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[g5oBakogER] RESEARCH ARTICLE * PHYLOGEOGRAPHY Early dispersal of domestic horses into the Great Plains and northern Rockies William Timothy Treal Taylor 1,2 *+, Pablo Librado 3 +, Mila Hunska Tasunke Icu (Chief Joseph American Horse) 4 +, Carlton Shield Chief Gover 1,5 +, Jimmy Arterberry 6 +, Anpetu Luta Wih (Antonia Loretta Afraid of Bear-Cook) 4 , Akil Nujipi (Harold Left Heron) 7 , Tanka Omniya (Robert Milo Yellow Hair) 4 , Mario Gonzalez (Nantan Hinapan) 4 , Bill Means 4,8 , Sam High Crane (Wapageya Mani) 9 ++, Mazasu (Wendell W. Yellow Bull) 4 , Barbara Dull Knife (Mah'piya Keyake Wih) 4,10 , Wakihyala Wih (Anita Afraid of Bear) 4 , Cruz Tecumseh Collin (Wanka'tuya Kiya) 4 , Chance Ward 2,11 , Theresa A. Pasqual 12 , Lorelei Chauvey 3 , Laure Tonasso-Calviere 3 , Stephanie Schiavinato 3 , Andaine Seguin-Orlando 3 , Antoine Fages 3,13 , Naveed Khan 3,14 , Clio Der Sarkissian 3 , Xuexue Liu 3 , Stefanie Wagner 3 , Beth Ginondidoy Leonard 15,16 , Bruce L. Manzano 17 , Nancy O'Malley 18 , Jennifer A. Leonard 19 , Eloisa Bernaldez-Sanchez 20 , Eric Barrey 21 , Lea Charliquart 22 , Emilie Robbe 23 , Thibault Denoblet 22 , Kristian Gregersen 23 , Alisa O. Vershinina 24 , Jaco Weinstock 25 , Petra RajicSikanjic 26 , Marjan Mashkour 27 , Irina Shingiray 28,29 , Jean-Marc Aury 30 , Aude Perdereau 30 , Saleh Alquraishi 31,32 , Ahmed H. Alfarhan 33,34 ,KhaledA.S.Al-Rasheid 32 , Tajana Trbojevic VukicVevic 35 , Marcel Buric 36 , Eberhard Sauer 37 ,MaryLucas 38 , Joan Brenner-Coltrain 39 ,JohnR.Bozell 40 , Cassidee A. Thornhill 41 , Victoria Monagle 42 ,AngelaPerri 43 ,CodyNewton 44 ,W.EugeneHall 45 , Joshua L. Conver 46 , Petrus Le Roux 47 ,SashaG.Buckser 1 , Caroline Gabe 48 , Juan Bautista Belardi 49 , Christina I. Barron-Ortiz 50 , Isaac A. Hart 39 , Christina Ryder 1 , Matthew Sponheimer 1 ,BethShapiro 51 , John Southon 52 ,JossHibbs 53 , Charlotte Faulkner 53 ,AlanOutram 54 ,LauraPattersonRosa 55 , Katelyn Palermo 56 ,MarinaSole 57 , Alice William 58 ,WayneMcCrory 59 , Gabriella Lindgren 57,60 , Samantha Brooks 61 , Camille Eche 62 , Cecile Donnadieu 62 , Olivier Bouchez 62 , Patrick Wincker 30 , Gregory Hodgins 63 , Sarah Trabert 64 , Brandi Bethke 65 , Patrick Roberts 38,66 , Emily Lena Jones 42 +, Yvette Running Horse Collin (Tasunke Iyanke Wih) 3,4 +, Ludovic Orlando 3 +* The horse is central to many Indigenous cultures across the American Southwest and the Great Plains. However, when and how horses were first integrated into Indigenous lifeways remain contentious, with extant models derived largely from colonial records. We conducted an interd isciplinary study of an assemblage of historic archaeological horse remains, integrating genomic, isotopic, radiocarbon, and paleopathological evidence. Archaeological and modern North American horses show strong Iberian genetic affinities, with later influx from British sources, but no Viking proximity. Horses rapidly spread from the south into the northern Rockies and central plains by the first half of the 17th century CE, likely through Indigenous exchange networks. They were deeply integrated into Indigenous societies be fore the arrival of 18th-century European observers, as reflected in herd management, ceremonial practices, and culture. T he spread of domestic horses and their integration into Indigenous societies con- tributed to profound social and ecolog- ical transformations across western North America. However, the mechanisms and timing of this transition are poorly understood. Horses and other members of the genus Equus originated in North America (1, 2). Horses and equids formed an important component of hu- man lifeways across the continent during the final Pleistocene (3-5), which is still encoded in some Indigenous oral traditions, including those of the Lakota (6). Although Western scholars commonly consider horses to have disappeared at lower latitudes b y the early Holocene, environmental DNA suggests their presence i n a rctic zones as late a s 5000 to 6000 years before the present (7, 8). Few ar- chaeozoological studies have carefully addres sed their possible persistence at lower latitudes during the Holocene. Viking colonizers brought horses as far as Greenland during the 10th to 14th centuries CE (9) and set tled along areas of the Newfoundland coast during the 11th century CE (10). There is, however, no direct evid ence that Viking horses reached settlements on the mainland (11). In- stead, most western scholars accept that horses were first reintroduced into the Americas by Spanish settlers in the late 15th century CE, reaching the mainland in the early 16th century CE with the Spanish colonization of Mexico (12).Duringthe17thto19thcenturiesCE, colonizing European powers, including the British, Spanish, and French (13, 14), and pos- sibly Russian and Chinese merchants (15) imported c onsiderable numbers of horses into western North America. Whereas horses would generally be cate- gorized as domestic commodities, Indigenous peoples often maintain different relationships with them. Lakota peoples attribute to horses a nationhood status equal to their own. The Lakota-horse relationship is thus one of great reverence, deeply embedded in their identity, spirituality, science, and cosmogony. Lakota peoples do not have concepts for "wild" and "domesticated." In fact, SungwakaNG--"the Horse Nation"--was neither controlled behind fences nor forced into breeding. Rather, the Lakota peoples strove to cultivate their environ- ment and adapt their lifeways to ensure that Sungwaka NG could live aligned with its natural systems. Within this nation-to-nation alliance, the horse enhanced the abilities of the Lakota with regard to hunting, mobility, healing, and more (16). Therefore, for the Lakota peoples, saying "our horse" never reflects ownership but rather responsibility for a sacred relativ e. European colonization entirely altered Indig- enous social dynamics, hierarchy, and lifeways, introducing profound changes to subsistence modes, movem ent, a nd warfar e (17 ). Many Indigenous peoples within the Great Plains and American Southwest developed horse- based pastoral or hunting economies and expanded transcontinental networks of raid- ing and exchange. Some becam e militarily dominant polities that maintained auton- omy and sovereignty into the end of the 19th century CE, with many maintaining this sov- ereignty today (18, 19). Historical models for the post-Columbian North American dispersal of horses and their integration into Indigenous cultures are almost exclusively derived from textual sources written by European observers dating largely to the 18 th and 