{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2025,11,20]],"date-time":"2025-11-20T18:49:46Z","timestamp":1763664586634,"version":"build-2065373602"},"reference-count":76,"publisher":"MDPI AG","issue":"6","license":[{"start":{"date-parts":[[2021,5,28]],"date-time":"2021-05-28T00:00:00Z","timestamp":1622160000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Genes"],"abstract":"<jats:p>The forced migration of millions of Africans during the Atlantic Slave Trade led to the emergence of new genetic and linguistic identities, thereby providing a unique opportunity to study the mechanisms giving rise to human biological and cultural variation. Here we focus on the archipelago of S\u00e3o Tom\u00e9 and Pr\u00edncipe in the Gulf of Guinea, which hosted one of the earliest plantation societies relying exclusively on slave labor. We analyze the genetic variation in 25 individuals from three communities who speak distinct creole languages (Forros, Principenses and Angolares), using genomic data from expanded exomes in combination with a contextual dataset from Europe and Africa, including newly generated data from 28 Bantu speakers from Angola. Our findings show that while all islanders display mixed contributions from the Gulf of Guinea and Angola, the Angolares are characterized by extreme genetic differentiation and inbreeding, consistent with an admixed maroon isolate. In line with a more prominent Bantu contribution to their creole language, we additionally found that a previously reported high-frequency Y-chromosome haplotype in the Angolares has a likely Angolan origin, suggesting that their genetic, linguistic and social characteristics were influenced by a small group of dominant men who achieved disproportionate reproductive success.<\/jats:p>","DOI":"10.3390\/genes12060833","type":"journal-article","created":{"date-parts":[[2021,5,28]],"date-time":"2021-05-28T11:33:20Z","timestamp":1622201600000},"page":"833","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":9,"title":["The Genes of Freedom: Genome-Wide Insights into Marronage, Admixture and Ethnogenesis in the Gulf of Guinea"],"prefix":"10.3390","volume":"12","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5912-1892","authenticated-orcid":false,"given":"Jo\u00e3o","family":"Almeida","sequence":"first","affiliation":[{"name":"CIBIO-Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Universidade do Porto, 4485-661 Vair\u00e3o, Portugal"},{"name":"CIIMAR\/CIMAR\u2014Interdisciplinary Centre of Marine and Environmental Research, University of Porto, 4450-208 Matosinhos, Portugal"}]},{"given":"Anne-Maria","family":"Fehn","sequence":"additional","affiliation":[{"name":"CIBIO-Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Universidade do Porto, 4485-661 Vair\u00e3o, Portugal"},{"name":"Department of Linguistic and Cultural Evolution, Max-Planck Institute for the Science of Human History, 07745 Jena, Germany"}]},{"given":"Margarida","family":"Ferreira","sequence":"additional","affiliation":[{"name":"CIBIO-Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Universidade do Porto, 4485-661 Vair\u00e3o, Portugal"},{"name":"Department of Medical Sciences, Institute of Biomedicine\u2014iBiMED, University of Aveiro, 3810-193 Aveiro, Portugal"}]},{"given":"Teresa","family":"Machado","sequence":"additional","affiliation":[{"name":"CIBIO-Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Universidade do Porto, 4485-661 Vair\u00e3o, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0763-9312","authenticated-orcid":false,"given":"Tjerk","family":"Hagemeijer","sequence":"additional","affiliation":[{"name":"Centro de Lingu\u00edstica da Universidade de Lisboa, 1600-214 Lisboa, Portugal"},{"name":"Faculdade de Letras, Universidade de Lisboa, 1600-214 Lisboa, Portugal"}]},{"given":"Jorge","family":"Rocha","sequence":"additional","affiliation":[{"name":"CIBIO-Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Universidade do Porto, 4485-661 Vair\u00e3o, Portugal"},{"name":"Departamento