ArticlePDF Available

Abstract and Figures

Advances in the isolation and sequencing of ancient DNA have begun to reveal the population histories of both people and dogs. Over the last 10,000 y, the genetic signatures of ancient dog remains have been linked with known human dispersals in regions such as the Arctic and the remote Pacific. It is suspected, however, that this relationship has a much deeper antiquity, and that the tandem movement of people and dogs may have begun soon after the domestication of the dog from a gray wolf ancestor in the late Pleistocene. Here, by comparing population genetic results of humans and dogs from Siberia, Beringia, and North America, we show that there is a close correlation in the movement and divergences of their respective lineages. This evidence places constraints on when and where dog domestication took place. Most significantly, it suggests that dogs were domesticated in Siberia by ∼23,000 y ago, possibly while both people and wolves were isolated during the harsh climate of the Last Glacial Maximum. Dogs then accompanied the first people into the Americas and traveled with them as humans rapidly dispersed into the continent beginning ∼15,000 y ago. archaeology | genetics | domestication | dogs | peopling of the Americas
Content may be subject to copyright.
Dog domestication and the dual dispersal of people
and dogs into the Americas
Angela R. Perri
, Tatiana R. Feuerborn
, Laurent A. F. Frantz
, Greger Larson
, Ripan S. Malhi
, David J. Meltzer
, and Kelsey E. Witt
Edited by Theodore G. Schurr, University of Pennsylvania, Philadelphia, PA, and accepted by Editorial Board Member Dolores R.
Piperno December 8, 2020 (received for review June 4, 2020)
Advances in the isolation and sequencing of ancient DNA have begun to reveal the population histories of
both people and dogs. Over the last 10,000 y, the genetic signatures of ancient dog remains have been
linked with known human dispersals in regions such as the Arctic and the remote Pacific. It is suspected,
however, that this relationship has a much deeper antiquity, and that the tandem movement of people and
dogs may have begun soon after the domestication of the dog from a gray wolf ancestor in the late
Pleistocene. Here, by comparing population genetic results of humans and dogs from Siberia, Beringia,
and North America, we show that there is a close correlation in the movement and divergences of their
respective lineages. This evidence places constraints on when and where dog domestication took place.
Most significantly, it suggests that dogs were domesticated in Siberia by 23,000 y ago, possibly while
both people and wolves were isolated during the harsh climate of the Last Glacial Maximum. Dogs then
accompanied the first people into the Americas and traveled with them as humans rapidly dispersed into
the continent beginning 15,000 y ago.
peopling of the Americas
Dogs were the first domesticated species and the only
animal known to enter into a domestic relationship
with people during the Pleistocene (14). Recent ge-
netic analyses of ancient dog remains and of the ar-
chaeological and genetic records of ancient people
have demonstrated that the spatiotemporal pattern-
ing of specific dog mitochondrial lineages are often
correlated with the known dispersal of human groups
at different times and places.
For instance, a study of mitochondrial signatures
derived from ancient Near Eastern and European
dogs has demonstrated that a specific haplogroup
arrived in Europe as dogs dispersed out of the Near
East along with farmers (5). The first dogs to arrive in
New Zealand did so with newly arriving Polynesians
(6). People and dogs also dispersed together in the
North American Arctic, where dogs carrying a specific
mitochondrial DNA (mtDNA) signature (haplogroup
A2a; refs. 7 and 8) accompanied Paleo-Inuit groups
as they moved into the region 5,000 y ago (5 ka).
Subsequently, the arrival of Inuit groups into the same
region 1 ka was accompanied by the introduction of
a dog population that carried novel mtDNA signatures
(A1a and A1b; ref. 7).
These correlations between the dispersals of peo-
ple and specific dog lineages may have begun much
Department of Archaeology, Durham University, Durham DH1 3LE, United Kingdom;
GLOBE Institute, University of Copenhagen, 1350
Copenhagen, Denmark;
The Qimmeq Project, University of Greenland, 3905 Nuussuaq, Greenland;
Archaeological Research Laboratory,
Department of Archaeology and Classical Studies, Stockholm University, 114 19 Stockholm, Sweden;
Department of Bioinformatics and Genetics,
Swedish Museum of Natural History, 114 18 Stockholm, Sweden;
Centre for Palaeogenetics, 114 18 Stockholm, Sweden;
Palaeogenomics Group,
Department of Veterinary Sciences, Ludwig Maximilian University, Munich D-80539, Germany;
School of Biological and Chemical Sciences, Queen
Mary University of London, London E1 4NS, United Kingdom;
The Palaeogenomics and Bio-Archaeology Research Network, Research Laboratory
for Archaeology and History of Art, University of Oxford, Oxford OX1 3QY, United Kingdom;
Department of Anthropology, University of Illinois at
UrbanaChampaign, Urbana, IL 61801;
Carl R. Woese Institute for Genomic Biology, University of Illinois at UrbanaChampaign, Urbana, IL 61801;
Department of Anthropology, Southern Methodist University, Dallas, TX 75205;
Lundbeck Foundation GeoGenetics Centre, GLOBE Institute,
University of Copenhagen, 1350 Copenhagen, Denmark;
Department of Ecology and Evolutionary Biology, Brown University, Providence, RI
02912; and
Center for Computational and Molecular Biology, Brown University, Providence, RI 02912
Author contributions: A.R.P., L.A.F.F., G.L., D.J.M., and K.E.W. designed research; A.R.P., T.R.F., L.A.F.F., G.L., R.S.M., D.J.M., and K.E.W. per-
formed research; A.R.P., T.R.F., L.A.F.F., G.L., R.S.M., D.J.M., and K.E.W. analyzed data; and A.R.P., T.R.F., L.A.F.F., G.L., R.S.M., D.J.M., and K.E.W.
wrote the paper.
The authors declare no competing interest.
This article is a PNAS Direct Submission. T.G.S. is a guest editor invited by the Editorial Board.
Published under the PNAS license.
To whom correspondence may be addressed. Email:,, or
This article contains supporting information online at
Published January 25, 2021.
PNAS 2021 Vol. 118 No. 6 e2010083118
Downloaded at MPI f. evolutionäre Anthropologie on January 25, 2021
earlier, perhaps soon after dogs became domesticated from a
gray wolf ancestor in Eurasia, though precisely where and how
many times that process took place remains unknown. The
archaeologically documented presence of dogs in the Americas
by at least 10 ka (9) suggests that dogs accompanied the early
human groups who moved from northeast Asia across the Bering
Land Bridge (Beringia) into the Americas. On the basis of current
archaeological and genetic evidence, this movement likely took
place before 15 ka (10, 11). Here, we take advantage of this re-
cord and newly available evidence from humans and dogs in late
Pleistocene Siberia, Beringia, and North America to assess the
likelihood that the first peopletoreachanddisperseacross
the Americas did so in tandem with their dogs. This analysis al-
lows us to better understand that dispersal process and pre-
sent a hypothesis for the temporal and geographic origins of
domestic dogs.
The First Dogs
Numerous archaeometric approaches have been applied to
document the interaction between wolves, dogs, and people in
order to establish the time frame and geography of dog domes-
tication. These studies have shown, first, that dogs were the ear-
liest animal domesticated, and the only species that entered into a
domestic relationship with people during the Pleistocene (1, 2, 12,
13). Second, the specific wolf population from which dogs derived
appears to be extinct (1, 14, 15). Finally, genetic and archaeo-
logical evidence from modern and ancient dogs and wolves
demonstrates that dog domestication took place in Eurasia
(1618). Many other aspects of dog domestication, including the
circumstances under which the relationship began, the time
frame, and the number and location(s) of potential independent
domestication regions, remain unresolved (1921).
The shift in humanwolf interactions that led to domestication
has been addressed through a wide variety of approaches, and
there are ongoing debates over when the first recognizably do-
mestic dog appeared. The earliest generally accepted dog dates
to 15 ka (from the site of Bonn-Oberkassel, discussed below).
However, claims for the existence of domestic dogs as early as 40
ka (2228) have been made on the basis of morphological (22,
2427), isotopic (22, 29), genetic (22, 28, 30), and contextual as-
sessments (24, 31) of ancient canid remains. Yet, none of these
potential domestication markers is fail-safe, owing to the fact that
wolves and early domesticated dogs can be difficult to distinguish
from each other.
For example, common morphological markers used to identify
domestication such as tooth crowding, skull size, and reduced
snout length often fail to clearly distinguish dogs from wolves
(3236). Isotopic signatures, and their dietary inferences, have
been used to identify early dogs, though these have also been
questioned given that the isotopic variation is often consistent
with wolf diets (3). In addition, genetic analyses of these proposed
early dogs have demonstrated that they do not belong to the
same lineages as ancient or modern dogs (30, 37). While genomic
estimates from multiple studies place the split time within wolf
lineages, including the one that ultimately gave rise to dogs, to
between 40 and 27 ka (14, 38), this timing is unlikely to reflect
the initiation of the domestication process (1, 39). Finally, sites
with purported domestic dogs have been questioned based on an
absence of carnivore gnawing (40) or pups (41), features taken to
indicate the presence of a living, breeding dog population.
Accordingly, claims for domestic dogs at the Belgian site of
Goyet (22, 42, 43) have been disputed since their cranial and
dental morphologies do not exclude the possibility that they are
wolves (3, 3236, 44, 45). mtDNA analyses of these canids also
showed that they belong to an ancient European wolf lineage that
is genetically highly divergent from any dog haplogroup (30).
