochondrial haplotype identified in ancient Phoenician remains from Carthage, North Africa.PLoS ONE11, e0155046 (2016).
Saupe, T. et al. Ancient genomes reveal structural shifts after the arrival of steppe-related ancestry in the Italian Peninsula.Curr. Biol.31, 2576–2591.e12 (2021).
Haber, M. et al. 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, 149–157 (2020).
Haber, M. et al. 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, 274–282 (2017).
Olalde, I. et al. The genomic history of the Iberian Peninsula over the past 8000 years.Science363, 1230–1234 (2019).
Zalloua, P. et al. Ancient DNA of Phoenician remains indicates discontinuity in the settlement history of Ibiza.Sci. Rep.8, 17567–17515 (2018).
Schuenemann, V. J. et al. Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods.Nat. Commun.8, 15694 (2017).
Villalba-Mouco, V. et al. Genomic transformation and social organization during the Copper Age–Bronze Age transition in southern Iberia.Sci. Adv.7, eabi7038 (2021).
Feldman, M. et al. Ancient DNA sheds light on the genetic origins of early Iron Age Philistines.Sci. Adv.5, eaax0061 (2019).
Agranat-Tamir, L. et al. The genomic history of the Bronze Age Southern Levant.Cell181, 1146–1157.e11 (2020).
Fregel, R. et al. Ancient genomes from North Africa evidence prehistoric migrations to the Maghreb from both the Levant and Europe.Proc. Natl Acad. Sci. USA115, 6774–6779 (2018).
Patterson, N., Price, A. L. & Reich, D. Population structure and eigenanalysis.PLoS Genet.2, e190 (2006).
Patterson, N. et al. Ancient admixture in human history.Genetics192, 1065–1093 (2012).
Harney, É., Patterson, N., Reich, D. & Wakeley, J. Assessing the performance of qpAdm: a statistical tool for studying population admixture.Genetics217, iyaa045 (2021).
Fantar, M. H. inFenicios y Territorio: Actas del II Seminario (ed. Prats, A. G.) 71–88 (Instituto Alicantino de Cultura Juan Gil-Albert Alfredo, 2000).
Fantar, M. H. Le cavalier marin de Kerkouane.Africa1, 19–32 (1988).
Fantar, M. H.Kerkouane: Cité Punique du Cap Bon (Tunisie) (Institut National d’Archéologie et d’Art, 1984).
Miles, R.Carthage Must Be Destroyed: The Rise and Fall of an Ancient Mediterranean Civilization (Allen Lane, 2010).
Fantar, M. Espaces culturels à Kerkouane.C. R. Acad. Sci. Inscr. Belles Lett.147, 817–824 (2003).
Fantar, M. H.Kerkouane: Une Cité Punique au Cap-Bon (Institut national d'archéologie et d'art, 1987).
Salas, A. et al. The making of the African mtDNA landscape.Am. J. Hum. Genet.71, 1082–1111 (2002).
Lazaridis, I. et al. Genetic origins of the minoans and mycenaeans.Nature548, 214–218 (2017).
Loogväli, E.-L. et al. Disuniting uniformity: a pied cladistic canvas of mtDNA haplogroup H in Eurasia.Mol. Biol. Evol.21, 2012–2021 (2004).
Skourtanioti, E. et al. Ancient DNA reveals admixture history and endogamy in the prehistoric Aegean.Nat. Ecol. Evol.7, 290–303 (2023).
Hunter, V. Review ofFamilies in Classical and Hellenistic Greece: Representations and Realities by Pomeroy, S. B.Phoenix 52, 395–398 (1998).
Gallego Llorente, M. et al. Ancient Ethiopian genome reveals extensive Eurasian admixture in eastern Africa.Science350, 820–822 (2015).
Fregel, R. et al. Mitogenomes illuminate the origin and migration patterns of the indigenous people of the Canary Islands.PLoS ONE14, e0209125 (2019).