19th centuries CE [e.g., (20, 21)]. These sources depict horses first spreading in appreciable numbers north from what is today the American Southwest after the Pueblo Revolt of 1680 CE, when Spanish settlers were temporarily expelled from much of New Mexico (22). Given that most of the continent north of New Mexico was terra incognita to European chroniclers, natural and cultural landscapes remained largely uncharacterized until the early 19th century CE (23). Furthermore, these Euro-Amer ican historic records are often rife with inaccuracies and strong anti-Indigenous biases, depreciating the fundamental rela- tionship between Indigenous peoples and horses (24). Despite representing a major source for understanding the timing and ways in which horses were managed, ridden, and integrated into early societies, archaeological remains of domestic horses from Indi genous contex ts are also overlooked (24). In this study, we ex- tensively surveyed existing archaeological collections to identify early historic horse specimens with potential for reconstructing early human-horse relationships across the American Southwest and Great Plains (Fig. 1). Togeth er, DNA, archaeozoological, and sta- ble isotope data support the introduction of RESEARCH Taylor et al., Science 379, 1316-1323 (2023) 31 March 2023 1of8 Downloaded from https://www.science.org on March 31, 2023 [DpqEwKREFx] Spanish-sourced domestic horses into Indig- enous societies across the plains before the first half of the 17th century CE. Results 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 2.06x to 12.24x, with substan- tial genome-wide sequence data for seven additional horse specimens, 0.06x to 0.96x, plus one donkey genome, 1.32x) (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 amedianboundarydateof~1544CE(Fig.1D and materials and methods section 2). Various models provided good measures of agreem ent (A model and A overall > 80 in all cases), and ex- cluding anomalous values did not meaningfully affect date estimates (materials and methods section 2). 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 1 Department of Anthropology, University of Colorado Boulder, Boulder, CO 80309, USA. 2 Museum of Natural History, University of Colorado Boulder, Boulder, CO 80309, USA. 3 Centre for Anthropobiology and Genomics of Toulouse (CAGT, CNRS UMR5288), University Paul Sabatier, Faculte de Medecine Purpan, 31000 Toulouse, France. 4 Oglala Lakota, Pine Ridge Reservation, SD 57770, USA. 5 Pawnee Nation of Oklahoma, Pawnee, OK 74058, USA. 6 Tribal Historian, Comanche Nation, Galindo Environmental Consulting LLC, Austin, TX 78757, USA. 7 Lakota, Pine Ridge Reservation, SD 5777 0, USA. 8 International Indian Treaty Council, San Francisco, CA 94103, USA. 9 Sicangu Lakota, Rosebud Indian Reservation, SD 57570, USA. 10 He'Sapa Unity Alliance Council of Elders, SD 57770, USA. 11 Cheyenne River Sioux Tribe (Lakota), Eagle Butte, SD 57625, USA. 12 Pueblo of Acoma, Acoma, NM 87034, USA. 13 Zoological Institute, Department of Environmental Sciences, University of Basel, 4051 Basel, Switzerland. 14 Department of Biotechnology, Abdul Wali Khan University, Mardan 23200, Pakistan. 15 Institute of Culture and Environment, Alaska Pacific University, Anchorage, AK 99508, USA. 16 Deg Xit'an (Athabasca n), Shageluk Tribe of Interior Alaska, Shageluk, AK 99665, USA. 17 Kentucky Archaeological Survey, Western Kentucky University, Bowling Green, KY 42101, USA. 18 W.S. Webb Museum of Anthropology, University of Kentucky, Bowling Green, KY 42101, USA. 19 Conservation and Evolutionary Genetics Group, Estacion Biologica de Donana (EBD-CSIC), 41092 Sevilla, Spain. 20 Laboratorio de Paleontologia y Paleobiologia, Instituto Andaluz del Patrimonio Historico, 41092 Sevilla, Spain. 21 Universite Paris-Saclay, INRAE, AgroParisTech, GABI UMR1313, Jouy-en-Josas, 78350 Paris, France. 22 Musee de l'Armee, Hotel des Invalides, 75007 Paris, France. 23 The Royal Danish Academy, Institute of Conservation, 1435 Copenhagen K, Denmark. 24 Department of Ecology and Evolutionary Biology, University of California, Santa Cruz, CA 95064, USA. 25 University of Southampton Faculty of Arts and Humanities (Archaeology), Southampton SO17 1BF, UK. 26 Institute for Anthropological Research, 10000 Zagreb, Croatia. 27 Centre National de Recherche Scientifique, Museum national d'Histoire naturelle, Archeozoologie, Archeobotanique (AASPE), CP 56, 75005 Paris, France. 28 Faculty of History, University of Oxford, Oxford OX1 2RL, UK. 29 Oxford Nizami Ganjavi Centre, Faculty of Oriental Studies, University of Oxford, Oxford OX1 2LE, UK. 30 Genoscope, Institut de Biologie Francois Jacob, CEA, CNRS, Universite d'Evry, Universite Paris-Saclay, 91000 Evry, France. 31 Biology Department, College of Science, Taif University, Taif 21944, Saudi Arabia. 32 Zoology Department, College of Science, King Saud University, Riyadh 12372, Saudi Arabia. 33 Biology Department, College of Science, Princess Nourah bint Abdulrahman University, Riyadh 11671, Saudi Arabia. 34 Department of Informati on Systems, College of Applied Sciences, Almaarefa University, Riyadh 13713, Saudi Arabia. 35 Department of Anatomy, Histology and Embryology, Faculty of Veterinary Medicine, University of Zagreb, 10000 Zagreb, Croatia. 36 Department of Archaeology, Faculty of Humanities and Social Sciences, University of Zagreb, 10000 Zagreb, Croatia. 37 School of History, Classics and Archaeology, University of Edinburgh, Edinburgh EH8 9AG, UK. 38 isoTROPIC Research Group, Max Planck Institute for Geoanthropology, 07745 Jena, Germany. 39 Department of Anthropology, University of Utah, Salt Lake City, UT 84112, USA. 40 Archaeological consultant, Omaha, NE 68131, USA. 41 Department of Anthropology, University of Wyoming, Laramie, WY 82071, USA. 42 Department of Anthropology, University of New Mexico, Albuquerque, NM 87131, USA. 43 Department of Anthropology, Texas A&M University, College Station, TX 77840, USA. 44 SWCA Environmental Consultants, Inc., Sheridan, WY 82801, USA. 45 Department of Entomology, University of Arizona, Tucson, AZ 85721, USA. 46 Department of Geogra phy, University of Colorado Boulder, Boulder, CO 80309, USA. 47 Department of Geological Sciences, University of Cape Town, Rondebosch 7700, South Africa. 48 Department of History, Anthropology, Philosophy, Political Science, and Department of Spanish, Adams State University, Alamosa, CO 81101, USA. 49 Universidad Nacional de la Patagonia Austral, Unidad Academica Rio Gallegos (ICASUR), Laboratorio de Arqueologia Dr. Luis A. Borrero, CONICET, 9400 Rio Gallegos, Santa Cruz, Argentina. 50 Quaternary Palaeontology Program, Royal Alberta Museum, Edmonton, AB T5J 0G2, Canada. 