Similarly, genetic analyses of proposed Paleolithic dogs from the
sites of Ulakhan Sular (23), Tumat (46), Razboinichya (27), Berelekh
(23), Kostenki 8 (23), and Eliseevichi (25) have shown these canids
to be more closely related to ancient and modern wolves than
they are to dogs (30, 37, 47).
At least one of the purported dogs from the Czech site of
ı(24, 48) has also shown a genetic affinity to wolves
over dogs (37). The domestic designation of these canids has
likewise been questioned based on analyses of their dental
and cranial morphologies (12, 3236, 44, 45). Isotopic (29)
and dental microwear (31) analyses of the diet of the pro-
posed dogs at the site may also fallwithintherangeofvaria-
tion of the local wolf populations (12). Additionally, P
lacks evidence for carnivore gnawing or pups (40, 41), raising
further questions about the presence of a dog population at
the site.
The challenge for all claims of late Pleistocene dogs has been
to show conclusively, across several lines of evidence, that
the specimen(s) in question can be clearly distinguished from
contemporaneous wolves (3). Here, we take a conservative ap-
proach and only include those canids whose taxonomic status is
unambiguously domestic.
The earliest generally accepted remains of a domestic dog,
based on a convergence of morphological, genetic, isotopic, and
contextual evidence, comes from the site of Bonn-Oberkassel in
Germany, dated to 15 ka (4, 30) (Fig. 1). The morphology and
genetics of this young dog clearly distinguish it from local wolves.
Its coburial with humans and the evidence for its care after suf-
fering an illness also suggest it was a dog. Claims for contempo-
raneous domestic dogs have also been made at sites in France
(49), Germany (50), Israel (51), Italy (52), and Switzerland (53).
Based on their morphology and context, additional poten-
tial dogs may be present at Pleistocene Siberian sites such as
Afontova Gora, Diuktai Cave, and Verkholenskaia Gora (9, 23),
although their status has yet to be established. In the Americas,
the earliest confirmed archaeological dog remains, based on
combined morphological, genetic, isotopic, and contextual evi-
dence, are from the Koster and Stilwell II sites, which have been
dated to 10 ka (9, 16).
From a genetic perspective, hundreds of ancient and modern
canid mitochondrial and nuclear genomes have been sequenced.
Analyses of the nuclear data indicate that all dogs represent a
genetically homogeneous group that possesses varying degrees
of ancestry from three major ancestral lineages: a western Eur-
asian lineage (mostly found in European, Indian, and African
dogs); an east Asian lineage (e.g., dingoes); and an Arctic lineage
(e.g., huskies and ancient American dogs) (16). A recent study of
dozens of ancient dog genomes suggests that these lineages
were all established by at least 11 ka (54).
mtDNA data indicate that the vast majority of modern dogs fall
into one of four monophyletic haplogroups (A, B, C, or D), with the
majority belonging to haplogroup A (SI Appendix, Fig. S1). Recent
ancient DNA studies demonstrated that all precontact dogs in the
Americas south of the Arctic possess a unique mitochondrial
haplogroup (A2b) that is nested within mitochondrial haplogroup
A and which has since virtually disappeared (0.5%; ref. 16) in
modern dogs inside and outside of the Americas (1618, 55).
Within A2b are four additional and well-supported monophyletic
PNAS Perri et al. Dog domestication and the dual dispersal of people and dogs into the Americas
Downloaded at MPI f. evolutionäre Anthropologie on January 25, 2021
subhaplogroups, A2b1 through A2b4 (16), which are only found in
the Americas. Of those four haplogroups, A2b1 is pan-American
(possessed by dogs from Californias Channel Islands to Argen-
tina), while the other three haplogroups are more geographically
restricted, given current data (Fig. 2).
Molecular clock analyses have revealed the timing associated
with splits within haplogroup A (SI Appendix, Fig. S1). First, the
split between lineages A1b and A2 at the base of the haplogroup
is estimated to date to 22.8 ka (95% CI 26 to 19.7 ka; ref. 7). This
timing, which represents the oldest known coalescence between
Fig. 1. Lineages and estimated population divergence times (hexagons) for dogs (Top, in red), and humans (Bottom, in blue), for the period from
24,000 to 10,000 y ago. Divergence times are shown as point estimates. Confidence intervals for the respective point estimates are shown in
the horizontal red bars (dogs) and bluebars (humans) in the center of the figure, which shows as well the span of the LGM, 23,000 to 19,000 y ago.
Perri et al. PNAS
Dog domestication and the dual dispersal of people and dogs into the Americas
Downloaded at MPI f. evolutionäre Anthropologie on January 25, 2021
two dog mitochondrial lineages, suggests that dogs were do-
mesticated several thousand years prior to their first appearance
in the archaeological record. This estimate may have been af-
fected by later gene flow between wolves and dogs, though this
possibility is less likely since these haplogroups have not been
found in any ancient or modern wolves, and because wolfdog
admixture appears to have been uncommon (54). Regardless, this
suggested early time frame indicates that dogs were likely do-
mesticated by the time humans crossed into the Americas.
The First People in the Americas
Our understanding of the origins and antiquity of the first people
to reach the Americas has advanced significantly over the past
decade through the identification of new archaeological sites and
the generation of ancient human genomes from individuals in the
Americas and northeast Asia.
Genomic evidence indicates that Native American ancestry
can be traced to a population that is currently estimated to have
diverged from an East Asian ancestor 30 ka (95% CI 36.4 to 26.8
ka; ref. 56) (all age estimates in this section are based on nuclear
DNA, except where noted). Around 24 ka (95% CI 27.9 to 20.9 ka
based on nuclear data and 24.9 to 18.4 ka based on mitochondrial
data; ref. 57), that population then split into at least two groups.
One group identified as Ancient Paleosiberians (APS) appears to
have remained in far northeast Asia, while the other group be-
came the basal branch of Native Americans (56). Both groups
subsequently, and separately, received gene flow (24 ka; Fig. 1)
from Ancient North Siberians (ANS), a population whose ances-
tors are detected archaeologically at the Yana RHS site in far
northern Siberia (31.6 ka; Fig. 2) and the Malta site near Lake
Baikal (24 ka; Fig. 2; ref. 56).
From the time of the Last Glacial Maximum (LGM, 23 to 19
ka), the basal branch of Native Americans appears to have been
isolated in northeast Asia, where they remained before departing
for the Americas. There is currently no evidence of subsequent
gene flow with other populations in the region, though this does
not preclude the possibility of interaction with other groups that is
not visible genetically. This period of isolation, by virtue of where
it is suspected to have occurred, is known as the Beringian
Standstill (58). Estimates of its duration based on nuclear DNA, Y
chromosome, and mtDNA vary, but overall they indicate this
episode may have lasted as little as 2,400 y to as long as 9,000 y
(10, 5759).
During this period of isolation, current evidence indicates that
perhaps 21 ka (95% CI 21.9 to 18.1 ka) this basal branch split into
at least two distinctive populations: Ancient Beringians (AB) and
Ancestral Native Americans (ANA) (60). Both the AB and ANA
populations crossed into eastern Beringia (present-day Alaska)
after their split, though the timing of their dispersal(s), whether
they moved simultaneously, and how long they may have main-
tained a degree of gene flow once they diverged from one an-
other remain unclear (60, 61). Although both populations reached
Alaska, to date no genomic evidence of the AB has been found
south of Alaska, or in any Alaskan populations after 9 ka (the
radiocarbon age of the AB individual from the Trail Creek Cave
site, Alaska; Fig. 2; ref. 10).
The ANA lineage, on the other hand, reached North America
south of the continental ice sheets, after diversifying 15.7 ka
(95% CI 17.5 to 14.6 ka; ref. 60) into northern (NNA) and southern
(SNA) branches (62). The NNA/SNA split must have taken place
well after ANA had physically separated from the population
represented by the AB lineage (57, 60), since NNA and SNA
groups are genetically equidistant to AB (otherwise, one or the
other of the groups would have been closer to AB). It is inferred
that this split took place as the ANA were moving south
from Alaska.
The estimated time of the split, and the archaeological evi-
dence for people in the Americas 15 ka (e.g., refs. 11 and 61),
implies that the route to the Americas south of the continental ice
sheets must have been along the Pacific coast. The alternativean
interior route between the ice sheets (the ice-free corridor)had
yet to open and did not yet support the plant and animal re-
sources necessary for human foragers (11, 63, 64). How much
earlier people may have arrived in the Americas is unclear:
Archaeological evidence provides only a minimum age, since the
oldest sites that have been found are not the oldest on the conti-
nent (61). Genetic estimates provide a maximum value, since
the peopling process must postdate the basal split of Native
Afontova Gora
Yana RHS
Trail Creek Cave Dog Haplogroup
Unknown canid
Human Ancestry
Ancient North Siberian (ANS)
Ancient Beringian (AB)
Fig. 2. A map depicting the sites and lineages mentioned in the text. Human and dog lineages are denoted (see references in the text).
PNAS Perri et al. Dog domestication and the dual dispersal of people and dogs into the Americas
Downloaded at MPI f. evolutionäre Anthropologie on January 25, 2021
Americans, which could have been as early as 27.9 ka. It is note-
worthy, however, that there are no archaeological sites in the
Americas that can be shown to securely predate or were occu-
pied during the LGM.
Once people arrived south of the ice sheets, the NNA branch
appears to have had a relatively limited geographic spread. The
SNA branch, however, radiated throughout the hemisphere, and
as they dispersed they diverged genetically, starting 14.1 ka
(95% CI 14.9 to 13.2 ka; refs. 10, 62, and 65).
