Spanish-sourced domestic horses into Indig-
enous societies across the plains before the
first half of the 17th century CE.
Indigenous societies incorporated horses before
the Pueblo Revolt
Of 33 early American equid specimens, we
successfully radiocarbon dated 29 and char-
acterized a total of 27 genetically, along with
six new specimens from Eurasia (producing
nine ancient genomes with an average depth-
of-coverage of 2.06× to 12.24×, with substan-
tial genome-wide sequence data for seven
additional horse specimens, 0.06× to 0.96×,
plus one donkey genome, 1.32×) (Fig. 1). Zonkey
software analyses (25) confirmed all specimens
as horses, except NW36 from Chupaderos,
Mexico, which is a donkey jennet (table S1).
Although a plateau in the radiocarbon cali-
bration curve prevent s easy discrimination be-
tween horses dating between 1670 CE and the
early 20th century CE, we identified three
horses from North American Indigenous con-
texts conclusively predating the Pueblo Revolt.
Near-infrared (NIR) spectrum analysis failed
to detect any external contaminants that could
have affected radiocarbon dating (materials
and methods section 3). The three specimens
include a juvenile horse burial from the site of
Blacks Fork in southwestern Wyoming, an adult
horse cranium from Kaw River , Kansas, and
isolated skeletal elements from the site of
Paa’ko, New Mexico, along with new analysis
of a previously dated specimen from Amer-
ican Falls Reservoir, Idaho, dated to between
1597 and 1657 CE (26), which we also assessed
with NIR spectroscopy (materials and methods
section 3). Assuming that the historic reinte-
gration of horses was bounded temporally by
the first presence of European horses on the
North American mainland (1519 CE), Bayesian
radiocarbon modeling suggests a date of be-
tween 1516 and 1599 CE (2s modeled ra nge)
for the initial adoption of horses by Indige-
nous societies in western North America, with
and materials and methods section 2). Various
models provided good measures of agreem ent
> 80 in all cases), and ex-
cluding anomalous values did not meaningfully
affect date estimates (materials and methods
Historic North American horses descend
primarily from Spanish genetic sources
Molecular phylogeny revealed that historic
and modern North American male horses
carried Y-chromosomal haplotypes belonging
to the “Crown group” (Fig. 2A), which became
dominant within the past ~1500 years, fol-
lowing the increasing popularity of oriental
stallions at the origin of most non-Asian do-
mestic bloodlines today, including Arabians,
Barbs, and Tho roughbreds (27). Mitochondrial
phylogenetic inference also rejected maternal
continuity from Late Pleistocene horses ex-
cavated both north and south of the North
American ice sheets (Fig. 2B). Furthermore,
BIONJ phylogenetic reconstruction based on
autosomal variation at ~7.5 million nucleotide
transversions supported a deep divergence be-
tween Late Pleistocene North American horses
and all present and past lineages identified in
Eurasia. This analysis placed both historic and
modern North American horses within the
genomic variation of modern domestic horses
(Fig. 2C). Combined, these phyloge netic recon-
structions portray historic and modern North
American horses as mainly descending from
domestic bloodlines that started spreading out-
side their native area of the Don-Volga region
no earlier than 4200 years ago (28).
Admixture graph modeling did not show
evidence of gene flow from Late Pleistocene
into historic or modern North American horses
(fig. S6.2). The individual ancestry profiles of
North American horses were consistent with
those found in recent domestic Eurasian blood-
lines, sporadically including a minor possible
contribution from Late Pleistocene North
American horses or related lineages (<0.73%)
(Fig. 2C). This ancestry was, however, not ex-
clusive to historic or modern North American
horses but instead shared across most Eurasian
lineages, including a ~4000-year-old horse from
Iberia, a ~5100-year-old horse from western
Beringia, and several ancient domestic speci-
mens such as a 1447 to 1621 CE sample from
Iran (Belgheis). Therefore, the minor ancestry
component detected likely reflects multiple
ancient contacts between Eurasia and North
America through the Beringian land bridge
during the past 830,000 years, in line with
previously reported studies (26) and also ap-
parent in mitochondrial phylogenies (Fig. 2B).
To fur ther characterize the main gene tic
sources of North American horses, we imple-
mented the qpAdm modeling rotation scheme
(29), considering either single or two-donor
sources among 37 populations. These included
Taylor et al., Science 379, 1316–1323 (2023) 31 March 2023 2of8
Department of Anthropology, University of Colorado Boulder, Boulder, CO 80309, USA.
Museum of Natural History, University of Colorado Boulder, Boulder, CO 80309, USA.
Anthropobiology and Genomics of Toulouse (CAGT, CNRS UMR5288), University Paul Sabatier, Faculté de Médecine Purpan, 31000 Toulouse, France.
Oglala Lakota, Pine Ridge Reservation, SD
Pawnee Nation of Oklahoma, Pawnee, OK 74058, USA.
Tribal Historian, Comanche Nation, Galindo Environmental Consulting LLC, Austin, TX 78757, USA.
Lakota, Pine Ridge
Reservation, SD 5777 0, USA.
International Indian Treaty Council, San Francisco, CA 94103, USA.
Sicangu Lakota, Rosebud Indian Reservation, SD 57570, USA.
He’Sapa Unity Alliance Council
of Elders, SD 57770, USA.
Cheyenne River Sioux Tribe (Lakota), Eagle Butte, SD 57625, USA.
Pueblo of Acoma, Acoma, NM 87034, USA.
Zoological Institute, Department of Environmental
Sciences, University of Basel, 4051 Basel, Switzerland.
Department of Biotechnology, Abdul Wali Khan University, Mardan 23200, Pakistan.