51 Department of Ecology and Evolutionary Biology and Howard Hughes Med ical Institute, University of California, Santa Cruz, CA 95060, USA. 52 Department of Earth System Science, University of California, Irvine, CA 92697, USA. 53 Dartmoor Hill Pony Association, Corndonford Farm, Poundsgate, Devon TQ13 7PP, UK. 54 Department of Archaeology, University of Exeter, Exeter EX4 4QE, UK. 55 Department of Agriculture and Industry, Sul Ross State University, Alpine, TX 79832, USA. 56 Department of Virology, Florida Department of Health, Jacksonville, FL 32202, USA. 57 Department of Animal Breeding and Genetics, Swedish University of Agricultural Sciences, SE-750 07 Uppsala, Sweden. 58 Xeni Gwet'in First Nations Government, 150-Milehouse, BC V0K 2G0, Canada. 59 McCrory Wildlife Services Ltd., New Denver, BC V0G 1S1, Canada. 60 Center for Animal Breeding and Genetics, Department of Biosystems, KU Leuven, 3001 Leuven, Belgium. 61 Department of Animal Science, UF Genetics Institute, University of Florida, Gainesville, FL 32610, USA. 62 Plateforme GeT-PlaGe, Genome et Transcriptome, US1426, Centre INRAe Occitanie, 31326 Auzeville, France. 63 UA Accelerator Mass Spectrometry Laboratory, University of Arizona, Tucson, AZ 85721, USA. 64 Department of Anthropology, University of Oklahoma, Norman, OK 73019, USA. 65 Oklahoma Archeological Survey, University of Oklahoma, Norman, OK 73019, USA. 66 Department of Archaeology, Max Planck Institute for Geoanthropology, 07745 Jena, Germany. *Corresponding author. Email: william.taylor@colorado.edu (W.T.T.T.); ludovic.orlando@univ-tlse3.fr (L.O.) +These authors contributed equally to this work. ++Deceased. RESEARCH | RESEARCH ARTICLE Downloaded from https://www.science.org on March 31, 2023 [ToWAAAAABj] Late Pleistocene North American horses as well as a representative panel of both modern and ancient domestic horses from around the globe. This analysis rejected Late Pleistocene North American horses as a possible source for both historic and modern North Amer ican horses but supported domestic bloodlines with almost exclusively European origins. Moreover , historic North American horses showed strong genetic affinities to three ancient do mestic horses from Spain (Jal5885), Iran (Belgheis), andFrance(Inva22),datedtobetweenthe 15th and 18 th centuries CE. This was true for all specimens except the one from Fort Boonesborough (Boones1/Kentucky, 1778 CE), which showed greater genetic affinity to British horses from the 18th century CE (Witter Place) (Fig.3A).Notably,theinfluenceofBritish bloodlines is also apparent among modern North American horses, which are commonly modeledasmixturesbetweenSpanish-like (Galiceno and Cartujano) an d British-like sources(ThoroughbredsandWelsh,Shetland, and Dartmoor Hill ponies) (table S3 and Fig. 3B). Th i s indicates a temporal shift in the genetic composition of North American horses tracing new inputs from growing British and, later, American presence in eastern North America, and the integration of these horses into In- digenous spheres of interaction. Historic North American horses show greater relatedness to the historic Iberian specimen (Jal5885) than to modern Welsh bloodlines, but modern North American horses show increase d Welsh related- ness (Fig. 3, B, C, and E). This is true for all except for feral horses isolated in the Santa Cruz Islands (STCZ3322 and STCZ3327), con- sistent with their alleged Spanish historical origins. Finally, both historic and most modern North American groups remain closer geneti- cally to Jal5885 than to Icelandic horses (Fig. 3D), and qpAdm modeling revealed ancestry pro- files incompatible with genetic contribution from a Viking specimen dating to between the 9th and 11th centuries CE (VHR102) (sup- plementary materials section 6). The genomic analyses described above focus on the minute fraction of the genome that is different across all horse lineages investigated. However, the Lakota find key instruction in the>99%ofthegenomefractionthatappears common to all. Pre-Pueblo Revolt contribution of horses to Indigenous beliefs, trade, and transport networks Archaeological specimens dated to the early 17th century CE show pathological and osteo- logical evidence of care, management, and use in transport. A horse phalanx from an Indigenous-affiliated context at Paa'ko Pueblo, New Mexico, and a metacarpal from American Falls Reservoir, Idaho, demonstrate the pres- ence of horses among Native communities as far north as Idaho by the first half of the 17th century CE. Furthermore, a foal from Blacks Fork, Wyoming, exhibits entheseal ossification of the nuchal ligament attachment at the rear of the skull at levels typically and exclusively found in horses used for transport or kept in confinement (30, 31). It also features a severe, healed cranial fracture that may have resulted from a kick caused by confineme nt in close Taylor et al., Science 379, 1316-1323 (2023) 31 March 2023 3of8 Fig. 1. Sample spatial and chronological distribution. (A) American samples. Processed archaeological and/or museum specimens are shown as red triangles, with those successfully sequenced shown as a plain circle; modern sp ecimens are indicated with squares. LP, Late Pleistocene [data from Vershinina et al.(26)]; ELEN, Equus lenensis (Siberia). (B) Ancient Eurasian samples in the comparative panel (table S1). Horses from western Beringia are colored in purple, the remaining in blue. Modern genomes are not projected on the map but are provided in table S1. (C) Geographic visualization of Bayesian radiocarbon dates for early horse dispersals. Produced in OxCal assuming a uniform prior , with results transferred and visualized in ArcGIS. For each specimen, point diameter corresponds to the portion of total probability distri bution withi n each time slice. Note tha t presence or absence of a dot does not necessarily indicate occupation during a given time slice (see materials and methods section 2 for detailed chronological modeling of sites thought to predate 1680 CE). (D) Modeled radiocarbon boundary for introduction of Spanish-sourced domestic horses into Indigenous societies in the western United States, based on analyzed radiocarbon dates. RESEARCH | RESEARCH ARTICLE Downloaded from https://www.science.org on March 31, 2023 [BxH0RYXZIX] proximity to other h orses (materials and methods section 4). Moreover, this foal was recovered in ritual features alongside coyotes (32), indicating that hor ses we re already integrated into existing social and ceremonia l traditions by the first half of the 17th century CE. Nuchal ossification along with charac- teristic damage linked to the use of