Reconciling Lineage Branching in Late-Pleistocene Humans
and Dogs
The Americas were one of the last regions of the world to be
settled by people, and based on the antiquity of dogs in North
America it is possible that the first people had canine companions
with them when entering this new landscape (9, 66). Dogs were
part of a larger cultural repertoire that may have assisted humans
in rapidly dispersing into and throughout the Northern Hemi-
sphere (67). Though people could have come to the Americas
without them, dogs must have entered along with people. It is
also reasonable to assume that, when human populations split
from one another, they took their dogs with them. Thus, by
aligning their respective population splits (Fig. 1), we can identify
the timing of their tandem late-Pleistocene movements.
Perhaps not surprisingly, there are challenges in comparing
divergence estimates obtained for dogs and humans. The dates
relevant to the introduction of dogs in the Americas were derived
from mitochondrial data and represent ancestral coalescence
events that predate population splits, especially if the ancestral
population was large. All ancient American dogs outside of the
Arctic, however, belong to the same lineage (A2b) which coa-
lesces with an ancient Siberian dog lineage 16.4 ka (95% CI 18.6
to 14.3 ka; refs. 7 and 16) (SI Appendix, Fig. S1). Although this
time provides an upper bound for the introduction of dogs in the
Americas, evidence for a population bottleneck associated with
the founding of the ancient American dog lineage (18) suggests
that this age may, in fact, be close to the population split between
ancient Siberian and American dogs. The deepest coalescence
event among A2b lineages dates back to 15 ka (95% CI 16.9 to
13.4 ka; ref. 7). Given that the A2b haplogroup is virtually absent
from outside of the Americas, it is likely that its deepest coales-
cence took place in the ancestral population of American dogs.
This time can, thus, be interpreted as a lower bound for diver-
gence between American and Siberian dogs.
This time frame is remarkably consistent with that of the first
peopling of the Americas (Fig. 1) and there are several key di-
vergence nodes in common. First, the deepest coalescence event
among dogs at 26 to 19.7 ka (16) is contemporaneous with the
split between APS and ANA/AB at 27.9 to 20.9 ka (56). This
correspondence suggests that dogs were already domesticated
around the time ANA ancestry was established. Second, the co-
alescence of the A2b (American dog) and A2a (Siberian and
American Arctic dog) lineages, at 18.6 to 14.3 ka (7, 16), and the
deepest coalescence event within A2b, at 16.9 to 13.4 ka (7),
overlap with the split time between the two major Native Ameri-
can lineages (NNA/SNA), at 17.5 to 14.6 ka (60). This indicates
that the radiation of the major human and dog lineages in the
Americas was contemporaneous, suggesting that they diverged
in tandem. This evidence, combined with the antiquity of dog
remains in North America (10 ka; ref. 9), and the lack of later
human migration into the Americas until the early to middle Ho-
locene (between 9 and 5 ka; ref. 68), indicates that dogs crossed
Beringia during the Pleistocene and were present south of the
continental ice sheets by the time the A2b lineage radiated at 15
ka, coincident with the widespread and rapid dispersal of the
SNA lineage.
Either ANA or AB could have brought dogs into the Americas
since there is archaeological evidence supporting the notion that
both groups crossed the land bridge. However, they may not have
crossed at the same time. AB are associated with a distinctive
microblade/microcore stone tool technology (69, 70) seen at the
eastern Siberian site of Diuktai Cave dated to 16.8 ka, from
which it spread northeastward into western Beringia, ultimately
reaching Alaska by 14.2 ka (site of Swan Point; ref. 71). Yet, by
that time people using a very different technology had already
been in the Americas south of the continental ice sheets for more
than a millennium and had begun to disperse throughout the
hemisphere (10). Although caution is always appropriate when
drawing links between stone tool traditions across space and time
(similar tools readily occur as a result of convergence), or between
stone tools and human populations, this evidence suggests that,
by the time AB arrived in Alaska ANA and dogs had already
passed through that region. This, in turn, implies that ANA were
the people who first brought dogs into the Americas.
Dog Domestication in Siberia
These parallels in the population divergences of humans and
dogs place constraints that allow us to reevaluate previously
proposed narratives for the origins of dogs and suggest a hy-
pothesis for the timing and geographic location of dog domes-
tication. On one hand, the split time estimated between wolf
lineages, including the one that gave rise to dogs, provides an
upper bound for domestication at 40 ka (14, 38). On the other
hand, since we established that dogs likely crossed Beringia with
the initial human arrivals, the archaeological evidence of people in
the Americas by 15 ka (11, 72, 73) provides a lower bound for
dog domestication. Combined with evidence that indicates that
dogs were not domesticated in the Americas (16), this points to
dogs having been present in Siberia prior to 15 ka.
Ancient human genome studies have identified multiple ge-
netically divergent groups that were in Siberia/western Beringia
within that time frame. This includes the ANS, APS, and the basal
branch of Native Americans, which, after 21 ka, split into ANA
and AB (Fig. 1). Ancient genomic data indicates that there was no
significant gene flow among these Siberian groups after 23 ka,
or with groups from outside Siberia from 39 ka (56). During this
same period there is also a paucity of archaeological sites in arctic
and subarctic Siberia and Beringia (71, 74, 75). Together, this
evidence suggests that human populations in the region must
have been small and living in relative isolation. They appear to
have remained so up to the time when ANA and AB (separately)
crossed into the Americas.
This evidence for little to no interaction with communities
outside of Siberia raises the question of how ANA acquired the
dogs that accompanied them into the Americas. One possible
explanation is that dogs were domesticated from a wolf pop-
ulation somewhere in Siberia or western Beringia during the late
Pleistocene, and before ANA crossed into the Americas. Previous
studies have suggested on the basis of genetic evidence that
dogs became domesticated in either East Asia (76), Europe (30),
Central Asia (77), or in more than one of these locations inde-
pendently (39). If dogs were domesticated in western Eurasia,
then their spread eastward into Siberia would have required a
far-ranging movement of people. Although possible, this seems
Perri et al. PNAS
Dog domestication and the dual dispersal of people and dogs into the Americas
Downloaded at MPI f. evolutionäre Anthropologie on January 25, 2021
unlikely given that western and eastern Eurasian human populations
had already diverged 39 ka (95% CI 45.8 to 32.2 ka) (56, 75, 78).
Any of the groups known to have been in Siberia during the
LGM (ANA, AB, APS, ANS, and their ancestral lineages; Fig. 1)
may have domesticated dogs. Dogs associated with ANA, how-
ever, do not represent a basal lineage but instead cluster with
Arctic dogs (7, 16, 67), suggesting they were not the initial do-
mesticated population. Likewise, although APS could have
domesticated dogs, there is no genomic evidence of their inter-
action with ANA (though meetings could have taken place and not
been recorded either archaeologically or genetically). Similarly,
dogs may have been domesticated by AB, but there is currently
no genetic evidence of their interacting with ANA. Nevertheless,
dogs could have been domesticated by their shared ancestral
lineage prior to its split at 21 ka.
By process of elimination and for several other reasons, ANS
therefore represents the more likely population to have initiated
the domestication process. For example, genomic analyses of
ANS individuals at the Siberian sites of Malta (24 ka) and
Afontova Gora (17 ka) show evidence for late Pleistocene gene
flow from these populations into both ancient Native American
(Fig. 1) and western Eurasian lineages (79). This provides a
mechanism for the transfer of dogs into different groups and thus
their movement both east and west following their domestication.
Potential late Paleolithic dogs have also been identified at Afon-
tova Gora (80, 81), possibly representing the basal lineage,
though the genomes of these canids have yet to be analyzed. This
scenario also fits with a recent study supporting a single origin for
domestic dogs (54) and reconciles the presence of dogs in
western Eurasia, the Near East, and the Americas by 15 ka.
Dog domestication in Siberia during the LGM provides a
plausible context for the process. Climatic conditions may have
brought human and wolf populations (37) into close proximity
within refugial areas, given their attraction to the same prey
species. Increasing interactions between the two, perhaps
resulting from the mutual scavenging of kills, or from wolves
drawn to the detritus of human campsites (82, 83), may have ini-
tiated a shift in the relationship between the species, eventually
leading to dog domestication. A number of recently identified
potential late-Pleistocene dogs from the region, including those
from Afontova Gora (80) and Diuktai Cave (9, 23), offer an op-
portunity to test this hypothesis.
The archaeological evidence for both early humans and dogs in
Siberia and the Americas is sparse. The ability to isolate and se-
quence ancient DNA from the few individuals that have been
recovered is gradually providing new insights into the populations
that initially moved east over the Bering Land Bridge and into the
Americas. Dog mitochondrial sequences reflect the history of just
a single locus, and genomic sequences are necessary to recon-
struct their population history. Nonetheless, the coalescence time
estimates of their mtDNA lineages suggest that dogs and humans
share a correlated history of population divergences and migra-
tion from Siberia into the Americas. More specifically, we suggest
that the first people to enter the Americas likely did so with their
dogs. The subsequent geographic dispersal and genetic diver-
gences within each population suggest that where people went,
dogs went.
The convergence of the early genetic histories of people and
dogs in Siberia and Beringia suggests that this may be the region
where humans and wolves first entered into a domestic relation-
ship. The oldest time to most recent common ancestor of the A
haplogroup suggests that this process had already begun by
2619.7 ka, which precedes the first unequivocal dogs in the
Eurasian archaeological record by 11,000 to 4,000 y. The vast
expanse of the region, combined with limited excavation, may
explain the absence of earlier dog remains in Siberia. Future anal-
yses of the handful of existing putative dogs, such as those from
the site of Afontova Gora (80), are necessary to test this hypothesis.