de Biologia, Faculdade de Ci\u00eancias, Universidade do Porto, 4169-007 Porto, Portugal"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7042-8948","authenticated-orcid":false,"given":"Magdalena","family":"Gay\u00e0-Vidal","sequence":"additional","affiliation":[{"name":"CIBIO-Centro de Investiga\u00e7\u00e3o em Biodiversidade e Recursos Gen\u00e9ticos, Universidade do Porto, 4485-661 Vair\u00e3o, Portugal"}]}],"member":"1968","published-online":{"date-parts":[[2021,5,28]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Curtin, P.D. (1998). The Rise and Fall of the Plantation Complex, Cambridge University Press.","DOI":"10.1017\/CBO9780511819414"},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Gelabert, P., Ferrando-Bernal, M., de-Dios, T., Mattorre, B., Campoy, E., Gorostiza, A., Patin, E., Gonz\u00e1lez-Mart\u00edn, A., and Lalueza-Fox, C. (2019). Genome-Wide Data from the Bubi of Bioko Island Clarifies the Atlantic Fringe of the Bantu Dispersal. BMC Genom., 20.","DOI":"10.1186\/s12864-019-5529-0"},{"key":"ref_3","unstructured":"Caldeira, A.M. (1999). Mulheres, Sexualidade e Casamento Em S\u00e3o Tom\u00e9 e Pr\u00edncipe (S\u00e9culos XV-XVIII), Edi\u00e7\u00f5es Cosmos."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"123","DOI":"10.14195\/0870-4112_3-4_6","article-title":"Do Ref\u00fagio Nos Picos da Ilha de S\u00e3o Tom\u00e9 \u00e0 Absor\u00e7\u00e3o Colonial: A Quest\u00e3o Dos Angolares [From the Refuge on the Peaks of S\u00e3o Tom\u00e9 Island to Colonial Absorption: The Issue of the Angolares]","volume":"4","author":"Caldeira","year":"2018","journal-title":"Biblos"},{"key":"ref_5","unstructured":"Tenreiro, F. (1961). A Ilha de S\u00e3o Tom\u00e9, Mem\u00f3rias da Junta de Investiga\u00e7\u00e3oes Cient\u00edficas do Ultramar."},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Klein, H.S. (1999). The Atlantic Slave Trade, Cambridge University Press.","DOI":"10.2307\/525264"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"L\u00f3pez, L.\u00c1., Gon\u00e7alves, P., and de Avelar, J.O. (2018). From Creoles to Portuguese. The Portuguese Language Continuum in Africa and Brazil, John Benjamins.","DOI":"10.1075\/ihll.20"},{"key":"ref_8","unstructured":"Havik, P.J., and Newitt, M. (2007). Castaways, Autochthons, or Maroons? The Debate on the Angolares of S\u00e3o Tom\u00e9 Island. Creole Societies in the Portuguese Colonial Empire, Bristol University Press."},{"key":"ref_9","first-page":"59","article-title":"Languages Portuguese-Based, Spanish-Based and French-Based Languages","volume":"Volume 2","author":"Michaelis","year":"2013","journal-title":"The Survey of Pidgin and Creole"},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Hagemeijer, T., and Zamora, A. (2016). Fa d\u2019Amb\u00f4: From Past to Present. Int. J. Sociol. Lang., 193\u2013209.","DOI":"10.1515\/ijsl-2016-0009"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1075\/jpcl.26.1.05hag","article-title":"The Gulf of Guinea Creoles: Genetic and Typological Relations","volume":"26","author":"Hagemeijer","year":"2011","journal-title":"J. Pidgin Creole Lang."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1163\/19589514-04901011","article-title":"Creole Languages and Genes: The Case of S\u00e3o Tom\u00e9 and Pr\u00edncipe","volume":"49","author":"Hagemeijer","year":"2019","journal-title":"Faits Lang."},{"key":"ref_13","unstructured":"Ryder, A. (1969). Benin and the Europeans 1485\u20131897, Longman."},{"key":"ref_14","unstructured":"Lorenzino, G. (1998). The Angolar Creole Portuguese of S\u00e3o Tom\u00e9: Its Grammar and Sociolinguistic History. [Ph.D. Thesis, City University of New York]."},{"key":"ref_15","first-page":"163","article-title":"L\u2019apport Lexical Bantou En Angolar","volume":"29","author":"Maurer","year":"1992","journal-title":"Afr. Arbeitspap."},{"key":"ref_16","unstructured":"Ferraz, L.I. (1979). The Creole of S\u00e3o Tom\u00e9, Witwatersrand University Press."