a metal bit, including erosion of the anterior cementum and enamel o f the lower second premolar, were also found in the horse from Kaw River, Kansas, dating to the same period (Fig. 4) (33). Several other pathological features reflect the useofametalcurbbitofthetypeusedbyearly Spanish colonists and later Mexican and Taylor et al., Science 379, 1316-1323 (2023) 31 March 2023 4of8 O GLA K O/P u ebl o an2 4 FO R T33 47 OGL AKO/C h o c t a w 78 M U ST10 99 MU S T 1096 CHI C O L T I N OG LA KO /P a iute54 I d ah o A m e r ica nF alls 3 88 +- 46 F O R T331 7 F O RT3 344 FO R T334 2 MG M R2726 O GLAKO / Ch oc t aw 75 OG L AK O / C h o ct a w73 O GLA K O / C h o c t a w 3 2 O GLA K O /C ho c taw 3 6 OG L A K O / C h ic k a s a w 8 3 OGL A K O / P a i u t e 88 O G L AK O /Pa i u te 8 7 OGLAKO /Sa ko w in 35 S T C Z 3 332 S TCZ3 3 8 O G L A KO / C h o ctaw7 1 OG L AK O / C h o c ta w9 W y om ing N atu ra l T ra p 2 5 1 6 0 +- 1 65 W yo m in g Na t u ra l T r a p 2 5 2 1 4 +- 1 5 3 W y om i n g Na t u r a l T r ap 2 53 56 +- 1 4 7 Wyom i ng N atu r al T r ap 2493 0+- 204 I d a h o J aw D ro ppe r 17 3 24 +- 9 5 NW3 / New Me xico NW 32/Kans as Bla c k s F o rk / Wy omin g M GM R293 9 FOR T33 5 9 FO R T3356 F O R T3 355 M GP L28 65 NW10 / Nebr a s k a 2 SA H 7 / A r gen t i na FO R T3 3 5 8 OGLAKO / Ute 1 6 O G L A KO/ C h o ct a w 72 O G L AKO/Cho c ta w 2 6 FO RT3 38 5 OGL A KO/ S a k o win 1 1 STCZ332 8 S TC Z33 2 7 OGL AKO/ Sa k o w i n 2 2 FO RT340 7 FO RT34 0 8 OGLAK O/Choctaw 81 OGL AK O/ Oji b w e 8 OGL A K O/ Oj i b we 1 2 O G LA K O / P u eb loa n 4 0 SA H 4 / A r g e n ti n a OG L A K O / S ako wi n 85 O G L AKO/U t e 6 8 S A H 2/A rg entin a OGLA K O/ S ak o wi n 70 NW 1 4 / W y o mi ng O GLAKO / A p ache 13 O G L A KO/Ap ache17 O GL A K O / Ojibw e 7 Ka wR ive r /Ka n sa s OGLAK O/O j i bw e 2 CHICO L T IN OGL A KO/ Ch o cta w78 S TCZ3 328 O G LAKO/Choc taw3 2 2 4 3 3 T R O F F O RT33 85 WURTB W01 HOL S 0 1 H OLS 00 4 T H O R TH O R3 4 7 5 W S T F 01 F R MO1 7 9 8 M U S T 10 96 YIL I 2 SW R M 310 Q R TR225 FRMO1 9 51 S W R M441 L I PI H OL S P1 AKTK001 A K TK 0 0 6 OG L AKO/Athab a s c 58 HANO0 1 H A NO02 HANO03 WSTF02 M A R VELE21 OTE2 M A R VE L E 32 L R 18 _ 9 NB44 V HR0 11 VH R037 MONG 26 28 PR40 B ZNK1002_4 BES T A5 LR18_16 U R 17_1 LR18 _ 6 2 R N131 CGG101 3 9 7 B O T A I1 BO T AI18_26 PRZW5 3 3 R US37 GR AL 7 GRAL9 B A T A GA I KSK16232 S P A IN3 9 Z A M9 Y G303 R US38 NW3 / N e w Mexi c o BELG H E I S G EP14 GV A 3 7 5 MG MR 272 6 MGPL2865 TUR140 SAH 4/Argen t ina INV A 22 N O R I 180 O G L AKO/iS ak ow i n11 H AFL0004 STCZ3 332 C AR T 5 3 2 C A RT 554 CART 5 3 9 C A R T53 4 MAR W AR A B948 ARAB 18 T RAK02 T RAK 0 1 BA CA 33 56 OGLAKO/ Ojibwe7 MZR1 UK1 9 NW26/Oklahoma DATO3 GE P 13 CGG1 0 1397 I SSYK 1 KYR H 8 J IZ I 4 L KZ T 1 4 G EP2 1 K H O T ONT VHR102 TU R14 5 MA RVELE 01 G VA1 2 2 JI Z I 3 L K Z T 28 P A VH2 ICELP5782 VHR0 17 K Y R H 1 0 M A R V ELE18 D A TO1 2 JEJU2 J E J U 3 NUST A R5 B A PSKA S AA1 4 5 7 7 G N O M Y C R O W N H A P L O G R O U P Bootstrap 90 92.5 95 97.5 100 THOR3903 THOR THOR3475 TRAK02 TRAK01 WURTBW01 SWRM310 WSTF02 HANO03 HANO02 HANO01 HOLS0004 HOLSP1 HOLS01 WSTF01 SWRM441 PAIN16 MIXD QRTR225 QRTR070 QRTR OGLAKO/Puebloan40 ARAB18 ARAB09 ARAB948 RTPN033 BELGHEIS AKTK006 AKTK03 AKTK001 STDB03 STDB01 STDB02 KawRiver/Kansas GALI3313 GALI3311 BACA3358 BACA3356 BACA3359 BACA3355 GALI3309 GALI3307 LIPI MARW FLCR3408 FLCR3407 OGLAKO/Athabascan58 Cartujano562 Cartujano534 Cartujano559 Cartujano532 Cartujano539 Cartujano561 Cartujano554 Cartujano547 SORR2 SORR1 JAL5886 MGPL2865 MGMR2726 MGMR2939 SAH4/Argentina SAH2/Argentina SAH7/Argentina NW32/Kansas NW10/Nebraska NW14/Wyoming NW26/Oklahoma OGLAKO/Apache17 OGLAKO/Apache13 STCZ3330 STCZ3327 STCZ3328 STCZ3332 OGLAKO/Paiute87 OGLAKO/Paiute54 OGLAKO/Paiute88 OGLAKO/OcetiSakowin22 OGLAKO/OcetiSakowin11 CHICOLTIN OGLAKO/Choctaw9 OGLAKO/Choctaw81 OGLAKO/Choctaw78 OGLAKO/Choctaw26 OGLAKO/Choctaw75 OGLAKO/Choctaw71 OGLAKO/Choctaw73 OGLAKO/Choctaw32 OGLAKO/Chickasaw83 OGLAKO/Choctaw36 MUST1099 MUST1096 OGLAKO/OcetiSakowin44 OGLAKO/OcetiSakowin35 OGLAKO/OcetiSakowin70 OGLAKO/Ute68 OGLAKO/Ute16 OGLAKO/OcetiSakowin85 OGLAKO/Choctaw72 OGLAKO/Puebloan24 OGLAKO/Ojibwe8 OGLAKO/Ojibwe7 OGLAKO/Ojibwe2 OGLAKO/Ojibwe12 NW3/NewMexico FORT3344 FORT3342 FORT3347 FORT3385 FORT3317 YILI2 FortBoonesborough/Kentucky UK18 HAFL0004 HAFL0002 HAFL0003 NORI180 BlacksFork/Wyoming INVA22 FRMO1951 FRMO0001 FRMO1798 UK15 UK17 UK16 WLSH007 WLSH006 UK19 DART5 5 DART5 1 DART6 7 GVA122 DUEL SHET041 SHET020 VHR017 VHR102 ICELP5782 VHR062 VHR037 VHR011 VHR010 MARVELE32 MARVELE01 MARVELE21 MARVELE18 OTE2 P132 GVA375 SAA1 GVA123 MZR1 MZH24 MZHT22 NIMU2 JIZI4 LKZT28 JIZI3 NIMU20 MZHT21 LKZT22 LKZT14 NIMU21 JIZI2 MIQI9916 JICH5 DATO28 DATO12 DATO3 MOGOWZ6 ELC21 MONGWMG8 YAQI29 MONG7754 JEJU2 JEJU1 JEJU3 MONG2629 MONG2628 OTOK16 ISSYK1 PAVH2 GEP14 KHOTONT NUSTAR5 YAKT7 YAKT6 CGG101397 YAKT2 BAPSKA GEP13 RUS37 GEORGIA2 KYRH8 KYRH10 TUR140 GEP21 TUR145 RUS38 UR17_41 NB44 UR17_1 AC8811 LR18_62 BESTA5 PR40 MOLDA1 UR17_26 LR18_9 BZNK1002_4 RN53 LO2018_15 LR18_16 KSK16232 CD2017 PRZW533 PRZW3879 PRZW339 BOTAI2018_26 BOTAI1 BOTAI2018_14 ZAM9 SPAIN39 UE2275 GRAL13 GRAL7 GRAL9 HOHLER3_3 HOHLER3_1 HOHLER2 RN131 RN130 RN109 CGG10022 CGG10023 BATAGAI YG303 YG188 DONK 0.00 0.25 0.50 0.75 1.00 Admixture Proportion LP Eastern Beringia ELEN (Western Beringia) Asian Far East LP Continental North America Historic North America Modern North America Historic South America Modern South America Central Asia, Mongolia and Europe DOM2 Origin of the sample (Inner circle) 10,000 20,000 30,000 40,000 50,000 Age of the sample (Outer circle) A B C Uniparental Markers DOM2 Fig. 2. Phylogenetic affinities. (A) Y-chromosomal maximum-likelihood tree (N = 110; 5244 nucleotide transversions). (B) Mitochondrial maximum-likelihood tree (N = 340; 16,406 base pairs). (C) Neighbor-joining tree based on ~7.5 million nucleotide transversions (N = 241; left), and the corresponding gene tic ancestry profiles (right). The ancestry proportions for K = 7 genetic componen ts were estimated using St ruct-f4 (28). Sample name s written as "OGLAKO/Apa che" represe nt moder n North Ame rican horses culturally associated with Apache communities. LP, Late Pleistocene; ELEN, Equus lenensis (Siberia); DOM2, modern domestic lineage descending from a lower Don-Volga genetic source expanding to Eurasia after 4200 years ago. RESEARCH | RESEARCH ARTICLE Downloaded from https://www.science.org on March 31, 2023 [06FgAAAAAY] Navajo craftsmen, such as the fracturing of the upper palate caused by the curb (34) and arthritis of the temporomandibular joint. Combined, these analyses indicate that early plains