Since their emergence from wolves, dogs have played a wide
variety of roles within human societies, many of which are spe-
cifically tied to the lifeways of cultures worldwide. Future ar-
chaeological research, combined with numerous scientific
techniques, will no doubt reveal how the emerging mutual rela-
tionship between people and dogs led to their successful
dispersal across the globe.
Data Availability. All study data are included in the article and/or
SI Appendix.
A.R.P. was funded by the European Unions Seventh Framework Programme for
research, technological development and demonstration under grant agree-
ment 609412. T.R.F. was funded by the European Unions EU Framework Pro-
gramme for research and innovation Horizon 2020 under grant agreement
676154, and funding for the Qimmeq project came from the Velux Foundations
and the Aage og Johanne Louis-Hansens Fond. L.A.F.F. and G.L. were sup-
ported by a European Research Council grants (ERC-2013-StG-337574-UN-
DEAD and ERC-2019-StG-853272-PALAEOFARM) and Natural Environmental
Research Council grants (NE/K005243/1 and NE/K003259/1). D.J.M. is sup-
ported by the Quest Archaeological Research Fund. K.E.W. was supported by
a Wenner Gren Foundation grant and with R.S.M. an NSF (BCS-1540336) grant
and is currently supported by NIH grant awarded to Dr. Emilia Huerta-Sanchez
(1R35GM128946-01). The authors thank Victor Moreno-Mayar, Yun S. Song, and
the editor and the reviewers for helpful advice.
1A. H. Freedman et al., Genome sequencing highlights the dynamic early history of dogs. PLoS Genet. 10, e1004016 (2014).
2G. Larson et al., Rethinking dog domestication by integrating genetics, archeology, and biogeography. Proc. Natl. Acad. Sci. U.S.A. 109, 88788883 (2012).
3A. Perri, A wolf in dogs clothing: Initial dog domestication and Pleistocene wolf variation. J. Archaeol. Sci. 68,14 (2016).
4L. Janssens et al., A new look at an old dog: Bonn-Oberkassel reconsidered. J. Archaeol. Sci. 92, 126138 (2018).
5M. Ollivier et al., Dogs accompanied humans during the Neolithic expansion into Europe. Biol. Lett. 14, 20180286 (2018).
6K. Greig et al., Complete mitochondrial genomes of New Zealands first dogs. PLoS One 10, e0138536 (2015).
7C. Ameen et al., Specialized sledge dogs accompanied Inuit dispersal across the North American Arctic. Proc. Biol. Sci. 286, 20191929 (2019).
8S. K. Brown, C. M. Darwent, B. N. Sacks, Ancient DNA evidence for genetic continuity in arctic dogs. J. Archaeol. Sci. 40, 12791288 (2013).
9A. Perri et al., New evidence of the earliest domestic dogs in the Americas. Am. Antiq. 84,6887 (2019).
10 J. V. Moreno-Mayar et al., Early human dispersals within the Americas. Science 362, eaav2621 (2018).
11 L. G. Davis et al., Late upper paleolithic occupation at Coopers Ferry, Idaho, USA, 16,000 years ago. Science 365, 891897 (2019).
12 A. Perri, A wolf in dog s clothing: Initial dog domestication and Pleistocene wolf variation. J. Archaeol. Sci. 68,14 (2016).
13 L. Janssens et al., A new look at an old dog: Bonn-Oberkassel reconsidered. J. Archaeol. Sci. 92, 126138 (2018).
14 P. Skoglund, E. Ersmark, E. Palkopoulou, L. Dal ´en, Ancient wolf genome reveals an early divergence of domestic dog ancestors and admixture into high-latitude
breeds. Curr. Biol. 25, 15151519 (2015).
PNAS Perri et al. Dog domestication and the dual dispersal of people and dogs into the Americas
Downloaded at MPI f. evolutionäre Anthropologie on January 25, 2021
15 L. A. F. Frantz et al., Genomic and archaeological evidence suggest a dual origin of domestic dogs. Science 352, 12281231 (2016).
16 M. N´
ıLeathlobhair et al., The evolutionary history of dogs in the Americas. Science 361,8185 (2018).
17 J. A. Leonard et al., Ancient DNA evidence for old world origin of new world dogs. Science 298, 16131616 (2002).
18 K. E. Witt et al., DNA analysis of ancient dogs of the Americas: Identifying possible founding haplotypes and reconstructing population histories. J. Hum. Evol. 79,
105118 (2015).
19 O. Thalmann, A. R. Perri, Paleogenomic inferences of dog domesticationin Paleogenomics: Genome-Scale Analysis of Ancient DNA, C. Lindqvist, O. P. Rajora,
Eds. (Springer International Publishing, 2019), pp. 273306.
20 G. Larson, D. G. Bradley, How much is that in dog years? The advent of canine population genomics. PLoS Genet. 10, e1004093 (2014).
21 L. A. F. Frantz, D. G. Bradley, G. Larson, L. Orlando, Animal domestication in the era of ancient genomics. Nat. Rev. Genet. 21, 449460 (2020).
22 M. Germonpr ´eet al., Fossil dogs and wolves from palaeolithic sites in Belgium, the Ukraine and Russia: Osteometry, ancient DNA and stable isotopes. J. Archaeol.
Sci. 36, 473490 (2009).
23 M. Germonpr ´eet al., Palaeolithic and prehistoric dogs and Pleistocene wolves from Yakutia: Identification of isolated skulls. J. Archaeol. Sci. 78,119 (2017).
24 M. Germonpr ´e, M. azni ˇckov ´a-Galeto v ´a, M. V. Sablin, Palaeolithic dog skulls at the Gravettian P
ısite, the Czech Republic. J. Archaeol. Sci. 39,184202
25 M. V. Sablin, G. A. Khlopachev, The earliest ice age dogs: Evidence from Eliseevichi 1. Curr. Anthropol. 43, 795799 (2002).
26 E. Camar ´os, S. C. Münzel, M. Cueto, F. Rivals, N. J. Conard, The evolution of Paleolithic hominincarnivore interaction written in teeth: Stories from the Swabian
Jura (Germany). J. Archaeol. Sci. Rep. 6, 798809 (2016).
27 N. D. Ovodov et al., A 33,000-year-old incipient dog from the Altai Mountains of Siberia: Evidence of the earliest domestication disrupted by the last glacial
maximum. PLoS One 6, e22821 (2011).
28 A. S. Druzhkova et al., Ancient DNA analysis affirms the canid from Altai as a primitive dog. PLoS One 8, e57754 (2013).
29 H. Bocherens et al., Reconstruction of the Gravettian food-web at P
ıI using multi-isotopic tracking (13C, 15N, 34S) of bone collagen. Quat. Int. 359-360,
211228 (2015).
30 O. Thalmann et al., Complete mitochondrial genomes of ancient canids suggest a European origin of domestic dogs. Science 342, 871874 (2013).
31 K. A. Prassack, J. DuBois, M. L ´azni ˇckov´a-Galetov ´a, M. Germonpr ´e, P. S. Ungar, Dental microwear as a behavioral proxy for distinguishing between canids at the
Upper Paleolithic (Gravettian) site of P
ı, Czech Republic. J. Archaeol. Sci. 115, 105092 (2020).
32 L. Janssens, A. Perri, P. Cromb ´e, S. Van Dongen, D. Lawler, An evaluation of classical morphologic and morphometric parameters reported to distinguish wolves
and dogs. J. Archaeol. Sci. Rep. 23, 501533 (2019).
33 A. G. Drake et al., Three-dimensional geometric morphometric analysis of fossil canid Mandibles and skulls. Sci. Rep. 7, 9508 (2017).
34 A. G. Drake, M. Coquerelle, G. Colombeau, 3D morphometric analysis of fossil canid skulls contradicts the suggested domestication of dogs during the late
Paleolithic. Sci. Rep. 5, 8299 (2015).
35 M. Boudadi-Maligne, G. Escarguel, A biometric re-evaluation of recent claims for Early Upper Palaeolithic wolf domestication in Eurasia. J. Archaeol. Sci. 45,
8089 (2014).
36 C. Ameen et al., A landmark-based approach for assessing the reliability of mandibular tooth crowding as a marker of dog domestication. J. Archaeol. Sci. 85,
4150 (2017).
37 L. Loog et al., Ancient DNA suggests modern wolves trace their origin to a Late Pleistocene expansion from Beringia. Mol. Ecol. 29, 15961610 (2020).
38 L. R. Botigu ´eet al., Ancient European dog genomes reveal continuity since the Early Neolithic. Nat. Commun. 8, 16082 (2017).
39 L. A. F. Frantz et al., Genomic and archaeological evidence suggest a dual origin of domestic dogs. Science 352, 12281231 (2016).
40 J. Wilczy ´nski et al., Friend or foe? Large canid remains from pavlovian sites and their archaeozoological context. J. Anthropol. Archaeol. 59, 101197
41 A. Perri, S. Sazelova, The role of large canids: Preliminary variabilities forming the population structure in Moravia (Dolni Vestonice II)in Doln ´
ıVestonice II:
Chronostratigraphy, Paleoethnology, Paleoanthropology, J. Svoboda, Ed. (Academy of Sciences of the Czech Republic, Institute of Archaeology, 2016),
pp. 138146.
42 M. Germonpr ´eet al., Palaeolithic dogs and the early domestication of the wolf: A reply to the comments of. J. Archaeol. Sci. 40, 786792 (2013).
