Fermat's Library | Early dispersal of domestic horses into the Great Plains and northern Rockies annotated/explained version.

Phylogeography is a field of study that combines principles from ev...

This research was a collaboration between archaeologists, geneticis...

> "Horses evolved in North America and dispersed to Eurasia across ...

The genus Equus is a genus of mammals that includes horses, asses,...

> "Despite representing a major source for understanding the timing...

Bayesian radiocarbon dating combines radiocarbon dating with Bayesi...

> "Our archaeological analyses show the dispersal of domestic horse...

> "Our findings have deep ramifications for our understanding of so...

RESEARCH ARTICLE

◥

PHYLOGEOGRAPHY

Early dispersal of domestic horses into the Great

Plains and northern Rockies

William Timothy Treal Taylor

1,2

*†, Pablo Librado

†, Mila Hunska Tašunke Icu (Chief Joseph American

Horse)

†, Carlton Shield Chief Gover

1,5

†, Jimmy Arterberry

†, Anpetu Luta Wih (Antonia Loretta

Afraid of Bear-Cook)

, Akil Nujipi (Harold Left Heron)

, Tanka Omniya (Robert Milo Yellow Hair)

Mario Gonzalez (Nantan Hinapan)

, Bill Means

4,8

, Sam High Crane (Wapageya Mani)

‡,

Mažasu (Wendell W. Yellow Bull)

, Barbara Dull Knife (Mah’piya Keyaké Wih)

4,10

Wakihyala Wih (Anita Afraid of Bear)

, Cruz Tecumseh Collin (Wanka’tuya Kiya)

, Chance Ward

2,11

Theresa A. Pasqual

, Lorelei Chauvey

, Laure Tonasso-Calviere

, Stéphanie Schiavinato

Andaine Seguin-Orlando

, Antoine Fages

3,13

, Naveed Khan

3,14

, Clio Der Sarkissian

, Xuexue Liu

Stefanie Wagner

, Beth Ginondidoy Leonard

15,16

, Bruce L. Manzano

, Nancy O’Malley

Jennifer A. Leonard

, Eloísa Bernáldez-Sánchez

, Eric Barrey

, Léa Charliquart

, Emilie Robbe

Thibault Denoblet

, Kristian Gregersen

, Alisa O. Vershinina

, Jaco Weinstock

Petra RajićŠikanjić

, Marjan Mashkour

, Irina Shingiray

28,29

, Jean-Marc Aury

, Aude Perdereau

Saleh Alquraishi

31,32

, Ahmed H. Alfarhan

33,34

,KhaledA.S.Al-Rasheid

, Tajana Trbojević Vukicˇević

Marcel Buric

, Eberhard Sauer

,MaryLucas

, Joan Brenner-Coltrain

,JohnR.Bozell

Cassidee A. Thornhill

, Victoria Monagle

,AngelaPerri

,CodyNewton

,W.EugeneHall

Joshua L. Conver

, Petrus Le Roux

,SashaG.Buckser

, Caroline Gabe

, Juan Bautista Belardi

Christina I. Barrón-Ortiz

, Isaac A. Hart

, Christina Ryder

, Matthew Sponheimer

,BethShapiro

John Southon

,JossHibbs

, Charlotte Faulkner

,AlanOutram

,LauraPattersonRosa

Katelyn Palermo

,MarinaSolé

, Alice William

,WayneMcCrory

, Gabriella Lindgren

57,60

Samantha Brooks

, Camille Eché

, Cécile Donnadieu

, Olivier Bouchez

, Patrick Wincker

Gregory Hodgins

, Sarah Trabert

, Brandi Bethke

, Patrick Roberts

38,66

, Emily Lena Jones

†,

Yvette Running Horse Collin (Tašunke Iyanke Wih)

3,4

†, Ludovic Orlando

†*

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 interdisciplinary 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 before the arrival of 18th-century European

observers, as reflected in herd management, ceremonial practices, and culture.

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, Šungwakaŋ—“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

Šungwaka ŋ 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

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 2.06× 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

amedianboundarydateof~1544CE(Fig.1D

and materials and methods section 2). Various

models provided good measures of agreem ent

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

Department of Anthropology, University of Colorado Boulder, Boulder, CO 80309, USA.

Museum of Natural History, University of Colorado Boulder, Boulder, CO 80309, USA.

Centre for

Anthropobiology and Genomics of Toulouse (CAGT, CNRS UMR5288), University Paul Sabatier, Faculté de Médecine Purpan, 31000 Toulouse, France.

Oglala Lakota, Pine Ridge Reservation, SD

57770, USA.

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.

Biology

Department, College of Science, Taif University, Taif 21944, Saudi Arabia.

Zoology Department, College of Science, King Saud University, Riyadh 12372, Saudi Arabia.

Biology Department,

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.

isoTROPIC

Research Group, Max Planck Institute for Geoanthropology, 07745 Jena, Germany.

Department of Anthropology, University of Utah, Salt Lake City, UT 84112, USA.

Archaeological consultant,

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,

USA.

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

85721, USA.

Department of Anthropology, University of Oklahoma, Norman, OK 73019, USA.

Oklahoma Archeological Survey, University of Oklahoma, Norman, OK 73019, USA.

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

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