horses were already used for mounted riding. Strontium isotope values for Blacks Fork, reflecting prenatal (dP4, 87 Sr/ 86 Sr = 0.70970) and postnatal (M1, 87 Sr/ 86 Sr = 0.70969) values for the foal, are directly in line with published values for modern fauna from the Gre en Riv er Basin, where the specimen was found (0.70950 to 0.71000) (35, 36). Although similar values characterize some stretches of the Colorado River tributary system farther to the south (37), the Blacks Fork values rule out most of Wyoming, including the border with Utah (38). Additionally, strontium isotope results from the Kaw horse, Kansas, are consistent with available values from northeastern Kansas (39) but indicate some mobility during the recorded sequence ( 87 Sr/ 86 Sr = 0.70905 near the crown, versus 87 Sr/ 86 Sr = 0.70930 near the root), which spans a 12- to 14-month period acro ss the horse'sfourthandfifthyearoflife,according to mineralization schedules ( 40). When con- sidered alongside stable isotopes of oxygen (d 18 O) and carbon (d 13 C), strontium isotope values suggest that the Kaw horse spent part of its life farther north, indicating gradual movement from an area matching reference data from South Dakota, Nebraska, and Iowa (Fig. 4 and materials and methods section 5). Therefore, is otope values from bo th early horses with complete dentition suggest that the animals were either raised and managed Taylor et al., Science 379, 1316-1323 (2023) 31 March 2023 5of8 Spanish-like French-like FortBoonesborough/Kentucky NW10/Nebraska BlacksFork/Wyoming NW32/Kansas KawRiver/Kansas NW3/NewMexico SAH4/Argentina SAH7/Argentina NW14/Wyoming* AB DC MGPL2865 FORT3344 FORT3347 MUST1096 FORT3317 FORT3342 FORT3385 FLCR3407 FLCR3408 OGLAKO/Choctaw36 OGLAKO/Ojibwe8 OGLAKO/Choctaw9* MUST1099 MGMR2726 MGMR2939 STCZ3327 STCZ3328 STCZ338 STCZ3332 BACA3355 BACA3356 OGLAKO/Paiute54 OGLAKO/Athabascan58 OGLAKO/Paiute68 OGLAKO/Choctaw71 OGLAKO/Choctaw72 OGLAKO/Choctaw75 OGLAKO/Choctaw81 OGLAKO/Chickasaw83 OGLAKO/Choctaw85 M U S T1099 M GMR2726 M GMR293 9 S TCZ332 7 S TCZ3328 S TCZ338 S TCZ3332 B A CA335 5 B A CA3356 OGLAKO/Paiute5 4 O GLAKO/Athabascan58 O GLAKO/Paiute68 OGLAKO/Choctaw7 1 OGLAKO/Choctaw72 O GLAK O /Choctaw7 5 OGLAKO/Choctaw8 1 OGLAKO/Chickasaw83 OGLAKO/Choctaw8 5 OGLAKO/Puebloan40 OGLAKO/Puebloan24 British-likeSpanish-like French-like British-like OGLAKO/Ojibwe7 OGLAKO/Ojibwe2 + OGLAKO/OcetiSakowin35 OGLAKO/OcetiSakowin11 OGLAKO/Choctaw32 OGLAKO/Apache13 OGLAKO/Apache17 OGLAKO/Choctaw73 OGLAKO/OcetiSakowin44 f 4 (S - URAL,X; CPONT,JAL5886) 0.001 0.002 0.003 0.004 0.005 0.001 0.002 0.003 0.004 0.005 FORT3385 BACA3359 BlacksFork/Wyoming STCZ3322 MGMR2726 STCZ3327 CHICOLTIN E -20246 Z-score f 4 (Historic NA, Modern NA; JAL5886, WELS) Avg Z-score = 2.31 95% CI 0.001 0.002 0.003 0.004 0.005 f 4 (S-URAL,X; CPONT,ICEL) f 4 (S-URAL,X; CPONT,WELS) BlacksFork/Wyoming OGLAKO/Ojibwe2 OGLAKO/Ojibwe12 MUST1096 Fig. 3. Temporal changes in the genomic makeup of American horses. (A) Ternary plots showing ancestry combinations of the historic horse genomes from North America. (B) Ternary plots showing ancestry combinations of the modern horse genomes from North America. Ancestry proportions were estimated using qpAdm modeling and rotating 37 possible donor sourc es. Models with highest P values are shown where multiple models could not be rejected. Asterisks depict two samples for which the ancestry profiles presente d correspond to the second best qpAdm model (table S3), while the cross accompany ing OGLAKO/Ojiwbe2 indicates that its best qpAdm model involves Galiceno and Icelandic horses as donor populations, instead of British sources (s upplementary materials section 6). (C) Relative genetic affinities of historic and modern North American horses to Jal5885 versus Welsh horses. Genetic affinities are estimated using f 4 -statistics in the form of (S-URAL, X; C-PONT, Jal5885 or Welsh), wh ere X represents historic and/or modern horses from North America, and C-PO NT horses from the third millennium CE showing the closest genomic affinities to modern DOM2 domesticates. Triangles show the two historic samples that returned modern radiocarbon dates (NW14/Wyoming and SAH7/Argentina). (D) Relative gen etic affinities of historic and modern North American horses to Jal5885 versus Icelandic horses. The calculations are the same as in (C), except that modern Welsh horse genomes were replaced by genomes from modern Icelandic horses and the VHR102 Viking sample (850 CE to 1050 CE). An additional qpAdm analysis considering modern Icelandic horses and the sample VHR102 as separate donors of genetic ancestry confirmed that the specimen OGLAKO/Ojibwe2 received introgre ssion from modern Icelandic horses, ruling out genetic contribution from relict populations of Viking horses (supplementary materials section 6). Error bars corr espond to twice the standard error estimated from jackknifing. (E) Distribution of Z-scores for f 4 -statistics of the form (Historic North American Horse, Modern North American Horse; Jal5885, Welsh). RESEARCH | RESEARCH ARTICLE Downloaded from https://www.science.org on March 31, 2023 [BYJddYaXAy] locally (Blacks Fork) or within a territory ex- tending even farther away from the European colonial sphere (Kaw). A d 13 C spike toward the end of the sampled section of the Kaw molar shows a strong input of C 4 -pathway plants, potentially reflecting winter foddering with the Indigenous domes- tic crop maize (materials and methods sec- tion 5), a traditi onal practice among Plains groups, including the Paw nee. Other situa- tions, such as herd movement south and west to rangeland higher in C 4 grasses or during events such as seasonal bison hunts, could ac- count for the enriched value but are not con- sistent with strontium and oxygen isotope d ata. Pawnee (Chaticks si Chaticks) villages typically relied on stores of maize to subsist through the winter (41), and the practice of winter fodder- ing horses with maize in the northern Missouri River region is documented ethnographically during the 18th and 19th centuries CE, includ- ing among the Pawnee (42, 43). Regardless of whether the Kaw specimen was affiliated with ancestral Pawnee or another Central Plains nation, our results indicate the presence of horses in Indigenous cultural and economic systems of the Missouri River drainage during the first half of the 17th century CE. Discussion Our archaeological analyses show the dispersal of domestic horses from Spanish settlements in the American Southwest to the northern Rockies and central Great Plains by the first half of the 17th century CE at the latest. They provide evidence of local raising and