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
3
, Mila Hunska Tašunke 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
,
Mažasu (Wendell W. Yellow Bull)
4
, Barbara Dull Knife (Mahpiya Keyaké Wih)
4,10
,
Wakihyala Wih (Anita Afraid of Bear)
4
, Cruz Tecumseh Collin (Wankatuya Kiya)
4
, Chance Ward
2,11
,
Theresa A. Pasqual
12
, Lorelei Chauvey
3
, Laure Tonasso-Calviere
3
, Stéphanie 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 OMalley
18
,
Jennifer A. Leonard
19
, Eloísa Bernáldez-Sánchez
20
, Eric Barrey
21
, Léa Charliquart
22
, Emilie Robbe
23
,
Thibault Denoblet
22
, Kristian Gregersen
23
, Alisa O. Vershinina
24
, Jaco Weinstock
25
,
Petra RajićŠikanjić
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 Trbojević Vukicˇević
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. Barrón-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
,MarinaSolé
57
, Alice William
58
,WayneMcCrory
59
, Gabriella Lindgren
57,60
,
Samantha Brooks
61
, Camille Eché
62
, Cécile 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 (Tašunke 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 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.
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 (35), 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
Lakotahorse 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 humanhorse 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, 13161323 (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
Paako, 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, 13161323 (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, Faculté de Médecine 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
HeSapa 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 Xitan (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, Estación Biológica
de Doñana (EBD-CSIC), 41092 Sevilla, Spain.
20
Laboratorio de Paleontología y Paleobiología, Instituto Andaluz del Patrimonio Histórico, 41092 Sevilla, Spain.
21
Université Paris-Saclay, INRAE,
AgroParisTech, GABI UMR1313, Jouy-en-Josas, 78350 Paris, France.
22
Musée de lArmée, Hôtel 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, Muséum national dHistoire naturelle,
Archéozoologie, Archéobotanique (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 François Jacob, CEA, CNRS, Université dEvry, Université Paris-Saclay, 91000 Évry, 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 Académica Río Gallegos (ICASUR), Laboratorio de Arqueología Dr. Luis A. Borrero, CONICET, 9400 Río 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 Medical 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 Gwetin 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, Génome 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
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.
PrePueblo 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 Paako 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, 13161323 (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.
(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
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, 13161323 (2023) 31 March 2023 4of8
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1
6
0
±
1
65
W
yo
m
in
g
Na
t
u
ra
l
T
r
a
p
2
5
2
1
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1
5
3
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y
om
i
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g
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t
u
r
a
l
T
r
ap
2
53
56
±
1
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7
Wyom
i
ng
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atu
r
al
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r
ap 2493
204
I
d
a
h
o
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aw
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ro
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r 17
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±
9
5
NW3
/
New
Me
xico
NW
32/Kans
as
Bla
c
k
s
F
o
rk
/
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omin
g
M
GM
R293
9
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T33
5
9
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R
T3356
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O
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H
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gen
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/
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1
6
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h
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ct
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72
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Z33
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7
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AKO/
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k
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n
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RT340
7
FO
RT34
0
8
OGLAK
O/Choctaw
81
OGL
AK
O/
Oji
b
w
e
8
OGL
A
K
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i
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1
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K
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L
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n
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GLAKO
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13
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L
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ache17
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GL
A
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7
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L
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R
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7
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9
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T
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AL
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P
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9
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c
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H
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3
7
5
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MR
272
6
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TUR140
SAH
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t
ina
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A
22
N
O
R
I
180
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L
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n11
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STCZ3
332
C
AR
T
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A
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554
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3
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A
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4
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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
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TU
R14
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G
VA1 2
2
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Z
I
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K
Z
T
28
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A
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ICELP5782
VHR0
17
K
Y
R
H
1
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M
A
R
V
ELE18
D
A
TO1
2
JEJU2
J
E
J
U
3
NUST
A
R5
B
A
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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 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,
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
horsesfourthandfifthyearoflife,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, 13161323 (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
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 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
4
-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
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
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 Paako 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, 13161323 (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 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]
RESEARCH | RESEARCH ARTICLE
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
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 datawhich
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
(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 USDAAshley
National Forest. Analysis of Blacks Fork material was completed
with permission from the USDAAshley 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 Unions Horizon 2020 research and
innovation programme (grant agreement 681605). Additional
funding was provided by the European Unions Horizon 2020
research and innovation programme under the Marie Skłodowska-
Curie (grant agreement 890702-MethylRIDE); the French
Government Investissement dAvenir 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
study are included in table S1. License information: Copyright ©
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 (53124)
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
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Copyright © 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
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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.