Arauna, L. R. et al. Recent historical migrations have shaped the gene pool of Arabs and Berbers in North Africa.Mol. Biol. Evol.34, 318–329 (2017).
Matisoo-Smith, E. et al. Ancient mitogenomes of Phoenicians from Sardinia and Lebanon: a story of settlement, integration, and female mobility.PLoS ONE13, e0190169 (2018).
Modi, A. et al. Genetic structure and differentiation from early Bronze Age in the Mediterranean island of Sicily: insights from ancient mitochondrial genomes.Front. Genet.13, 945227 (2022).
Modi, A. et al. The mitogenome portrait of Umbria in Central Italy as depicted by contemporary inhabitants and pre-Roman remains.Sci. Rep.10, 10700 (2020).
Aneli, S. et al. The genetic origin of Daunians and the pan-Mediterranean southern Italian Iron Age context.Mol. Biol. Evol.39, msac014 (2022).
Serventi, P. et al. Iron Age Italic population genetics: the Piceni from Novilara (8th–7th century BC).Ann. Hum. Biol.45, 34–43 (2018).
Sarno, S. et al. Insights into Punic genetic signatures in the southern necropolis of Tharros (Sardinia).Ann. Hum. Biol.48, 247–259 (2021).
Moorjani, P. et al. The history of African gene flow into Southern Europeans, Levantines, and Jews.PLoS Genet.7, e1001373 (2011).
Koptekin, D. et al. Spatial and temporal heterogeneity in human mobility patterns in Holocene Southwest Asia and the East Mediterranean.Curr. Biol.33, 41–57.e15 (2023).
Antonio, M. L. et al. Stable population structure in Europe since the Iron Age, despite high mobility. Preprint atbioRxivhttps://doi.org/10.1101/2022.05.15.491973 (2022).
Botigué, L. R. et al. Gene flow from North Africa contributes to differential human genetic diversity in southern Europe.Proc. Natl Acad. Sci. USA110, 11791–11796 (2013).
Fentress, E. inThe Hellenistic West: Rethinking the Ancient Mediterranean (eds Prag, J. & Quinn, J.) 157–178 (Cambridge Univ. Press, 2013).
Sereno, P. C. et al. Lakeside cemeteries in the Sahara: 5000 years of holocene population and environmental change.PLoS ONE3, e2995 (2008).
Sommer, M. inMoney, Trade and Trade Routes in Pre-Islamic North Africa (eds Dowler, A. & Galvin, E. R.) 61–64 (British Museum, 2011).
Hérodote.Textes relatifs à l’histoire de l’Afrique du Nord. Hérodote [L. IV, ch. 168–199; l. II, ch. 31–33; l. IV, ch. 42–43], par Stéphane Gsell. [Texte, traduction, commentaire. Fragments d’Hécatée relatifs à la Libye.] (A. Jourdan, 1915).
Hodos, T. Colonial engagements in the global Mediterranean Iron Age.Camb. Archaeol. J.19, 221–241 (2009).
Whittaker, C. R. inImperialism in the Ancient World:The Cambridge University Research Seminar in Ancient History (eds Garnsey, P. & Whittaker, C.) 59–90 (Cambridge Univ. Press, 1979).
Freund, K. P. A long-term perspective on the exploitation of Lipari obsidian in central Mediterranean prehistory.Quat. Int.468, 109–120 (2018).
Pinhasi, R., Fernandes, D. M., Sirak, K. & Cheronet, O. Isolating the human cochlea to generate bone powder for ancient DNA analysis.Nat. Protoc.14, 1194–1205 (2019).
Pinhasi, R. et al. Optimal ancient DNA yields from the inner ear part of the human petrous bone.PLoS ONE10, e0129102 (2015).
Meyer, M. & Kircher, M. Illumina sequencing library preparation for highly multiplexed target capture and sequencing.Cold Spring Harb. Protoc.2010, pdb.prot5448 (2010).