43 P. Galeta, M. L ´azni ˇckov´a-Galetov ´a, M. Sablin, M. Germonpr´e, Morphological evidence for early dog domestication in the European Pleistocene: New evidence
from a randomization approach to group differences. Anat. Rec. (Hoboken) 304,4262 (2020).
44 S. J. Crockford, Y. V. Kuzmin, Comments on Germonpr ´e et al., Journal of Archaeological Science 36, 2009 Fossil dogs and wolves from Palaeolithic sites in
Belgium, the Ukraine and Russia: osteometry, ancient DNA and stable isotopes, and Germonpr ´e, azki ˇ
ckov ´a-Galetov ´a, and Sablin, Journal of Archaeological
Science 39, 2012 Palaeolithic dog skulls at the Gravettian P
ısite, the Czech Republic..J. Archaeol. Sci. 39, 27972801 (2012).
45 D. F. Morey, In search of paleolithic dogs: A quest with mixed results. J. Archaeol. Sci. 52,300307 (2014).
46 А.В.Кандыба,С.Е.Федоров,А.И.Дмитриев,Местонахождение Сыалах-новый археологический объект позднег
онеоплейстоцена Сибирской Арктики.Проблемы археологии 21,9093 (2015).
47 J. Ramos-Madrigal et al., Genomes of Pleistocene Siberian wolves uncover multiple extinct wolf lineages. Curr. Biol. In press.
48 M. Germonpr ´e, M. L ´azni ˇckov ´a-Galetov ´a, R. J. Losey, J. Räikkönen, M. V. Sablin, Large canids at the Gravettian P
ısite, the Czech Republic: The mandible.
Quat. Int. 359-360, 261279 (2015).
49 M. Pionnier-Capitan et al., New evidence for upper palaeolithic small domestic dogs in south-western Europe. J. Archaeol. Sci. 38, 21232140 (2011).
50 R. Musil, Domestication of wolves in central European Magdalenian sitesin Dogs Through T ime: An Archaeological Perspective, S. J. Crockford, Ed. (BAR
International Series, British Archaeological Reports, Oxford, 2000), vol. 889, pp. 2128.
51 E. Tchernov, F. F. Valla, Two new dogs, and other natufian dogs, from the southern levant. J. Archaeol. Sci. 24,6
595 (1997).
52 F. Boschin et al., The first evidence for Late Pleistocene dogs in Italy. Sci. Rep. 10, 13313 (2020).
53 P. Morel et al., Un Campement Magdal ´enien au Bord du Lac de Neuch ˆatel: ´
Etude Arch ´eozoologique (Secteur 1) (Mus ´ee cantonal darch ´eologie, 1997).
54 A. Bergström et al., Origins and genetic legacy of prehistoric dogs. Science 370, 557564 (2020).
55 B. van Asch et al., Pre-Columbian origins of native American dog breeds, with only limited replacement by European dogs, confirmed by mtDNA analysis. Proc.
Biol. Sci. 280, 20131142 (2013).
56 M. Sikora et al., The population history of northeastern Siberia since the Pleistocene. Nature 570,182188 (2019).
57 B. Llamas et al., Ancient mitochondrial DNA provides high-resolution time scale of the peopling of the Americas. Sci. Adv. 2, e1501385 (2016).
58 E. Tamm et al., Beringian standstill and spread of Native American founders. PLoS One 2, e829 (2007).
59 T. Pinotti et al., Y chromosome sequences reveal a short Beringian standstill, rapid expansion, and early population structure of native American founders. Curr.
Biol. 29, 149157.e3 (2019).
60 J. V. Moreno-Mayar et al., Terminal Pleistocene Alaskan genome reveals first founding population of Native Americans. Nature 553, 203207 (2018).
61 D. J. Meltzer, First Peoples in a New World: Populating Ice Age America (Cambridge University Press, 2021).
62 M. Rasmussen et al., The genome of a Late Pleistocene human from a Clovis burial site in western Montana. Nature 506, 225229 (2014).
63 T. D. Dillehay et al., Monte verde: Seaweed, food, medicine, and the peopling of south America. Science 320, 784786 (2008).
64 D. Froese, J. M. Young, S. L. Norris, M. Margold, Availability and viability of the ice-free corridor and pacific coast routes for the peopling of the Americas. SAA
Archaeol. Rec. 19,2733 (2019).
65 C. Posth et al., Reconstructing the deep population history of Central and South America. Cell 175, 11851197.e22 (2018).
Perri et al. PNAS
Dog domestication and the dual dispersal of people and dogs into the Americas
Downloaded at MPI f. evolutionäre Anthropologie on January 25, 2021
66 S. J. Fiedel, Mans best friendmammoths worst enemy? A speculative essay on the role of dogs in paleoindian colonization and megafaunal extinction. World
Archaeol. 37,1125 (2005).
67 M. S. Sinding et al., Arctic-adapted dogs emerged at the Pleistocene-Holocene transition. Science 368, 14951499 (2020).
68 P. Flegontov et al., Palaeo-Eskimo genetic ancestry and the peopling of Chukotka and North America. Nature 570,236240 (2019).
69 B. A. Potter, J. D. Irish, J. D. Reuther, H. J. McKinney, New insights into eastern beringian mortuary behavior: A terminal Pleistocene double infant burial at upward
sun river. Proc. Natl. Acad. Sci. U.S.A. 111, 1706017065 (2014).
70 B. A. Potter, J. D. Irish, J. D. Reuther, C. Gelvin-Reymiller, V. T. Holliday, A terminal Pleistocene child cremation and residential structure from eastern Beringia.
Science 331, 10581062 (2011).
71 T. Goebel, B. Potter, First tracesin The Oxford Handbook of the Prehistoric Arctic, T. M. Friesen, O. K. Mason, Eds. (Oxford University Press, 2016), p. 223.
72 M. R. Waters et al., Pre-Clovis projectile points at the debra L. Friedkin site, Texas-implications for the late Pleistocene peopling of the Americas. Sci. Adv. 4,
eaat4505 (2018).
73 D. L. Jenkins et al., Geochronology, archaeological context, and DNA at the paisley caves. Paleoamerican Odyssey 32, 485510 (2013).
74 Y. V. Kuzmin, S. G. Keates, Siberia and neighboring regions in the last glacial maximum: Did people occupy northern Eurasia at that time? Archaeol. Anthropol.
Sci. 10, 111124 (2018).
75 K. E. Graf, Siberian odysseyin Paleoamerican Odyssey, K. E. Graf, C. V. Ketron, M. R. Waters, Eds. (Texas A&M University Press, 2014), pp. 6580.
76 G.-D. Wang et al., Out of southern east Asia: The natural history of domestic dogs across the world. Cell Res. 26,2133 (2016).
77 L. M. Shannon et al., Genetic structure in village dogs reveals a Central Asian domestication origin. Proc. Natl. Acad. Sci. U.S.A. 112, 1363913644 (2015).
78 M. Lipson, D. Reich, A working model of the deep relationships of diverse modern human genetic lineages outside of Africa. Mol. Biol. Evol. 34, 889902 (2017).
79 M. Raghavan et al., Upper Palaeolithic Siberian genome reveals dual ancestry of Native Americans. Nature 505,8791 (2014).
80 Y. V. Kuzmin, Mammalian fauna from palaeolithic sites in the upper Yenisei river basin (southern Siberia): Review of the current zooarchaeological evidence. Int.
J. Osteoarchaeol. 21, 218228 (2011).
81 M. Germonpr ´e, M. V. Sablin, Humans and mammals in the Upper Palaeolithic of Russiain The Oxford Handbook of Zooarchaeology, H. Russ, U. Albarella,
K. Vickers, M. Rizzetto, S. Viner-Daniels, Eds. (Oxford University Press, 2017), pp. 2538.
82 S. J. Olsen, Origins of the Domestic Dog: The Fossil Record (University of Arizona Press, 1985).
83 M. A. Zeder, Pathways to animal domesticationin Biodiversity in Agriculture: Domestication, Evolution, and Sustainability, J. R. Harlan et al., Eds. (Cambridge
University Press, 2012), pp. 227259.
PNAS Perri et al. Dog domestication and the dual dispersal of people and dogs into the Americas
Downloaded at MPI f. evolutionäre Anthropologie on January 25, 2021
... The prevailing view on the beginnings of animal domestication is that the first taxon domesticated by humans was the wolf (Canis lupus), giving rise to domestic dogs (C. familiaris) [1][2][3][4][5][6][7][8]. Scholars and scientists have widely contended that the domestication of a now-extinct variant of this wild carnivore was initiated somewhere in Late Pleistocene Eurasia, including eastern Asia (e.g., China), by a non-sedentary population of anatomically modern human (AMH) hunter-gatherers, certainly by at least 15-16 thousand years (ka) ago [9], but possibly up to 40 ka ago according to some authorities [10]. ...
... In the case of the wolf, two competing hypotheses have been proposed to explain how a non-sedentary population of Late Pleistocene foragers could have domesticated this large predatory canid [45][46][47][48]. The first hypothesis is that the wolf 'selfdomesticated' by establishing a commensal relationship with mobile groups of AMH hunter-gatherers [4,7,47,[49][50][51][52][53][54][55][56][57][58]. The second hypothesis, now increasingly seen by scholars as being more tenable than the first (but cf. ...