Institute of Culture and Environment, Alaska Pacific
University, Anchorage, AK 99508, USA.
Deg Xit’an (Athabasca n), Shageluk Tribe of Interior Alaska, Shageluk, AK 99665, USA.
Kentucky Archaeological Survey, Western Kentucky University,
Bowling Green, KY 42101, USA.
W.S. Webb Museum of Anthropology, University of Kentucky, Bowling Green, KY 42101, USA.
Conservation and Evolutionary Genetics Group, Estación Biológica
de Doñana (EBD-CSIC), 41092 Sevilla, Spain.
Laboratorio de Paleontología y Paleobiología, Instituto Andaluz del Patrimonio Histórico, 41092 Sevilla, Spain.
Université Paris-Saclay, INRAE,
AgroParisTech, GABI UMR1313, Jouy-en-Josas, 78350 Paris, France.
Musée de l’Armée, Hôtel des Invalides, 75007 Paris, France.
The Royal Danish Academy, Institute of Conservation, 1435
Copenhagen K, Denmark.
Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, CA 95064, USA.
University of Southampton Faculty of Arts and Humanities
(Archaeology), Southampton SO17 1BF, UK.
Institute for Anthropological Research, 10000 Zagreb, Croatia.
Centre National de Recherche Scientifique, Muséum national d’Histoire naturelle,
Archéozoologie, Archéobotanique (AASPE), CP 56, 75005 Paris, France.
Faculty of History, University of Oxford, Oxford OX1 2RL, UK.
Oxford Nizami Ganjavi Centre, Faculty of Oriental
Studies, University of Oxford, Oxford OX1 2LE, UK.
Genoscope, Institut de Biologie François Jacob, CEA, CNRS, Université d’Evry, Université Paris-Saclay, 91000 Évry, France.
Department, College of Science, Taif University, Taif 21944, Saudi Arabia.
Zoology Department, College of Science, King Saud University, Riyadh 12372, Saudi Arabia.
College of Science, Princess Nourah bint Abdulrahman University, Riyadh 11671, Saudi Arabia.
Department of Informati on Systems, College of Applied Sciences, Almaarefa University, Riyadh
13713, Saudi Arabia.
Department of Anatomy, Histology and Embryology, Faculty of Veterinary Medicine, University of Zagreb, 10000 Zagreb, Croatia.
Department of Archaeology, Faculty of
Humanities and Social Sciences, University of Zagreb, 10000 Zagreb, Croatia.
School of History, Classics and Archaeology, University of Edinburgh, Edinburgh EH8 9AG, UK.
Research Group, Max Planck Institute for Geoanthropology, 07745 Jena, Germany.
Department of Anthropology, University of Utah, Salt Lake City, UT 84112, USA.
Omaha, NE 68131, USA.
Department of Anthropology, University of Wyoming, Laramie, WY 82071, USA.
Department of Anthropology, University of New Mexico, Albuquerque, NM 87131, USA.
Department of Anthropology, Texas A&M University, College Station, TX 77840, USA.
SWCA Environmental Consultants, Inc., Sheridan, WY 82801, USA.
Department of Entomology,
University of Arizona, Tucson, AZ 85721, USA.
Department of Geogra phy, University of Colorado Boulder, Boulder, CO 80309, USA.
Department of Geological Sciences, University of Cape
Town, Rondebosch 7700, South Africa.
Department of History, Anthropology, Philosophy, Political Science, and Department of Spanish, Adams State University, Alamosa, CO 81101, USA.
Universidad Nacional de la Patagonia Austral, Unidad Académica Río Gallegos (ICASUR), Laboratorio de Arqueología Dr. Luis A. Borrero, CONICET, 9400 Río Gallegos, Santa Cruz, Argentina.
Quaternary Palaeontology Program, Royal Alberta Museum, Edmonton, AB T5J 0G2, Canada.
Department of Ecology and Evolutionary Biology and Howard Hughes Medical Institute,
University of California, Santa Cruz, CA 95060, USA.
Department of Earth System Science, University of California, Irvine, CA 92697, USA.
Dartmoor Hill Pony Association, Corndonford Farm,
Poundsgate, Devon TQ13 7PP, UK.
Department of Archaeology, University of Exeter, Exeter EX4 4QE, UK.
Department of Agriculture and Industry, Sul Ross State University, Alpine, TX 79832,
Department of Virology, Florida Department of Health, Jacksonville, FL 32202, USA.
Department of Animal Breeding and Genetics, Swedish University of Agricultural Sciences, SE-750
07 Uppsala, Sweden.
Xeni Gwet’in First Nations Government, 150-Milehouse, BC V0K 2G0, Canada.
McCrory Wildlife Services Ltd., New Denver, BC V0G 1S1, Canada.
Center for Animal
Breeding and Genetics, Department of Biosystems, KU Leuven, 3001 Leuven, Belgium.
Department of Animal Science, UF Genetics Institute, University of Florida, Gainesville, FL 32610, USA.
Plateforme GeT-PlaGe, Génome et Transcriptome, US1426, Centre INRAe Occitanie, 31326 Auzeville, France.
UA Accelerator Mass Spectrometry Laboratory, University of Arizona, Tucson, AZ
Department of Anthropology, University of Oklahoma, Norman, OK 73019, USA.
Oklahoma Archeological Survey, University of Oklahoma, Norman, OK 73019, USA.
of Archaeology, Max Planck Institute for Geoanthropology, 07745 Jena, Germany.
*Corresponding author. Email: email@example.com (W.T.T.T.); firstname.lastname@example.org (L.O.) †These authors contributed equally to this work. ‡Deceased.
RESEARCH | RESEARCH ARTICLE
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