},{"key":"ref_17","unstructured":"Livraria, J.O. (1953). Aventura e Rotina: Sugest\u00f5es de Uma Viagem \u00e0 Procura Das Constantes Portuguesas de Car\u00e1ter e A\u00e7\u00e3o, Topbooks."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1002\/ajpa.1330260508","article-title":"The Anthropological Genetics of the Black Caribs (Garifuna) of Central America and the Caribbean","volume":"26","author":"Crawford","year":"1983","journal-title":"Yearb. Phys. Anthropol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1002\/ajpa.20117","article-title":"Shipwrecks and Founder Effects: Divergent Demographic Histories Reflected in Caribbean MtDNA","volume":"128","author":"Salas","year":"2005","journal-title":"Am. J. Phys. Anthropol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"453","DOI":"10.2307\/3171955","article-title":"Quilombos on S\u00e3o Tom\u00e9, or in Search of Original Sources","volume":"23","author":"Vansina","year":"1996","journal-title":"Hist. Afr."},{"key":"ref_21","first-page":"177","article-title":"A Linguistic Appraisal of Angolar","volume":"Volume 2","author":"Ferraz","year":"1974","journal-title":"Memoriam Antonio Jorge Dias"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"134","DOI":"10.1086\/524762","article-title":"Human Microevolution and the Atlantic Slave Trade: A Case Study from S\u00e4o Tom\u00e9","volume":"49","author":"Coelho","year":"2008","journal-title":"Curr. Anthropol."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Coelho, M., Sequeira, F., Luiselli, D., Beleza, S., and Rocha, J. (2009). On the Edge of Bantu Expansions: MtDNA, y Chromosome and Lactase Persistence Genetic Variation in Southwestern Angola. BMC Evol. Biol.","DOI":"10.1186\/1471-2148-9-80"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1002\/ajpa.23378","article-title":"Matriclans Shape Populations: Insights from the Angolan Namib Desert into the Maternal Genetic History of Southern Africa","volume":"165","author":"Oliveira","year":"2018","journal-title":"Am. J. Phys. Anthropol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1353\/hub.2002.0036","article-title":"The Peopling of S\u00e3o Tom\u00e9 (Gulf of Guinea): Origins of Slave Settlers and Admixture with the Portuguese","volume":"74","author":"Seco","year":"2002","journal-title":"Hum. Biol."},{"key":"ref_26","unstructured":"1000 Genomes Project, Consortium, Auton, A., Brooks, L.D., Durbin, R.M., Garrison, E.P., Kang, H.M., Korbel, J.O., Marchini, J.L., McCarthy, S., and McVean, G.A. (2015). A Global Reference for Human Genetic Variation. Nature."},{"key":"ref_27","unstructured":"Andrews, S. (2021, May 27). FastQC: A Quality Control Tool for High Throughput Sequence Data; [Software]; 2010. Available online: http:\/\/www.bioinformatics.babraham.ac.uk\/projects\/fastqc."},{"key":"ref_28","unstructured":"Joshi, N., and Fass, J. (2021, May 27). Sickle: A Sliding-Window, Adaptive, Quality-Based Trimming Tool for FastQ Files (Version 1.33); [Software]; 2011. Available online: https:\/\/github.com\/najoshi\/sickle."},{"key":"ref_29","unstructured":"Li, H. (2013). Aligning Sequence Reads, Clone Sequences and Assembly Contigs with BWA-MEM. arxiv."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"2078","DOI":"10.1093\/bioinformatics\/btp352","article-title":"The Sequence Alignment\/Map Format and SAMtools","volume":"25","author":"Li","year":"2009","journal-title":"Bioinformatics"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"2987","DOI":"10.1093\/bioinformatics\/btr509","article-title":"A Statistical Framework for SNP Calling, Mutation Discovery, Association Mapping and Population Genetical Parameter Estimation from Sequencing Data","volume":"27","author":"Li","year":"2011","journal-title":"Bioinformatics"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1038\/ng.806","article-title":"A Framework for Variation Discovery and Genotyping Using Next-Generation DNA Sequencing Data","volume":"43","author":"Depristo","year":"2011","journal-title":"Nat. Genet."