Full-text available
The Sulawesi warty pig (S. celebensis) is a wild and still-extant suid that is endemic to the Indonesian island of Sulawesi. It has long been theorised that S. celebensis was domesticated and/or deliberately introduced to other islands in Indonesia prior to the advent of the Neolithic farming transition in the region. Thus far, however, there has been no empirical support for this idea, nor have scientists critiqued the argument that S. celebensis was a pre-Neolithic domesticate in detail. Here, it is proposed that early foragers could have formed a relationship with S. celebensis that was similar in essence to the close association between Late Pleistocene foragers in Eurasia and the wild wolf ancestors of domestic dogs. That is, a longstanding practice of hunter-gatherers intensively socialising wild-caught S. celebensis piglets for adoption into human society as companion animals (‘pets’) may have altered the predator–prey dynamic, brought aspects of wild pig behaviour and reproduction under indirect human selection and control, and caused changes that differentiated human-associated pigs from their solely wild-living counterparts.
... We chose to work with dogs as an initial study species for this inter-species helping investigation for two main reasons. First, dogs have a unique coevolutionary history with humans as they are by far the earliest known domesticated species (appearing genetically distinct from wolves as early as 23,000 years ago; Nagasawa et al., 2015;Perri et al., 2021). Second, dogs have evolved to read human social cues (Hare and Tomasello, 1999) and naturally seek out human help when they encounter problems: Dogs' help-seeking behaviors such as reaching and 'begging' for items they cannot access (Marshall-Pescini et al., 2017) resemble the actions of human experimenters in a classic experimental helping paradigm (Warneken and Tomasello, 2006). ...
Full-text available
Two capacities considered foundational in human cooperation are prosocial motivation and goal-reading abilities that enable helping. Children exhibit both proclivities by age 2 in interactions with other humans, but interactions with nonhuman species on whom we have been interdependent for millennia are unstudied. We tested the hypothesis that children’s goal-reading and prosocial propensities extend to other animals. We predicted children would help pet dogs access objects that dogs attempted to reach but could not reach themselves. We studied 97 children between 2 and 3 years of age living in a small mid-western US city, 44 of whom had dogs as household pets. In a quasi-naturalistic setting, we introduced children to 1 of 3 friendly pet dogs who remained within a small, porous enclosure while a treat or toy was placed outside it. Dogs reacted naturally, either showing interest in accessing the item (e.g., pawing, begging) or ignoring it. Measures of dog and child behavior during sessions were coded blindly with high reliability. Children provided dogs with out-of-reach items twice as often when dogs showed interest rather than ignored items, indicating sensitivity to the dog’s goals. Additionally, children were more generally likely to provide dogs with items if children lived with pet dogs, if dogs were more lively and engaged rather than subdued and if the item was a treat rather than a toy. These findings lend support to our hypothesis that children’s early-developing proclivities for goal-reading and prosociality extend beyond humans to other animals.
... Des traces d'ADN peuvent être préservées dans des tissus anciens, le plus souvent dans les tissus calcifiés (os, dents), les tissus kératinisés (sabots, bois, ongles, cheveux) ou encore dans l'environnement (sols...) (Damgaard et al., 2015;Ellegaard et al., 2020;Gelabert et al., 2021). Il est possible de replacer des espèces éteintes au sein des arbres phylogénétiques des espèces à partir de fragments d'ossements, reconstruire leur évolution, ainsi qu'identifier les migrations, extensions, rétrécissements et remplacements des populations humaines, animales et végétales (Brunel et al., 2020;Grange et al., 2018;Kistler, 2012;Massilani et al., 2016;Mikić, 2015;Perri et al., 2021;van der Valk et al., 2021;Verdugo et al., 2019). Il est ainsi possible d'établir les éventuelles relations entre extinction des espèces et changements climatiques et environnementaux du passé. ...
Les genres Bos et Bison constituent la sous-tribu des Bovina et incluent plusieurs lignées de bovins. Les espèces domestiques et sauvages appartenant à ces deux genres auraient divergé au Pléistocène inférieur. L’histoire génomique évolutive récente de Bos et Bison ne reflète qu’imparfaitement les liens phylogénétiques anciens car ceux-ci sont éclipsés par les événements récents, en particulier la quasi-extermination aux 19ème et 20ème siècles des bisons américains (Bison bison) et bisons européens (Bison bonasus), respectivement, ainsi que la disparition au 17ème siècle de l’aurochs, ancêtre des bovins domestiques actuels. Dans le but de clarifier l’histoire évolutive de ces deux genres, nous avons mis en place une approche paléogénomique permettant de séquencer l’ADN ancien extrait à partir de spécimens fossiles des genres Bos et Bison. L’objectif était de reconstruire les mitogénomes et les génomes nucléaires de restes fossiles datant du Pléistocène à l’Holocène, témoins des événements évolutifs précédant les événements les plus récents qui ont remodelé la diversité génétique des populations modernes. Nous avons ainsi analysé des restes fossiles du Néolithique jusqu’à l’ère moderne, témoins de l’impact de la domestication sur la structuration génétique des bovins. Nous avons adapté les méthodes d’analyse de l’ADN aux caractéristiques particulières de l’ADN ancien pour maximiser la récupération de l’ADN ancien à partir des restes fossiles et convertir efficacement les fragments d’ADN contenus dans l’extrait fossile en banques génomiques pour le séquençage de nouvelle génération (NGS). Nous avons aussi optimisé un protocole de capture du génome mitochondrial de bovins anciens pour augmenter son efficacité avec des échantillons mal conservés. L’optimisation méthodologique nous a permis d’obtenir la séquence de 20 mitogénomes complets de bisons européens et bisons des steppes datant du Pléistocène moyen jusqu’aux aux temps modernes et 98 mitogénomes complets d’aurochs et de bovins domestiques eurasiatiques et africains datant du Pléistocène supérieur jusqu’au Moyen Âge. Nous avons reconstruit l’histoire évolutive de Bos et Bison à travers une analyse phylogénétique qui rassemble 279 mitogénomes anciens et 671 mitogénomes modernes des Bovina. Ceci nous a permis d’établir une datation robuste des radiations phylogénétiques et de reconstruire les dynamiques des populations bovines pendant les 50 000 dernières années.Nous avons identifié un nouvel haplogroupe mitochondrial d’aurochs rassemblant des échantillons du Pléistocène supérieur originaire de l’Europe de l’Ouest. Cet haplogroupe s’est séparé le premier des lignées d’aurochs européens et proche-orientaux, des bovins domestiques eurasiatiques et africains ainsi que des zébus. Ceci met en évidence une dynamique complexe des populations bovines ancestrales au Pléistocène moyen et supérieur ainsi que les liens partagés entre les populations eurasiatiques et du sous-continent indien. L’analyse d’échantillons précédant et suivant le dernier maximum glaciaire, entre 30 000 et 12 000 ans, et les modélisations démographiques utilisant les mitogénomes anciens et actuels des différentes lignées ont permis d’évaluer l’impact de cette glaciation sur la dynamique des tailles effectives des populations d’aurochs. Notre étude permet de réévaluer les causes du goulot d’étranglement précédemment attribué à la domestication des aurochs. Nous avons aussi caractérisé en parallèle les génomes d’une vingtaine de bisons qui permettront de comparer l’évolution de ces deux lignées cousines, l’une ayant été domestiquée et l’autre non. L’ensemble du travail réalisé permet d’établir une base solide pour étudier les changements génomiques associés à la domestication des bovins.
... Après des millénaires de construction de niches basées sur la chasse, la cueillette, la pêche et la recherche de nourriture, les communautés de diverses régions du monde se sont lancées dans des trajectoires de production alimentaire qui, dans certains cas, ont permis l'émergence de sociétés complexes, d'agglomérations urbaines et d'empires semant ainsi les graines de la mondialisation actuelle . Grâce aux récentes et intensives recherches en archéozoologie et en paléogénomique, il est maintenant admis que, mis à part la domestication du chien au sein des sociétés de chasseurs-cueilleurs aux alentours de 23 000 BP (Perri et al., 2021), la domestication des animaux d'élevage d'importance mondiale s'est produite au sein de communautés sédentaires engagées dans l'agriculture dans trois berceaux indépendants . Le plus ancien centre de domestication animale est localisé en Asie du Sud-Ouest où les moutons (Ovis aries), chèvres (Capra hircus), porcs (Sus scrofa) et bovins sont les formes domestiquées apparues entre 10 500 et 10 000 BP (Colledge et al., 2013). ...