veteri- nary care of horses, likely foddering with do- mestic maize, and use of horses in transport by Indigenous peoples by this time. A directly dated radiocarbon specimen from Paa'ko Pueblo in northern New Mexico shows that horses reached the region via Indigenous groups before Spanish colonization of the American Southwest, as previously hypothesized (44, 45). Moreover , our new tempora l framework shows tha t horse s were present across the plains long before any documented European presence in the Rockies or the central plains. Despite their Taylor et al., Science 379, 1316-1323 (2023) 31 March 2023 6of8 Fig. 4. Horse herding in the 17 th century CE. (A) Osteological indicators of bridling and riding: (1) palatal damage from curb bit; (2) bit damage to anterior margin of lower second premolar; (3) osteoarthritis at the temporoman dibular joint; (4) entheseal changes to the nuchal ligament attachment site; and (5) remodeling of the premaxilla. (B) Osteological indicators of human activity: (1) Healed kick fracture in young foal, suggesting health care. The inset shows heali ng at 50x magnification. (2) Ossification of the nuchal ligament, indicating us e in transport or confinement and/or tethering. (C) Modeled northward dispersal of hors es. Model based on archaeological discoveries and consideration of historically documented cultural developments and migrations. (D) Isotopic evidence for fodd ering. Stable carbon and oxygen and strontium isotope sampling locations for the Kaw River horse, as measured in millimeters from the root (lower-right third m olar). Sampling locations are represented on the x axis as the midpoint of a 2-mm sec tion (e.g., the section from 0 to 2 mm is shown as "1" on the graph). Filled tria ngles show strontium isotope values, with stable carbon values represented by filled circles, and stable oxygen values as open circles. [Kaw horse image credit: E. Sc ott] RESEARCH | RESEARCH ARTICLE Downloaded from https://www.science.org on March 31, 2023 [0fgAAAABJR] Ibe ri an genetic makeup and earlier arguments attributing one of these horses to Spanish ex- ploration (46), strontium, carbon, and oxygen isotope results suggest that these animals were raised and died locally. Osteological analyses also provide some indication that horse dis- persal was tied to broader economic links; the Kaw River horse was indeed controlled with a European-style metal curb bit, and the Blacks Fork horse body was cut with metal tools (32). Therefore, a possible mechanism of horse dis- persal was exchange across Indigenous net- works at the margins of New Mexico and the American S outhwest (47). Once acquired, horses could have moved via ancient trade routes based on kinship ties and social networks es- tablished throughout the Great Plains and Rockies millennia before European contact (48). The dispersal of Puebloan groups toward the northeast, from Spanish New Mexico into western Kansas, providesanothermechanism for the transmission of domestic horses into the Central Plains. Our findings have deep ramifications for our understanding of social dynamics in the Great Plains during a period of disruptive social changes for Indigenous peoples. The area of southwestern Wyoming including Blacks Fork is considered to be a homeland for the Shoshonean-speaking ancestral Comanche (Nu - mu - nu - u - ), who migrated to the southern plains of Texas, New Mexico, Colorado, and Oklahoma before the early 18th century CE (49, 50). The drive to acquire horses from Spanish New Mexico is often cited as a likely mechanism for this transcontinental move- ment (50). However, our new data--which align with some Comanche and Shoshone oral accounts (51) (materials and methods section 7)--suggest that ancestral Comanche had already integrated horse raising, ritual practices, and transport into their lifeways at least a full half century before their southward migration, effectively moving to the southern plains as horse herders. Once in the southern plains, the Comanche were able to marshal these advantages, along with their herding and equestrian skills, to build an empire on horse and bison trade by the middle of the 18th century CE (19). By the time Europeans arrived in southwestern Wyoming, the area was already a critical "secondary diffusion center" for horse transmission to Northern Plains groups (17, 52). Considering the small body of archaeol ogical data availabl e, our findings raise the possibility of rapid, non- European transmission of horses farther north- ward, including the Columbia Plateau, the Canadian Rockies, and the middle and upper Missouri regions. DNA data from horses currently caretaken and historically protected by the Oglala Lakota were included in this study. All protocols for the transmission of sacred and traditional knowledge were followed, as outlined by our Lakota Elder Knowledge Keeper Internal Re- view Board. They provide further context of the findings, which clarifies the Lakota cultu re and their relationship with the horse. Chief Joe American Horse states: "Horses have been part of us since long before other cultures came to our lands, and we are a part of them. The Horse Nation is our relative. We always protect our relatives and the next seven generations. We stand with the horse and we will alwa ys do so however it has evolved through its journey. That is what being Lakota is" (original quote, in Lakota, provided in materials and methods section 7). This study established that Indige- nous peoples were living and interacting with the horse before the Pueblo Revolt of 1680 CE, which was the earliest date accepted by Western science. H owever, current genetic evidence shows that the horses caretaken by Indigenous peoples from as early as the firsthalfofthe17thcenturyCEdonotsharean excess of genetic ancestry with Late Pleisto- cene North American horses. Given that the Horse Nation is foundational to Lakota life- ways (16), one possible implication of this find- ing is that relationships of the kind developed by Lakota peoples could have already been in place by the Late Pleistocene. Such life manage- ment practices may even have extended to other members of the horse family at that time. Testing these implications requires fur- ther paleontological, archaeological, genetic, and ethnographic research. Dr. Antonia Loret ta Afraid of Bear-Cook adds: "The Horse Nation always chose their own mates. Bringing in new blood is a replenishment and renewal process of life that we celebrate. It strengthens our we (the life force from our blood). No matter how our horses may have transformed, or where they are around the world, we will always call to them. Toget her we are home" (original quote, in Lakota, provided in materials and methods section 7). This study demonstrates that colonization did not just drastically affect Indigenous peoples but also their horses, whose genetics captures an ancestry shift from Span- ish to British bloodlines. 