},{"key":"ref_33","doi-asserted-by":"crossref","unstructured":"Van der Auwera, G.A., Carneiro, M.O., Hartl, C., Poplin, R., del Angel, G., Levy-Moonshine, A., Jordan, T., Shakir, K., Roazen, D., and Thibault, J. (2013). From FastQ Data to High-Confidence Variant Calls: The Genome Analysis Toolkit Best Practices Pipeline. Curr. Protoc. Bioinforma., 43.","DOI":"10.1002\/0471250953.bi1110s43"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1297","DOI":"10.1101\/gr.107524.110","article-title":"The Genome Analysis Toolkit: A MapReduce Framework for Analyzing Next-Generation DNA Sequencing Data","volume":"20","author":"McKenna","year":"2010","journal-title":"Genome Res."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"2156","DOI":"10.1093\/bioinformatics\/btr330","article-title":"The Variant Call Format and VCFtools","volume":"27","author":"Danecek","year":"2011","journal-title":"Bioinformatics"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"2867","DOI":"10.1093\/bioinformatics\/btq559","article-title":"Robust Relationship Inference in Genome-Wide Association Studies","volume":"26","author":"Manichaikul","year":"2010","journal-title":"Bioinformatics"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1038\/ng.608","article-title":"Common SNPs Explain a Large Proportion of the Heritability for Human Height","volume":"42","author":"Yang","year":"2010","journal-title":"Nat. Genet."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"Lawson, D.J., Hellenthal, G., Myers, S., and Falush, D. (2012). Inference of Population Structure Using Dense Haplotype Data. PLoS Genet., 8.","DOI":"10.1371\/journal.pgen.1002453"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1084","DOI":"10.1086\/521987","article-title":"Rapid and Accurate Haplotype Phasing and Missing-Data Inference for Whole-Genome Association Studies by Use of Localized Haplotype Clustering","volume":"81","author":"Browning","year":"2007","journal-title":"Am. J. Hum. Genet."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"116","DOI":"10.1016\/j.ajhg.2015.11.020","article-title":"Genotype Imputation with Millions of Reference Samples","volume":"98","author":"Browning","year":"2016","journal-title":"Am. J. Hum. Genet."},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Leslie, S., Winney, B., Hellenthal, G., Davison, D., Boumertit, A., Day, T., Hutnik, K., Royrvik, E.C., Cunliffe, B., and Lawson, D.J. (2015). The Fine-Scale Genetic Structure of the British Population. Nature.","DOI":"10.1038\/nature14230"},{"key":"ref_42","doi-asserted-by":"crossref","unstructured":"Van Dorp, L., Balding, D., Myers, S., Pagani, L., Tyler-Smith, C., Bekele, E., Tarekegn, A., Thomas, M.G., Bradman, N., and Hellenthal, G. (2015). Evidence for a Common Origin of Blacksmiths and Cultivators in the Ethiopian Ari within the Last 4500 Years: Lessons for Clustering-Based Inference. PLoS Genet., 11.","DOI":"10.1371\/journal.pgen.1005397"},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Huson, D.H., and Bryant, D. (2006). Application of Phylogenetic Networks in Evolutionary Studies. Mol. Biol. Evol., 254\u2013267.","DOI":"10.1093\/molbev\/msj030"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1655","DOI":"10.1101\/gr.094052.109","article-title":"Fast Model-Based Estimation of Ancestry in Unrelated Individuals","volume":"19","author":"Alexander","year":"2009","journal-title":"Genome Res."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1086\/519795","article-title":"PLINK: A Tool Set for Whole-Genome Association and Population-Based Linkage Analyses","volume":"81","author":"Purcell","year":"2007","journal-title":"Am. J. Hum. Genet."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"2817","DOI":"10.1093\/bioinformatics\/btw327","article-title":"Pong: Fast Analysis and Visualization of Latent Clusters in Population Genetic Data","volume":"32","author":"Behr","year":"2016","journal-title":"Bioinformatics"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"583","DOI":"10.1093\/genetics\/132.2.583","article-title":"Estimation of Levels of Gene Flow from DNA Sequence Data","volume":"132","author":"Hudson","year":"1992","journal-title":"Genetics"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1038\/ng2116","article-title":"Measurement of the Human Allele Frequency Spectrum Demonstrates Greater Genetic Drift in East Asians than in Europeans","volume":"39","author":"Keinan","year":"2007","journal-title":"Nat. Genet."