Les lamas (Lama glama Linnaeus, 1758) et les alpagas (Vicugna pacos Linnaeus, 1758) ont été domestiqués à partir de deux espèces sauvages : les guanacos (Lama guanicoe Müller, 1776) et les vigognes (Vicugna vicugna Molina, 1782). Les camélidés domestiques ont joué un rôle économique et symbolique majeur dans le développement des sociétés préhispaniques. Cependant, la documentation du processus de domestication des camélidés sud-américains (CSA) reste un défi. Pour obtenir de nouvelles informations sur ce processus, nous nous sommes concentrés sur la zone des Andes centrales et le site de Telarmachay (Puna de Junín, Pérou). Sur la longue séquence chronologique de ce site, de 7000 à 150 av. J.-C., des preuves de contrôle/management ont été suggérées vers 3000-1800 av. J.-C., proposant un processus de domestication des CSA antérieur de 2000 ans aux autres centres de domestication de la zone andine. Afin d'explorer l’intensification des interactions entre sociétés humaines et camélidés, nous avons produit de nouvelles datations 14C des ossements de CSA et utilisé une approche multicritère associant des profils de mortalité, la morphométrie morphométrique 3D du talus et une analyse des teneurs en isotopes stables du carbone et de l’azote (δ13C et δ15N) du collagène de l’os. Nos résultats nous permettent d’étayer l’hypothèse d’un processus ancien de domestication des CSA in situ à Telarmachay dès le 5ème millénaire, entre 5106/4851 et 4328/4129 av. J.-C. La morphométrie géométrique 3D du talus nous permettant de suivre l’évolution phénotypique des deux lignées de camélidés le long de la séquence chronologique du site, nous avons pu observer, dès cette période, la présence de formes hybrides de taille intermédiaire entre les vigognes et les guanacos. Ces premiers changements de la diversité phénotypique sont associés à des profils de mortalité témoignant d’une phase de changement d’exploitation des camélidés avec la forte représentation des camélidés ayant entre 0 et 3 mois. C’est à la phase suivante, entre 4270/4060 à 3626/3054 av. J.-C, que l’on observe un changement marqué de la variabilité du talus, interprété comme la conséquence d’une contrainte plus forte sur la mobilité des vigognes avec potentiellement une mise en captivité, comme le suggère la présence d’enclos empierrés à proximité du site. Enfin, si la réduction de la mobilité des guanacos semble moins claire, des modifications squelettiques ont quand même été mises en évidence dès le 5ème millénaire. D’après les valeurs du δ13C et δ15N, les camélidés de Telarmachay ont vécu dans la puna autour du site et n’ont pas subi de changement majeur dans leur alimentation et leur environnement durant 4000 ans. En revanche, une légère variation a été mise en évidence dès le 3ème millénaire, entre 3179/2495 et 2226/1854 av. J.-C., qui pourrait concorder avec le contrôle de la mobilité des camélidés. Ainsi, l’accroissement du contrôle culturel des CSA à partir du 5ème millénaire av. J.-C. soutient l’hypothèse de l’ancienneté du processus de domestication des camélidés andins dans la région de Junín. Nous proposons que l’écosystème de la Puna du Junin fût un écosystème particulièrement propice à l’intensification des interactions entre les sociétés de chasseurs-cueilleurs et celles des camélidés à l’Holocène.
... 91 In future studies, ancient genomes will provide insight into temporal and geographic origins of variants identified in this study. [92][93][94][95] Importantly, this study focuses on behavioral diversification that postdates domestication, the nature, timing, and precise geographical origins of which remain controversial. [96][97][98] Nonetheless, ongoing efforts to resolve outstanding questions in dog domestication will be critical for understanding the nature of canine behavioral diversification relative to domestication-related behavioral shifts. ...
Article Domestic dog lineages reveal genetic drivers of behavioral diversification Graphical abstract Highlights d High-dimensional data analysis reveals canine lineages, resolving breed relationships d Canine behavioral diversification predates modern breed formation d Ancient non-coding variation drives working role-related dog behaviors d Canine genetic diversity is associated with neurodevelopmental gene co-expression A framework for understanding the relationships between canine breeds allows for the identification of genetic drivers of the behaviors that define them. SUMMARY Selective breeding of domestic dogs has generated diverse breeds often optimized for performing specialized tasks. Despite the heritability of breed-typical behavioral traits, identification of causal loci has proven challenging due to the complexity of canine population structure. We overcome longstanding difficulties in identifying genetic drivers of canine behavior by developing a framework for understanding relationships between breeds and the behaviors that define them, utilizing genetic data for over 4,000 domestic, semi-feral, and wild canids and behavioral survey data for over 46,000 dogs. We identify ten major canine genetic line-ages and their behavioral correlates and show that breed diversification is predominantly driven by non-coding regulatory variation. We determine that lineage-associated genes converge in neurodevelopmental co-expression networks, identifying a sheepdog-associated enrichment for interrelated axon guidance functions. This work presents a scaffold for canine diversification that positions the domestic dog as an unparalleled system for revealing the genetic origins of behavioral diversity.
Full-text available
Evolution has at regular intervals developed new ways to transmit information to the next generation: GENETIC TRANSMISSION (DNA): 1: asexual, 2: sexual; SOCIAL TRANSMISSION (teaching): 3: parental feedback, 4: tool transfer, 5: referential gesture, 6: demonstration, 7: communication of concepts, 8: explanation of relationships between concepts, 9: narration; TECHNOLOGICAL TRANSMISSION (via writing): 10: hand writing, 11: mechanical printing, 12: computer-based. The pattern of these transmission methods is the same as in nonlinear, dynamic, period-doubling systems as described by Chaos Theory, namely at intervals decreasing by the factor 4.66920…, known as the Feigenbaum constant delta.
Full-text available
Valerii Manko and Guram Chkhatarashvili published their article in the “Arheologia”, No. 2, 2022. In the paper, they discussed the migration of bearers of four Neolithic flint industries from Southwest Asia through the Caucasus to the south of Eastern Europe from the final Pleistocene to the early Atlantic. According to the authors, stable connections between these remote areas led to the emergence of four “information networks”, which they called “Cultural-Historical Regions” (CHR). The authors believe that the first region of such type in human history was the Pre-Pottery Neolithic A (PPNA) in the Near East. Therefore, they call the “theoretical basis” of their study “the idea of understanding the Neolithic as an epoch of the formation of global information networks, within which innovations created in the Near and Middle East were disseminated.” V. Manko began to develop the described theoretical views in 2010 when he wrote that the reason for the emergence of the CHR is the ability to communicate, formed due to the mental changes of inhabitants of the PPNA large settlements. The statement about specific psyche and worldview as the basis of Neolithic has been expressed as an idea of Post-Processual archaeology long before V. Manko announced it. In particular, Trevor Watkins developed this concept in detail. However, V. Manko does not mention works by any post-processualists in his publications. The statement about the formation of the ability to communicate only in the Neolithic is V. Manko’s novelty. He based it on one reference to a publication of Alexey N. Sorokin, who allegedly claimed that the bearers of different flint industries did not contact each other in the central part of European Russia in the Mesolithic. V. Manko misinterpreted this particular subjective observation and gave it the meaning of a global pattern. Thus, his definition of the Neolithic is controversial, because of using this erroneous premise. Generally, V. Manko’s theoretical reasoning is full of contradictions, logical errors, terminological chaos, and rhetoric in the postmodernism style. It is noteworthy that V. Manko himself does not fully adhere to his previous theoretical views in his later works.
Human history is inextricably linked to the introduction of desirable heritable traits in plants and animals. Selective breeding (SB) predates our historical period and has been practiced since the advent of agriculture and farming more than ten thousand years ago. Since the 1970s, methods of direct plant and animal genome manipulation are constantly being developed. These are collectively described as "genetic engineering" (GE). Plant GE aims to improve nutritional value, insect resistance and weed control. Animal GE has focused on livestock improvement and disease control. GE applications also involve medical improvements intended to treat human disease. The scientific consensus built around marketed products of GE organisms (GEOs) is usually well established, noting significant benefits and low risks. GEOs are exhaustively scrutinized in the EU and many non-EU countries for their effects on human health and the environment, but scrutiny should be equally applied to all previously untested organisms derived directly from nature or through selective breeding. In fact, there is no evidence to suggest that natural or selectively bred plants and animals are in principle safer to humans than GEOs. Natural and selectively bred strains evolve over time via genetic mutations that can be as risky to humans and the environment as the mutations found in GEOs. Thus, previously untested plant and animal strains aimed for marketing should be proven useful or harmful to humans only upon comparative testing, regardless of their origin. Highlighting the scientific consensus declaring significant benefits and rather manageable risks provided by equitably accessed GEOs, can mitigate negative predispositions by policy makers and the public. Accordingly, we provide an overview of the underlying technologies and the scientific consensus to help resolve popular myths about the safety and usefulness of GEOs.
Full-text available
The article presents a literary review of scientific works related to the topic of the role of dogs in human life. The union of man and dog, as scientists report, dates back to ancient times, probably 14000 to 16000 years ago. In modern conditions, dogs occupy one of the main places next to people in almost all spheres of life - from protection of the rights and freedoms of citizens to sports with the use of dogs, as well as non-standard methods of treatment with the active participation of dogs. Dogs were the first animals domesticated by man. In this regard, they managed to overtake cows, sheep and goats. Of course, dogs were great guards and helpers during the hunt, but much more important was that you could communicate with them. They were devoted to their master and understood him better than other animals. A dog's ability to track prey and warn of danger was enough for primitive man. It freed a person from the fear of the unknown. It is difficult to overestimate the role of dogs in human life, because they perform many functions, in particular, they are actively used for hunting, as shepherds, in the work of the police and the military, in everyday life for people with disabilities, and dog therapy is also widely distributed, that is, work with injured people due to certain circumstances.
What causes Ice Ages? How did we learn about them? What were their affects on the social history of humanity? Allan Mazur's book tells the appealing history of the scientific 'discovery' of Ice Ages. How we learned that much of the Earth was repeatedly covered by huge ice sheets, why that occurred, and how the waning of the last Ice Age paved the way for agrarian civilization and, ultimately, our present social structures. The book discusses implications for the current 'controversies' over anthropogenic climate change, public understanding of science, and (lack of) 'trust in experts'. In parallel to the history and science of Ice Ages, sociologist Mazur highlights why this is especially relevant right now for humanity. Ice Ages: Their Social and Natural History is an engrossing combination of natural science and social history: glaciology and sociology writ large.
