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Abstract

SignificanceChickens are the world's most numerous domestic animal. In order to understand when, where, and how they first became associated with human societies, we critically assessed the domestic status of chicken remains described in >600 sites in 89 countries, and evaluated zoogeographic, morphological, osteometric, stratigraphic, contextual, iconographic, and textual data. Although previous studies have made claims for an early origin of chickens, our results suggest that unambiguous chickens were not present until ∼1650 to 1250 BCE in central Thailand. A correlation between early chickens and the first appearance of rice and millet cultivation suggests that the production and storage of these cereals may have acted as a magnet, thus initiating the chicken domestication process.
The biocultural origins and dispersal of domestic chickens
Joris Peters
a,b,1,2
, Oph
elie Lebrasseur
c,d,e,1
, Evan K. Irving-Pease
d,f
, Ptolemaios Dimitrios Paxinos
a
, Julia Best
g,h
, Riley Smallman
i
,
C
ecile Callou
j
, Armelle Gardeisen
k
, Simon Trixl
a,l
, Laurent Frantz
m,n
, Naomi Sykes
i
, Dorian Q. Fuller ()
o,p
, and Greger Larson
d,2
Edited by Fiona Marshall, Washington University in St. Louis, Saint Louis, MO; received December 13, 2021; accepted March 8, 2022
Though chickens are the most numerous and ubiquitous domestic bird, their origins,
the circumstances of their initial association with people, and the routes along which
they dispersed across the world remain controversial. In order to establish a robust spa-
tial and temporal framework for their origins and dispersal, we assessed archaeological
occurrences and the domestic status of chickens from 600 sites in 89 countries by
combining zoogeographic, morphological, osteometric, stratigraphic, contextual, icono-
graphic, and textual data. Our results suggest that the rst unambiguous domestic
chicken bones are found at Neolithic Ban Non Wat in central Thailand dated to
1650 to 1250 BCE, and that chickens were not domesticated in the Indian Subconti-
nent. Chickens did not arrive in Central China, South Asia, or Mesopotamia until
the late second millennium BCE, and in Ethiopia and Mediterranean Europe by
800 BCE. To investigate the circumstances of their initial domestication, we corre-
lated the temporal spread of rice and millet cultivation with the rst appearance of
chickens within the range of red junglefowl species. Our results suggest that agricultural
practices focused on the production and storage of cereal staples served to draw arboreal
red junglefowl into the human niche. Thus, the arrival of rice agriculture may have rst
facilitated the initiation of the chicken domestication process, and then, following their
integration within human communities, allowed for their dispersal across the globe.
domestication jchickens jdispersal jhuman niche
Despite the global ubiquity and cultural importance of chickens (Gallus gallus domesticus L.,
1758), the timing and circumstances of their domestication and subsequent dispersal remain
both obscure and controversial (1). Although the spatiotemporal patterns of chicken dispersal
following their domestication have been addressed by several studies (24), two key publica-
tions (5, 6) pertaining to the early history of chicken domestication are almost always cited
but rarely challenged (SI Appendix, Table S1). In brief, the rst argues for a Southeast Asian
and possible South Indian origin for chickens (5), and the second claims that domestic
chickens rst appeared in northern China before following a northern trajectory toward
Europe (6). Both of these oft-cited syntheses summarized, but did not critically assess either
the original osteological and stratigraphic data or the cultural implications for the presence of
chickens across the Old World.
Inferring chicken origins and dispersal have primarily been hampered by a paucity of
archaeological remains, and more specically, issues related to excavation and recovery
biases, faunal identication, and dating (7). Excavations that do not consistently
employ ne sieving, for example, are unlikely to systematically retrieve chicken bones.
When bird remains are recovered, condent identication can be problematic in the
absence of reference collections, since chicken bones are difcult to distinguish from
other related galliform species. Although early Holocene bird remains from northern
China were initially claimed to be chickens (8), a reanalysis of those bones based upon
discrete osteomorphological criteria demonstrated that the specimens were derived
from pheasants (1, 9). More generally, ongoing hybridization between wild red jungle-
fowl (G. gallus) populations and those birds living in human settlements (10) compli-
cates the task of identifying early poultry husbandry in the archaeological record.