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ACK NOW LED GME NTS We are grateful to the scholars, elders, and leaders who make up the Lakota Review Team and who supervised the Oglala Institutional Review Board process for this research, as well as their counterparts from other Indigenous nations who supported this research and interpretation. Wopila Tanka to the Horse Nation and each of the Indigenous nations who served as cultural caretakers for these horses. Special thanks to M. Sims, S. Olsen, and C. Beard of the KU Natural History Museum for facilitating access to the Kaw specimen (KUVP 347; imagery provided courtesy of E. Scott); to B. Peecook and A. Commendador for facilitating access to the American Falls specimen (IMNH 71004/25895, a Bureau of Reclamation specimen under the care of the Idaho Museum of Natural History, Idaho State University); and to D. Walker and the University of Wyoming Archaeological Repository (Office of the Wyoming State Archaeologist, 1000 E. University Ave., Dept. 3431, Laramie, WY 82071, USA) for providing imagery and facilitating access to collections at Blacks Fork (48SW8319), located in Flaming Gorge National Recreation Area, managed by the USDA-Ashley National Forest. Analysis of Blacks Fork material was completed with permission from the USDA-Ashley National Forest, coordinated through J. Rust, Heritage Program Leader. Additional thanks to B. Britt (the Maxwell Museum of Anthropology), D. Gifford-Gonzalez, and K. Kjaer for facilitating research and access to additional collections. Funding: Research was funded by an award from the National Science Foundation (Collaborative Research: Horses and Human Societies in the American West, Awards 1949305, 1949304, and 1949283) and from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant agreement 681605). Additional funding was provided by the European Union's Horizon 2020 research and innovation programme under the Marie Sklodowska- Curie (grant agreement 890702-MethylRIDE); the French Government "Investissement d'Avenir" FRANCE GENOMIQUE (ANR-10-INBS-09); the Universidad Nacional de la Patagonia Austral (PIP 29/A476); the Resea rchers Supporti ng Projects at King Saud University, Riyadh, Saudi Arabia (NSRSP2022R1), Taif University, Taif, Saudi Arabia (NSTURSP2022), Princess Nourah bint Abdulrahman University, Riyadh, Saudi Arabia (NSPNURSP2022), and Almaar efa University, Ri yadh 13713, Saudi Arabia (NSTUMA/1); and the Marie Sklodowska Curie programme (101027750-HOPE). P.R. and M.L. thank the Max Planck Society for funding for the isotope analyses. Author contributions: Designed and coordinated the study: W.T.T.T., E.L.J., and L.O. Performed radiocarbon dating: J.S. and G.H. Performed ancient DNA laboratory work: L.Chau., L.T.-C., S.S., A.S.-O., A.F., N.K., C.D.S., X.L., S.W., and Y.R.H.C., with input from L.O. Performed ancient DNA computational analyses: P.L. and L.O. Provided material, reagents, and methods: W.T.T.T., P.R., B.L.M., N.O., J.A.L., E.B.-S., E.B., L.Char., E.R., T.D., K.G., A.O.V., J.W., P.R.S., M.M., I.S., J.-M.A., A.Perd., S.A., A.H.A., K.A.S.A.-R., T.T.V., M.B., E.S., C.N., J.R.B., C.A.T., W.E.H., C.G., J.B.B., C.I.B.-O., C.R., M.Sp., B.S., J.S., A.O., L.P.R ., K.P., M.So., A.W., W.M., G.L., S.B., C.E., C.D., O.B., P.W., G.H., S.T., B.B., E.L.J., Y.R.H.C., and L.O. Interpreted data: W.T.T.T., P.L., J. A., C.W., J.B.-C., J.R.B., C.A.T., V.M., A.Perr . , W.E.H., J.L.C., P.L.R., S.G.B., J.B.B., C.R., M.Sp., G.H., S.T., B.B., P.R., E.L.J., Y.R.H.C., and L.O. Wrote the supplementary text: W.T.T.T., P.L., M.M., E.S., V.M., A.Pe rr., C.N., S.G.B., J.B.B., C.I.B.-O., I.A.H., S.T., B.B., E.L.J., Y.R .H.C., and L.O., with input from all coauthors. Wrote the article: W.T.T.T., B.B., E.L.J., Y.R.H.C., and L.O., with input from all coauthors. Competing interests: The authors declare that they have no competing interests. Data and materials availability: The sequence data generated in this study are available for download at the European Nucleotide Archive (accession no. PRJEB56773). The repository information for each individual sample and all other data used in t his study are included in table S1. License information: Copyright (c) 2023 the authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original US government works. https://www.science.org/about/science- licenses-journal-article-reuse SUPPLEMENTARY MATERIALS science.org/doi/10.1126/science.adc9691 Materials and Methods Figs. S1.1 to S6.4 Tables S1.1 to S3.3, S5.1, and S6.1 to S6.3 References (53-124) MDAR Reproducibility Checklist View/request a protocol for this paper from Bio-protocol. Submitted 15 May 2022; accepted 14 February 2023 10.1126/science.adc9691 Taylor et al., Science 379, 1316-1323 (2023) 31 March 2023 8of8 RESEARCH | RESEARCH ARTICLE Downloaded from https://www.science.org on March 31, 2023 [7ZbxzQAADA] Use of this article is subject to the Terms of service Science (ISSN ) is published by the American Association for the Advancement of Science. 1200 New York Avenue NW, Washington, DC 20005. The title Science is a registered trademark of AAAS. Copyright (c) 2023 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works Early dispersal of domestic horses into the Great Plains and northern Rockies William Timothy Treal Taylor, Pablo Librado, , Carlton Shield Chief Gover, Jimmy Arterberry, , , , , Bill Means, , , , , , Chance Ward, Theresa A. Pasqual, Lorelei Chauvey, Laure Tonasso-Calviere, Stphanie Schiavinato, Andaine Seguin- Orlando, Antoine Fages, Naveed Khan, Clio Der Sarkissian, Xuexue Liu, Stefanie Wagner, Beth Ginondidoy Leonard, Bruce L. Manzano, Nancy OMalley, Jennifer A. Leonard, Elosa Bernldez-Snchez, Eric Barrey, La Charliquart, Emilie Robbe, Thibault Denoblet, Kristian Gregersen, Alisa O. Vershinina, Jaco Weinstock, Petra Raji ikanji, Marjan Mashkour, Irina Shingiray, Jean-Marc Aury, Aude Perdereau, Saleh Alquraishi, Ahmed H. Alfarhan, Khaled A. S. Al-Rasheid, Tajana Trbojevi Vukievi, Marcel Buric, Eberhard Sauer, Mary Lucas, Joan Brenner-Coltrain, John R. Bozell, Cassidee A. Thornhill, Victoria Monagle, Angela Perri, Cody Newton, W. Eugene Hall, Joshua L. Conver, Petrus Le Roux, Sasha G. Buckser, Caroline Gabe, Juan Bautista Belardi, Christina I. Barrn-Ortiz, Isaac A. Hart, Christina Ryder, Matthew Sponheimer, Beth Shapiro, John Southon, Joss Hibbs, Charlotte Faulkner, Alan Outram, Laura Patterson Rosa, Katelyn