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...

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> "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...

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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

Downloaded from https://www.science.org on March 31, 2023

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

(Galiceño 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 specimens

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.

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

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

GLA

O/P

ebl

an2

T33

OGL

AKO/C

ST10

1096

CHI

iute54

ica

alls 3

T331

RT3

344

T334

R2726

GLAKO

w73

GLA

taw

OGL

/Pa

OGLAKO

/Sa

332

TCZ3

ctaw7

ing

atu

p 2

Wyom

atu

ap 2493

0±

204

ppe

r 17

NW3

New

xico

32/Kans

Bla

omin

R293

FOR

T33

T3356

355

L28

NW10

Nebr

gen

OGLAKO

Ute

KO/

AKO/Cho

RT3

OGL

KO/

win

STCZ332

Z33

OGL

AKO/

RT340

RT34

OGLAK

O/Choctaw

OGL

Oji

OGL

loa

ako

AKO/U

2/A

entin

OGLA

GLAKO

ache

KO/Ap

ache17

Ojibw

ive

/Ka

OGLAK

O/O

CHICO

OGL

KO/

cta

w78

TCZ3

328

LAKO/Choc

taw3

RT33

WURTB

W01

HOL

OLS

MO1

YIL

310

TR225

FRMO1

M441

AKTK001

AKO/Athab

HANO0

NO02

HANO03

WSTF02

VELE21

OTE2

NB44

HR0

R037

MONG

PR40

ZNK1002_4

BES

LR18_16

17_1

LR18

N131

CGG101

AI18_26

PRZW5

US37

GRAL9

KSK16232

IN3

G303

US38

NW3

Mexi

BELG

EP14

272

MGPL2865

TUR140

SAH

4/Argen

ina

INV

180

AKO/iS

n11

AFL0004

STCZ3

332

554

CART

T53

MAR

B948

ARAB

RAK02

RAK

OGLAKO/

Ojibwe7

MZR1

UK1

NW26/Oklahoma

DATO3

CGG1

1397

SSYK

KYR

EP2

ONT

VHR102

R14

RVELE

VA1 2

VH2

ICELP5782

VHR0

ELE18

TO1

JEJU2

NUST

PSKA

AA1

Bootstrap

92.5

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)

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 genetic ancestry

profiles (right). The ancestry proportions for K = 7 genetic components 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

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,

Sr/

Sr = 0.70970)

and postnatal (M1,

Sr/

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 (

Sr/

Sr = 0.70905 near the crown,

versus

Sr/

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

O) and carbon (d

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*

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

T1099

GMR2726

GMR293

TCZ332

TCZ3328

TCZ338

TCZ3332

CA335

CA3356

OGLAKO/Paiute5

GLAKO/Athabascan58

GLAKO/Paiute68

OGLAKO/Choctaw7

OGLAKO/Choctaw72

GLAK

/Choctaw7

OGLAKO/Choctaw8

OGLAKO/Chickasaw83

OGLAKO/Choctaw8

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

−

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

−20246

Z-score f

(Historic NA, Modern NA; JAL5886, WELS)

Avg Z-score = 2.31

95% CI

0.001 0.002 0.003 0.004 0.005

(S−URAL,X; CPONT,ICEL)

(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 sources. Models

with highest P values are shown where multiple models could not be rejected.

Asterisks depict two samples for which the ancestry profiles presented correspond

to the second best qpAdm model (table S3), while the cross accompanying

OGLAKO/Ojiwbe2 indicates that its best qpAdm model involves Galiceño and

Icelandic horses as donor populations, instead of British sources (supplementary

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

-statistics in the form of (S-URAL, X; C-PONT, Jal5885 or Welsh), where X

represents historic and/or modern horses from North America, and C-PONT 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 genetic

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 introgression

from modern Icelandic horses, ruling out genetic contribution from relict populations

of Viking horses (supplementary materials section 6). Error bars correspond to

twice the standard error estimated from jackknifing. (E) Distribution of Z-scores for

-statistics of the form (Historic North American Horse, Modern North American

Horse; Jal5885, Welsh).

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locally (Blacks Fork) or within a territory ex-

tending even farther away from the European

colonial sphere (Kaw).

A d

C spike toward the end of the sampled

section of the Kaw molar shows a strong input

of C

-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

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

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 temporomandibular 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 healing at 50×

magnification. (2) Ossification of the nuchal ligament, indicating use in transport or

confinement and/or tethering. (C) Modeled northward dispersal of horses. Model

based on archaeological discoveries and consideration of historically documented

cultural developments and migrations. (D) Isotopic evidence for foddering. Stable

carbon and oxygen and strontium isotope sampling locations for the Kaw River

horse, as measured in millimeters from the root (lower-right third molar). Sampling

locations are represented on the x axis as the midpoint of a 2-mm section (e.g.,

the section from 0 to 2 mm is shown as “1” on the graph). Filled triangles show

strontium isotope values, with stable carbon values represented by filled circles, and

stable oxygen values as open circles. [Kaw horse image credit: E. Scott]

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Downloaded from https://www.science.org on March 31, 2023

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

–

), 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 wé

(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. Dr. Antonia Loretta

Afraid of Bear-Cook thus reminds us that the

current genetic makeup of the horse does

not change the Lakota responsibility toward

Šungwakaŋ. In fact, for the Lakota and other

Indigenous peoples with deep ancestral rela-

tionships with the horse, their evolution was

not to be feared or controlled, but rather some-

thing to be respected. Protecting horses now,

regardles s of their origins, is in fact protecti ng

the Lakota and other Indigenous lifeways. A

commitment to protecting these is a com-

mitment to protect all life.

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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. Kjær 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 Skłodowska-

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 Skłodowska 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.Š., 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.Perr.,

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 this

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

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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

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Discussion

This research was a collaboration between archaeologists, geneticists, and scientists/historians from the Lakota, Comanche and Pawnee nations. 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) > "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." 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. " > "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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