Acknowledgements
The authors thank J. Víctor Moreno-Mayar for helpful suggestions, and
the ISOGG, YFull and FamilyTree DNA teams for their work curating the human Y -SNP tree
and making it publicly available for the research community. The Lundbeck Foundation
GeoGenetics Centre is supported by the Lundbeck Foundation (grant nos. R302-2018-2155,
R155-2013-16338), the Novo Nordisk Foundation (grant no. NNF18SA0035006), the
Wellcome Trust (grant no. UNS69906), Carlsberg Foundation (grant no . CF18 -0024), the
Danish National Research Foundation (grant nos. DNRF94, DNRF174), the University of
Copenhagen (KU2016 programme) . All authors were supported by the Novo Nordisk
Foundation and Wellcome Trust (AEGIS project) and the Danish National Research
Foundation Center for Ancient Environmental Genomics (CAEG).
Competing interests: The authors declare no competing interests.
.CC-BY-NC 4.0 International licenseavailable under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
The copyright holder for this preprintthis version posted November 20, 2025. ; https://doi.org/10.1101/2025.11.20.689455doi: bioRxiv preprint
References
1. Cortez, D. et al. Origins and functional evolution of Y chromosomes across mammals.
Nature 508, 488–493 (2014).
2. Hughes, J. F . & Page, D. C. The Biology and Evolution of Mammalian Y Chromosomes.
Annual Review of Genetics 49, 507–527 (2015).
3. Karmin, M. et al. A recent bottleneck of Y chromosome diversity coincides with a global
change in culture. Genome Res 25, 459–466 (2015).
4. Poznik, G. D. et al. Punctuated bursts in human male demography inferred from 1,244
worldwide Y-chromosome sequences. Nat Genet 48, 593–599 (2016).
5. Jobling, M. A. & Tyler-Smith, C. Human Y-chromosome variation in the genome-
sequencing era. Nat Rev Genet 18, 485–497 (2017).
6. Petr, M. et al. The evolutionary history of Neanderthal and Denisovan Y chromosomes.
Science 369, 1653–1656 (2020).
7. Taylor, W. T. T. et al. Early dispersal of domestic horses into the Great Plains and
northern Rockies. Science 379, 1316–1323 (2023).
8. Bozlak, E. et al. Refining the evolutionary tree of the horse Y chromosome. Sci Rep 13,
8954 (2023).
9. Underhill, P . A. et al. Y chromosome sequence variation and the history of human
populations. Nat Genet 26, 358–361 (2000).
10. Y-Chromosome Consortium. A Nomenclature System for the Tree of Human Y-
Chromosomal Binary Haplogroups. Genome Res. 12, 339–348 (2002).
.CC-BY-NC 4.0 International licenseavailable under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
The copyright holder for this preprintthis version posted November 20, 2025. ; https://doi.org/10.1101/2025.11.20.689455doi: bioRxiv preprint
11. Fowler, C. et al. A high-resolution picture of kinship practices in an Early Neolithic
tomb. Nature 1–4 (2021) doi:10.1038/s41586-021-04241-4.
12. Seersholm, F . V . et al. Repeated plague infections across six generations of Neolithic
Farmers. Nature 632, 114–121 (2024).
13. Poznik, G. D. Identifying Y-chromosome haplogroups in arbitrarily large samples of
sequenced or genotyped men. 088716 Preprint at https://doi.org/10.1101/088716
(2016).
14. Ralf, A., Montiel González, D., Zhong, K. & Kayser, M. Yleaf: Software for Human Y-
Chromosomal Haplogroup Inference from Next-Generation Sequencing Data. Mol Biol
Evol 35, 1291–1294 (2018).
15. Jagadeesan, A. et al. HaploGrouper: a generalized approach to haplogroup
classification. Bioinformatics 37, 570–572 (2021).
16. Martiniano, R., De Sanctis, B., Hallast, P . & Durbin, R. Placing Ancient DNA Sequences
into Reference Phylogenies. Mol Biol Evol 39, msac017 (2022).
17. Köster, J. & Rahmann, S. Snakemake—a scalable bioinformatics workflow engine.
Bioinformatics 28, 2520–2522 (2012).
18. Li, H. et al. The Sequence Alignment/Map format and SAMtools. Bioinformatics 25,
2078–2079 (2009).
19. Poznik, G. D. et al. Sequencing Y Chromosomes Resolves Discrepancy in Time to
Common Ancestor of Males versus Females. Science 341, 562–565 (2013).
20. Zhao, H. et al. CrossMap: a versatile tool for coordinate conversion between genome
assemblies. Bioinformatics 30, 1006–1007 (2014).
.CC-BY-NC 4.0 International licenseavailable under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
The copyright holder for this preprintthis version posted November 20, 2025. ; https://doi.org/10.1101/2025.11.20.689455doi: bioRxiv preprint
21. Danecek, P . et al. The variant call format and VCFtools. Bioinformatics 27, 2156–2158
(2011).
22. Briggs, A. W. et al. Patterns of damage in genomic DNA sequences from a Neandertal.
PNAS 104, 14616–14621 (2007).
23. Gansauge, M.-T. & Meyer, M. Single-stranded DNA library preparation for the
sequencing of ancient or damaged DNA. Nat Protoc 8, 737–748 (2013).
24. Huerta-Cepas, J., Serra, F . & Bork, P . ETE 3: Reconstruction, Analysis, and Visualization
of Phylogenomic Data. Mol Biol Evol 33, 1635–1638 (2016).
25. Czech, L., Barbera, P . & Stamatakis, A. Genesis and Gappa: processing, analyzing and
visualizing phylogenetic (placement) data. Bioinformatics 36, 3263–3265 (2020).
26. Antonio, M. L. et al. Ancient Rome: A genetic crossroads of Europe and the
Mediterranean. Science 366, 708–714 (2019).
27. de Barros Damgaard, P . et al. The first horse herders and the impact of early Bronze Age
steppe expansions into Asia. Science 360, eaar7711 (2018).
28. Cassidy, L. M. et al. A dynastic elite in monumental Neolithic society. Nature 582, 384–
388 (2020).
29. Felkel, S. et al. The horse Y chromosome as an informative marker for tracing sire lines.
Sci Rep 9, 6095 (2019).
30. Remer, V . et al. Y-Chromosomal Insights into Breeding History and Sire Line
Genealogies of Arabian Horses. Genes 13, 229 (2022).
.CC-BY-NC 4.0 International licenseavailable under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
The copyright holder for this preprintthis version posted November 20, 2025. ; https://doi.org/10.1101/2025.11.20.689455doi: bioRxiv preprint
31. Radovic, L. et al. The global spread of Oriental Horses in the past 1,500 years through
the lens of the Y chromosome. Proceedings of the National Academy of Sciences 121,
e2414408121 (2024).
.CC-BY-NC 4.0 International licenseavailable under a
(which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made
The copyright holder for this preprintthis version posted November 20, 2025. ; https://doi.org/10.1101/2025.11.20.689455doi: bioRxiv preprint