Palermo, Marina Sol, Alice William, Wayne McCrory, Gabriella Lindgren, Samantha Brooks, Camille Ech, Ccile Donnadieu, Olivier Bouchez, Patrick Wincker, Gregory Hodgins, Sarah Trabert, Brandi Bethke, Patrick Roberts, Emily Lena Jones, , and Ludovic Orlando Science, 379 (6639), . DOI: 10.1126/science.adc9691 Making a horse culture Horses evolved in North America and dispersed to Eurasia across the Bering Land Bridge. They continued to evolve and were domesticated in Eurasia, but, as far as we know, they became extinct in North America by the late Pleistocene and were then reintroduced by European colonizers. Taylor et al. looked at the genetics of horses across the Old and New Worlds and studied archaeological samples. They found no evidence for direct Pleistocene ancestry of North American horses, but they did find that horses of European descent had been integrated into indigenous cultures across western North America long before the arrival of Europeans in that region. --SNV View the article online https://www.science.org/doi/10.1126/science.adc9691 Permissions https://www.science.org/help/reprints-and-permissions Downloaded from https://www.science.org on March 31, 2023 Please enable JavaScript to view the comments powered by Disqus. Discussion The genus Equus is a genus of mammals that includes horses, asses, and zebras. All species are herbivores, and most are grazers. ! [Taxonomic tree](https://imgur.com/bq1yWyo.png) For more: https:// en.wikipedia.org/wiki/Equus_(genus) Bayesian radiocarbon dating combines radiocarbon dating with Bayesian statistical analysis to obtain more accurate and refined estimates of the age of artifacts or samples. It incorporates additional information and uncertainties to provide a more precise range of possible ages, taking into account factors like calibration curves, measurement errors, and prior knowledge. The method is particularly useful in complex contexts where multiple sources of evidence need to be considered to obtain reliable age estimates. For more on the method: https:// www.cambridge.org/core/journals/radiocarbon/article/ bayesian-analysis-of-radiocarbon-dates/ F622173B70F9C1597F2738DEFC597114 > "Our archaeological analyses show the dispersal of domestic horses from Spanish settlements. in the American Southwest to the northern Rockies and central Great Plains by the first half of the 17th century CE at the latest. They provide evidence of local raising and veterinary care of horses, likely foddering with domestic maize, and use of horses in transport by Indigenous peoples by this time. A directly dated radiocarbon specimen from Paa'ko Pueblo in northern New Mexico shows that horses reached the region via Indigenous groups before Spanish colonization of the American Southwest, as previously hypothesized (44, 45). Moreover, our new temporal framework shows that horses were present across the plains long before any documented European presence in the Rockies or the central plains. " This research was a collaboration between archaeologists, geneticists, and scientists/historians from the Lakota, Comanche and Pawnee nations. > "Despite representing a major source for understanding the timing and ways in which horses were managed, ridden, and integrated into early societies, archaeological remains of domestic horses from Indigenous contexts are also overlooked (24). In this study, we ex- tensively surveyed existing archaeological collections to identify early historic horse specimens with potential for reconstructing early human-horse relationships across the American Southwest and Great Plains (Fig. 1). Together, DNA, archaeozoological, and stable isotope data support the introduction of Spanish-sourced domestic horses into Indigenous societies across the plains before the first half of the 17th century CE." > "Horses evolved in North America and dispersed to Eurasia across the Bering Land Bridge. They continued to evolve and were domesticated in Eurasia, but, as far as we know, they became extinct in North America by the late Pleistocene and were then reintroduced by European colonizers. Taylor et al. looked at the genetics of horses across the Old and New Worlds and studied archaeological samples. They found no evidence for direct Pleistocene ancestry of North American horses, but they did find that horses of European descent had been integrated into indigenous cultures across western North America long before the arrival of Europeans in that region." -SNV in original Science article: https://www.science.org/doi/10.1126 /science.adc9691 > "Our findings have deep ramifications for our understanding of social dynamics in the Great Plains during a period of disruptive social changes for Indigenous peoples. The area of southwestern Wyoming including Blacks Fork is considered to be a homeland for the Shoshonean-speaking ancestral Comanche who migrated to the southern plains of Texas, New Mexico, Colorado, and Oklahoma before the early 18th century CE (49, 50). The drive to acquire horses from Spanish New Mexico is often cited as a likely mechanism for this transcontinental movement (50). However, our new data--which align with some Comanche and Shoshone oral accounts (51) (materials and methods section 7)--suggest that ancestral Comanche had already integrated horse raising, ritual practices, and transport into their lifeways at least a full half century before their southward migration, effectively moving to the southern plains as horse herders. " Phylogeography is a field of study that combines principles from evolutionary biology and biogeography to investigate the historical processes that have shaped the geographical distribution of species and their genetic variation. It seeks to understand the evolutionary history of species and populations by analyzing their genetic data in the context of geographic patterns. Phylogeography examines the spatial distribution of genetic lineages within and among populations, aiming to uncover the historical events that have influenced their divergence and dispersal. By studying the genetic variation of organisms, researchers can infer the movement of species across geographic regions, the formation of populations, and the effects of past climatic and environmental changes on species' distributions. This discipline utilizes molecular genetic techniques, such as DNA sequencing, to compare genetic data from different populations or species. It also incorporates information from other fields, including paleontology, geology, and climatology, to reconstruct the evolutionary and ecological processes that have shaped the current distribution of organisms. 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