Renaud, G., Slon, V., Duggan, A. T. & Kelso, J. Schmutzi: estimation of contamination and endogenous mitochondrial consensus calling for ancient DNA.Genome Biol.16, 224 (2015).
Allen Ancient DNA Resource: Downloadable Genotypes of Present-day and Ancient DNA Data (David Reich Lab, 2021);https://reich.hms.harvard.edu/allen-ancient-dna-resource-aadr-downloadable-genotypes-present-day-and-ancient-dna-data (2021).
Mallick, S. et al. The Allen Ancient DNA Resource (AADR): a curated compendium of ancient human genomes. Preprint atbioRxivhttps://doi.org/10.1101/2023.04.06.535797 (2023).
Kuhn, M., Manuel, J., Jakobsson, M. & Günther, T. Estimating genetic kin relationships in prehistoric populations.PLoS ONE13, e0195491 (2018).
Maier, R. & Patterson, N. admixtools2 (Github, 2022).
ESRI. World Basemap v2.World Topographic Map (ArcGIS Online, 2017);https://www.arcgis.com/home/item.html?id=7dc6cea0b1764a1f9af2e679f642f0f5
Acknowledgements
This project was partially supported by the Stanford Interdisciplinary Graduate Fellowship (H.M.M.) and grants from the Stanford Archaeology Centre (H.M.M.), the Europe Center Austria Exchange Program (H.M.M.) and the Stanford Anthropology Department (H.M.M.), a National Science Foundation Graduate Research Fellowship (M.A.), the Austrian Science Fund (FWF) M3108-G (S.S.), the Howard Hughes Medical Institute (J.K.P.), a grant from the National Institutes of Health RO1 HG011432 (C.L.W.), the Italian Ministry of Foreign Affairs and International Cooperation (A.C., M.L., F.C., F.L.P., F.G.), the Istituto Per l’Oriente CA Nallino (A.C., M.L., F.C., F.L.P., F.G.), and a Ministro dell’Istruzione, Università e Ricerca (MIUR) project grant via the International Association for the Mediterranean and Oriental Studies (A.C., M.L., F.C., F.L.P., F.G.). We thank all members of the Pritchard and Pinhasi labs for their thoughtful and valuable feedback; P. W. Faber and the University of Chicago Genomics Facility for providing sequencing services for the ancient genomes reported here.
Author information
Ron Pinhasi
Present address: Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria
Hannah M. Moots
Present address: Department of Human Genetics, University of Chicago, Chicago, IL, USA
Victoria Oberreiter, Brina Zagorc & Olivia Cheronet
Present address: Human Evolution and Archaeological Sciences, University of Vienna, Vienna, Austria
These authors contributed equally: Margaret Antonio, Susanna Sawyer, Jeffrey P. Spence, Victoria Oberreiter, Clemens L. Weiß, Michaela Lucci, Yahia Mehdi Seddik Cherifi.
These authors jointly supervised this work: Ziyue Gao, Mounir Fantar, Alfredo Coppa, Jonathan K. Pritchard, Ron Pinhasi.