Full-text available
Extant Canis lupus genetic diversity can be grouped into three phylogenetically distinct clades: Eurasian and American wolves and domestic dogs.1 Genetic studies have suggested these groups trace their origins to a wolf population that expanded during the last glacial maximum (LGM)1 , 2 , 3 and replaced local wolf populations.4 Moreover, ancient genomes from the Yana basin and the Taimyr peninsula provided evidence of at least one extinct wolf lineage that dwelled in Siberia during the Pleistocene.3 5 Previous studies have suggested that Pleistocene Siberian canids can be classified into two groups based on cranial morphology. Wolves in the first group are most similar to present-day populations, although those in the second group possess intermediate features between dogs and wolves.6 7 However, whether this morphological classification represents distinct genetic groups remains unknown. To investigate this question and the relationships between Pleistocene canids, present-day wolves, and dogs, we resequenced the genomes of four Pleistocene canids from Northeast Siberia dated between >50 and 14 ka old, including samples from the two morphological categories. We found these specimens cluster with the two previously sequenced Pleistocene wolves, which are genetically more similar to Eurasian wolves. Our results show that, though the four specimens represent extinct wolf lineages, they do not form a monophyletic group. Instead, each Pleistocene Siberian canid branched off the lineage that gave rise to present-day wolves and dogs. Finally, our results suggest the two previously described morphological groups could represent independent lineages similarly related to present-day wolves and dogs.
Full-text available
Dog domestication was multifaceted Dogs were the first domesticated animal, likely originating from human-associated wolves, but their origin remains unclear. Bergstrom et al. sequenced 27 ancient dog genomes from multiple locations near to and corresponding in time to comparable human ancient DNA sites (see the Perspective by Pavlidis and Somel). By analyzing these genomes, along with other ancient and modern dog genomes, the authors found that dogs likely arose once from a now-extinct wolf population. They also found that at least five different dog populations ∼10,000 years before the present show replacement in Europe at later dates. Furthermore, some dog population genetics are similar to those of humans, whereas others differ, inferring a complex ancestral history for humanity's best friend. Science , this issue p. 557 ; see also p. 522
Full-text available
The identification of the earliest dogs is challenging because of the absence and/or mosaic pattern of morphological diagnostic features in the initial phases of the domestication process. Furthermore, the natural occurrence of some of these characters in Late Pleistocene wolf populations and the time it took from the onset of traits related to domestication to their prevalence remain indefinite. For these reasons, the spatiotemporal context of the early domestication of dogs is hotly debated. Our combined molecular and morphological analyses of fossil canid remains from the sites of Grotta Paglicci and Grotta Romanelli, in southern Italy, attest of the presence of dogs at least 14,000 calibrated years before present. This unambiguously documents one of the earliest occurrence of domesticates in the Upper Palaeolithic of Europe and in the Mediterranean. The genetic affinity between the Palaeolithic dogs from southern Italy and contemporaneous ones found in Germany also suggest that these animals were an important common adjunct during the Late Glacial, when strong cultural diversification occurred between the Mediterranean world and European areas north of the Alps. Additionally, aDNA analyses indicate that this Upper Palaeolithic dog lineage from Italy may have contributed to the genetic diversity of living dogs.
Full-text available
In this paper we discuss recent claims that dogs were first domesticated from wild wolves in the Middle Upper Paleolithic (MUP), about 27 ka BP. According to our data, we think the presence of large canids at the Pavlovian/ MUP sites is a result of hunting specialization and not a sign of an early process of dog domestication. Our interpretation is supported by the following observations, whose implications we discuss: (1) Pavlovian faunal assemblages from seven sites in Moravia contain relatively high numbers of large canids; (2) gnaw-marking by large canids occurs with low frequency on the animal bones in these assemblages; (3) the bones of Pavlovian large canids in the sites often have cut marks from skinning, dismembering, and filleting. Whatever the reasons MUP people had for killing wolves, such as for food or for the skins, the effect would have reduced competition for prey between humans and wolves. The relatively high frequency of wolves at Pavlovian sites may have been a side effect of settlement aggregation and long-term occupations of sites, which could have attracted wolves to the settlements, and thus increased the need for humans to reduce their numbers.
Full-text available
The domestication of animals led to a major shift in human subsistence patterns, from a hunter–gatherer to a sedentary agricultural lifestyle, which ultimately resulted in the development of complex societies. Over the past 15,000 years, the phenotype and genotype of multiple animal species, such as dogs, pigs, sheep, goats, cattle and horses, have been substantially altered during their adaptation to the human niche. Recent methodological innovations, such as improved ancient DNA extraction methods and next-generation sequencing, have enabled the sequencing of whole ancient genomes. These genomes have helped reconstruct the process by which animals entered into domestic relationships with humans and were subjected to novel selection pressures. Here, we discuss and update key concepts in animal domestication in light of recent contributions from ancient genomics.
Full-text available
Grey wolves (Canis lupus) are one of the few large terrestrial carnivores that have maintained a wide geographic distribution across the Northern Hemisphere throughout the Pleistocene and Holocene. Recent genetic studies have suggested that, despite this continuous presence, major demographic changes occurred in wolf populations between the late Pleistocene and early Holocene, and that extant wolves trace their ancestry to a single late Pleistocene population. Both the geographic origin of this ancestral population and how it became widespread remain unknown. Here, we used a spatially and temporally explicit modelling framework to analyse a dataset of 90 modern and 45 ancient mitochondrial wolf genomes from across the Northern Hemisphere, spanning the last 50,000 years. Our results suggest that contemporary wolf populations trace their ancestry to an expansion from Beringia at the end of the Last Glacial Maximum, and that this process was most likely driven by Late Pleistocene ecological fluctuations that occurred across the Northern Hemisphere. This study provides direct ancient genetic evidence that long‐range migration has played an important role in the population history of a large carnivore, and provides an insight into how wolves survived the wave of megafaunal extinctions at the end of the last glaciation. Moreover, because late Pleistocene grey wolves were the likely source from which all modern dogs trace their origins, the demographic history described in this study has fundamental implications for understanding the geographical origin of the dog.
Over 15,000 years ago, a band of hunter-gatherers became the first people to set foot in the Americas. They soon found themselves in a world rich in plants and animals, but also a world still shivering itself out of the coldest depths of the Ice Age. The movement of those first Americans was one of the greatest journeys undertaken by ancient peoples. In this book, David Meltzer explores the world of Ice Age Americans, highlighting genetic, archaeological, and geological evidence that has revolutionized our understanding of their origins, antiquity, and adaptation to climate and environmental change. This fully updated edition integrates the most recent scientific discoveries, including the ancient genome revolution and human evolutionary and population history. Written for a broad audience, the book can serve as the primary text in courses on North American Archaeology, Ice Age Environments, and Human evolution and prehistory.
The antiquity of the wolf/dog domestication has been recently pushed back in time from the Late Upper Paleolithic (14,000 years ago) to the Early Upper Paleolithic (EUP; 36,000 years ago). Some authors questioned this early dog domestication claiming that the putative (EUP) Paleolithic dogs fall within the morphological range of recent wolves. In this study, we reanalyzed a data set of large canid skulls using unbalanced-and balanced-randomized discriminant analyses to assess whether the putative Paleolithic dogs are morphologically unique or whether they represent a subsample of the wolf morpho-population. We evaluated morphological differences between 96 specimens of the 4 a priori reference groups (8 putative Paleolithic dogs, 41 recent northern dogs, 7 Pleisto-cene wolves, and 40 recent northern wolves) using discriminant analysis based on 5 ln-transformed raw and allometrically size-adjusted cranial measurements. Putative Paleolithic dogs are classified with high accuracies (87.5 and 100.0%, cross-validated) and randomization experiment suggests that these classification rates cannot be exclusively explained by the small and uneven sample sizes of reference groups. It indicates that putative Upper Paleolithic dogs may represent a discrete canid group with morphological signs of domestication (a relatively shorter skull and wider palate and braincase) that distinguish them from sym-patric Pleistocene wolves. The present results add evidence to the view that these specimens could represent incipient Paleolithic dogs that were involved in daily activities of European Upper Paleolithic forager groups. K E Y W O R D S
Sled dog arctic adaptations go far back Dogs have been used for sledding in the Arctic as far back as ∼9500 years ago. However, the relationships among the earliest sled dogs, other dog populations, and wolves are unknown. Sinding et al. sequenced an ancient sled dog, 10 modern sled dogs, and an ancient wolf and analyzed their genetic relationships with other modern dogs. This analysis indicates that sled dogs represent an ancient lineage going back at least 9500 years and that wolves bred with the ancestors of sled dogs and precontact American dogs. However, gene flow between sled dogs and wolves likely stopped before ∼9500 years ago. Science , this issue p. 1495
Morphological and genetic evidence put dog domestication during the Paleolithic, sometime between 40,000 and 15,000 years ago, with identification of the earliest dogs debated. We predict that these earliest dogs (referred to herein as protodogs), while potentially difficult to distinguish morphologically from wolves, experienced behavioral shifts, including changes in diet. Specifically, protodogs may have consumed more bone and other less desirable scraps within human settlement areas. Here we apply Dental Microwear Texture Analysis (DMTA) to canids from the Gravettian site of P�redmostí (approx. 28,500 BP), which were previously assigned to the Paleolithic dog or Pleistocene wolf morphotypes. We test whether these groups separate out significantly by diet- related variation in microwear patterning. Results are consistent with differences in dietary breadth, with the Paleolithic dog morphotype showing evidence of greater durophagy than those assigned to the wolf morphotype. This supports the presence of two morphologically and behaviorally distinct canid types at this middle Upper Paleolithic site. Our primary goal here was to test whether these two morphotypes expressed notable differences in dietary behavior. However, in the context of a major Gravettian settlement, this may also support evidence of early stage dog domestication. Dental microwear is a behavioral signal that may appear generations before morphological changes are established in a population. It shows promise for distinguishing protodogs from wolves in the Pleistocene and domesticated dogs from wolves elsewhere in the archaeological record.