},{"key":"ref_49","unstructured":"Kolde, R. (2021, May 27). Pheatmap: Pretty Heatmaps; 2013. [Software]. Available online: http:\/\/CRAN.R-project.org\/package=pheatmap."},{"key":"ref_50","unstructured":"Bernstein, F. (1931). Die Geographische Verteilung der Blutgruppen Und Ihre Anthropologische Bedeutung. Comitato Italiano per lo Studio dei Problemi della Populazione, Instituto Poligrafico dello Stato."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"2529","DOI":"10.1016\/j.cub.2017.07.002","article-title":"Parallel Trajectories of Genetic and Linguistic Admixture in a Genetically Admixed Creole Population","volume":"27","author":"Verdu","year":"2017","journal-title":"Curr. Biol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"786","DOI":"10.1073\/pnas.0909559107","article-title":"Genome-Wide Patterns of Population Structure and Admixture in West Africans and African Americans","volume":"107","author":"Bryc","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1002\/humu.21382","article-title":"HaploGrep: A Fast and Reliable Algorithm for Automatic Classification of Mitochondrial DNA Haplogroups","volume":"32","author":"Pacher","year":"2011","journal-title":"Hum. Mutat."},{"key":"ref_54","first-page":"34","article-title":"Haplogroup Prediction from Y-STR Values Using a Bayesian-Allele- Frequency Approach","volume":"2","author":"Athey","year":"2006","journal-title":"J Genetic Genealogy"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"743","DOI":"10.1093\/genetics\/141.2.743","article-title":"Mitochondrial Portraits of Human Populations Using Median Networks","volume":"141","author":"Bandelt","year":"1995","journal-title":"Genetics"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1093\/oxfordjournals.molbev.a026036","article-title":"Median-Joining Networks for Inferring Intraspecific Phylogenies","volume":"16","author":"Bandelt","year":"1999","journal-title":"Mol. Biol. Evol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1177\/117693430500100003","article-title":"Arlequin (Version 3.0): An Integrated Software Package for Population Genetics Data Analysis","volume":"1","author":"Excoffier","year":"2005","journal-title":"Evol. Bioinforma."},{"key":"ref_58","first-page":"935","article-title":"Origin and Evolution of Native American MtDNA Variation: A Reappraisal","volume":"59","author":"Forster","year":"1996","journal-title":"Am. J. Hum. Genet."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Macaulay, V., Soares, P., and Richards, M.B. (2019). Rectifying Long-Standing Misconceptions about the \u03c1 Statistic for Molecular Dating. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0212311"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"520","DOI":"10.1002\/humu.20254","article-title":"Mutation Rates at Y Chromosome Specific Microsatellites","volume":"26","author":"Calafell","year":"2005","journal-title":"Hum. Mutat."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1413","DOI":"10.1038\/ejhg.2014.285","article-title":"Y-Chromosome Descent Clusters and Male Differential Reproductive Success: Young Lineage Expansions Dominate Asian Pastoral Nomadic Populations","volume":"23","author":"Balaresque","year":"2015","journal-title":"Eur. J. Hum. Genet."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1002\/ajpa.20188","article-title":"Cross-Cultural Estimation of the Human Generation Interval for Use in Genetics-Based Population Divergence Studies","volume":"128","author":"Fenner","year":"2005","journal-title":"Am. J. Phys. Anthropol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"543","DOI":"10.1126\/science.aal1988","article-title":"Dispersals and Genetic Adaptation of Bantu-Speaking Populations in Africa and North America","volume":"356","author":"Patin","year":"2017","journal-title":"Science"},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Semo, A., Gay\u00e0-Vidal, M., Fortes-Lima, C., Alard, B., Oliveira, S., Almeida, J., Prista, A., Damasceno, A., Fehn, A.