Authors and Affiliations
Stanford Archaeology Center, Stanford University, Stanford, CA, USA
Hannah M. Moots
Department of Anthropology, Stanford University, Stanford, CA, USA
Hannah M. Moots
Biomedical Informatics Program, Stanford University, Stanford, CA, USA
Margaret Antonio
Department of Evolutionary Anthropology, University of Vienna, Vienna, Austria
Susanna Sawyer, Victoria Oberreiter, Yahia Mehdi Seddik Cherifi, Elisa Praxmeier, Brina Zagorc, Olivia Cheronet, Lea Demetz & Daniel Fernandes
Human Evolution and Archaeological Sciences, University of Vienna, Vienna, Austria
Susanna Sawyer
Department of Genetics, Stanford University, Stanford, CA, USA
Jeffrey P. Spence, Clemens L. Weiß & Jonathan K. Pritchard
Dipartimento di Biologia Ambientale, Sapienza Università di Roma, Rome, Italy
Michaela Lucci, Francesco La Pastina & Francesco Genchi
Cardiolo-Oncology Research Collaborative Group (CORCG), Faculty of Medicine, Benyoucef Benkhedda University, Algiers, Algeria
Yahia Mehdi Seddik Cherifi
Molecular Pathology, University Paul Sabatier Toulouse III, Toulouse, France
Yahia Mehdi Seddik Cherifi
Department of Oriental Studies, Sapienza University of Rome, Rome, Italy
Francesco Genchi
Department of History and Art History, Utrecht University, Utrecht, the Netherlands
Kadir T. Özdoğan
LBEIG, Population Genetics and Conservation Unit, Department of Cellular and Molecular Biology–Faculty of Biological Sciences, University of Sciences and Technology Houari Boumediene, Algiers, Algeria
Selma Amrani
Museo delle Civiltà, Italian Ministry of Culture, Rome, Italy
Francesca Candilio
Museo Nazionale Etrusco di Tarquinia, Direzione Generale Musei Lazio, Rome, Italy
Daniela De Angelis
Soprintendenza Archeologia, belle arti e paesaggio per le province di Sassari e Nuoro, Sassari, Italy
Gabriella Gasperetti
CIAS, Department of Life Sciences, University of Coimbra, Coimbra, Portugal
Daniel Fernandes
Department of Genetics, University of Pennsylvania, Perelman School of Medicine, Philadelphia, PA, USA
Ziyue Gao
Département des Monuments et des Sites Antiques–Institut National du Patrimoine INP, Tunis, Tunisia
Mounir Fantar
Department of Biology, Stanford University, Stanford, CA, USA
Alfredo Coppa
Dipartimento di Storia Antropologia Religioni Arte Spettacolo, Sapienza Università di Roma, Rome, Italy
Alfredo Coppa
Department of Genetics, Harvard Medical School, Cambridge, MA, USA
Jonathan K. Pritchard
Department of Human Genetics, University of Chicago, Chicago, IL, USA
Ron Pinhasi
- Hannah M. Moots
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- Margaret Antonio
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Contributions
R.P., J.K.P, A.C. and M.F. designed research; R.P., S.S., V.O., E.P., O.C., K.T.O., L.D., H.M.M. and D.F. performed and supervised laboratory work; A.C. and M.F. designed the collection strategy for archaeological material; M.F., A.C., M.L., F.L.P., F.G., F.C., D.D.A., G.G., H.M.M., Y.M.S.C. and S.A. assembled skeletal material and provided archaeological background; H.M.M., M.A., S.S., J.P.S., V.O., C.L.W., E.P., B.Z. and Z.G. curated and analysed data with input from J.K.P., R.P., A.C., D.F. and Y.M.S.C.; H.M.M., J.K.P., R.P., J.P.S., M.A., S.S., C.L.W., Y.M.S.C. and Z.G. wrote the paper with input from all collaborators.
Corresponding authors
Correspondence toJonathan K. Pritchard orRon Pinhasi.
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Nature Ecology & Evolution thanks Rosa Fregel, Josephine Crawley Quinn and the other, anonymous, reviewer(s) for their contribution to the peer review of this work.
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Supplementary information
Supplementary Information
Supplementary Figs. 1–15, Discussion/Text, Table 1, and captions for Datasets 1–5.
Supplementary Data 1
Newly reported ancient individuals.
Supplementary Data 2
Ancient individual genomes used in analyses.
Supplementary Data 3
AMS dating and isotope analysis results.
Supplementary Data 4
Admixture and cladistic analysis using qpAdm and qpWave.
Supplementary Data 5
Admixture analysis using qpAdm, including Levantine source populations.
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Moots, H.M., Antonio, M., Sawyer, S.et al. A genetic history of continuity and mobility in the Iron Age central Mediterranean.Nat Ecol Evol7, 1515–1524 (2023). https://doi.org/10.1038/s41559-023-02143-4
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