-M., and Schlebusch, C. (2020). Along the Indian Ocean Coast: Genomic Variation in Mozambique Provides New Insights into the Bantu Expansion. Mol. Biol. Evol., 37.","DOI":"10.1101\/697474"},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Nielsen, R., Akey, J.M., Jakobsson, M., Pritchard, J.K., Tishkoff, S., and Willerslev, E. (2017). Tracing the Peopling of the World through Genomics. Nature, 302\u2013310.","DOI":"10.1038\/nature21347"},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"Ceballos, F.C., Joshi, P.K., Clark, D.W., Ramsay, M., and Wilson, J.F. (2018). Runs of Homozygosity: Windows into Population History and Trait Architecture. Nat. Rev. Genet., 220\u2013234.","DOI":"10.1038\/nrg.2017.109"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Kirin, M., McQuillan, R., Franklin, C.S., Campbell, H., Mckeigue, P.M., and Wilson, J.F. (2010). Genomic Runs of Homozygosity Record Population History and Consanguinity. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0013996"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1525\/aa.1956.58.1.02a00080","article-title":"Double Descent and Its Correlates among the Herero of Ngamiland","volume":"58","author":"Gibson","year":"1956","journal-title":"Am. Anthropol."},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Price, R. (1996). Maroon Societies. Rebel Slave Communities in the Americas, The John Hopkins University Press.","DOI":"10.56021\/9780801854965"},{"key":"ref_70","unstructured":"Seibert, G. (1999). Comrades, Clients, and Cousins. Colonialism, Socialism and Democratization in S\u00e3o Tom\u00e9 and Pr\u00edncipe, CNWS Publication."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1086\/367774","article-title":"The Genetic Legacy of the Mongols","volume":"72","author":"Zerjal","year":"2003","journal-title":"Am. J. Hum. Genet."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"11645","DOI":"10.1073\/pnas.0710158105","article-title":"Male Dominance Rarely Skews the Frequency Distribution of Y Chromosome Haplotypes in Human Populations","volume":"105","author":"Lansing","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1002\/ajpa.10180","article-title":"Hopi Indians, \u201cCultural\u201d Selection, and Albinism","volume":"121","author":"Hedrick","year":"2003","journal-title":"Am. J. Phys. Anthropol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"815","DOI":"10.1126\/science.170.3960.815","article-title":"Lessons from a \u201cPrimitive\u201d People","volume":"170","author":"Neel","year":"1970","journal-title":"Science"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1002\/ajpa.1330350206","article-title":"Demographic Structure of a Primitive Population: A Simulation","volume":"35","author":"MacCluer","year":"1971","journal-title":"Am. J. Phys. Anthropol."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Heyer, E., Sibert, A., and Austerlitz, F. (2005). Cultural Transmission of Fitness: Genes Take the Fast Lane. Trends Genet., 234\u2013239.","DOI":"10.1016\/j.tig.2005.02.007"}],"container-title":["Genes"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4425\/12\/6\/833\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T06:09:51Z","timestamp":1760162991000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4425\/12\/6\/833"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,5,28]]},"references-count":76,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2021,6]]}},"alternative-id":["genes12060833"],"URL":"https:\/\/doi.org\/10.3390\/genes12060833","relation":{},"ISSN":["2073-4425"],"issn-type":[{"type":"electronic","value":"2073-4425"}],"subject":[],"published":{"date-parts":[[2021,5,28]]}}}