The low density of excavated archaeological sites, specically within the distribution
of red junglefowl, means that the earliest shifts in the relationship between people and
these birds remain opaque. Additionally, chicken bones are prone to taphonomic loss
through scavenger activity (11) and stratigraphic displacement (12). The latter can lead
to invalid claims regarding their dating and cultural context, and a recent program of
radiocarbon dating demonstrated that numerous early claims for the rst appearance of
chickens in Europe were spurious (13).
Regarding their geographic origin, genetic studies have demonstrated that of the
four extant junglefowl species, chickens were primarily derived from the red junglefowl
(14, 15). A more recent study analyzed 863 genomes from modern Gallus specimens,
including all ve subspecies of red junglefowl, and identied the subspecies Gallus gallus
Signicance
Chickens are the worldsmost
numerous domestic animal. In
order to understand when, where,
and how they rst became
associated with human societies,
we critically assessed the domestic
status of chicken remains
described in >600 sites in 89
countries, and evaluated
zoogeographic, morphological,
osteometric, stratigraphic,
contextual, iconographic, and
textual data. Although previous
studies have made claims for an
early origin of chickens, our results
suggest that unambiguous
chickens were not present until
1650 to 1250 BCE in central
Thailand. A correlation between
early chickens and the rst
appearance of rice and millet
cultivation suggests that the
production and storage of these
cereals may have acted as a
magnet, thus initiating the chicken
domestication process.
Author contributions: J.P., E.K.I.-P., and G.L. designed
research; J.P., O.L., E.I-.P., P.D.P., J.B., R.S., C.C., A.G.,
S.T., L.F., N.S., D.Q.F., and G.L. performed research; J.P.
analyzed data; and J.P., O.L., E.I-.P., P.D.P., J.B., R.S.,
C.C., A.G., S.T., L.F., N.S., D.Q.F., and G.L. wrote the
paper.
The authors declare no competing interest.
This article is a PNAS Direct Submission.
Copyright © 2022 the Author(s). Published by PNAS.
This open access article is distributed under Creative
Commons Attribution-NonCommercial-NoDerivatives
License 4.0 (CC BY-NC-ND).
1
J.P. and O.L. contributed equally to this work.
2
To whom correspondence may be addressed. Email:
joris.peters@palaeo.vetmed.uni-muenchen.de or greger.
larson@arch.ox.ac.uk.
This article contains supporting information online at
http://www.pnas.org/lookup/suppl/doi:10.1073/pnas.
2121978119/-/DCSupplemental.
Published June 6, 2022.
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RESEARCH ARTICLE
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ANTHROPOLOGY OPEN ACCESS
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spadiceus as the most likely progenitor of domestic chickens
(16). This result suggests that the chicken domestication pro-
cess began within the distribution of this subspecies in South-
east Asia (Fig. 1 and SI Appendix, Fig S1). Genomic analyses
further suggested that the divergence between the ancestral
population of modern domestic chickens and G. gallus spadiceus
occurred between 12,800 and 6,200 y ago. Importantly, this
range cannot be equated with the initiation of a domestication
process. Instead, this timeframe represents the divergence
between G. gallus spadiceus and the lineage from which domes-
tic chickens were derived (17), and thus represents an upper
bound on the chicken domestication timeframe.
Here, in order to establish a robust spatial and temporal
framework for the early occurrence of chickens both within and
beyond the range of red junglefowl, including Africa and Ocea-
nia, we reevaluated reports of chicken remains from >600 archae-
ological sites in 89 countries. We did so by assessing the claims
for chickens in their chrono-cultural and geographic contexts, and
wherever possible, by reassessing the taxonomic determination of
existing specimens by measuring either published photographs or
the actual bones. We then combined zoogeographic, contextual,
and osteometric data to conrm or question the domestic status of
the birds. We also critically reviewed the stratigraphic position of
each of the remains and assessed their intrusive potential. We com-
plemented these efforts by investigating iconographic, written, and
linguistic records pertaining to chickens. In addition, we interpreted
these records within the context of the ecological characteristics and
distributions of all the jungle fowl subspecies.
These analyses allowed us to generate two datasets: a compre-
hensive table of archaeological remains consisting of chickens that
were condently assigned as domestic using conservative measures
(SI Appendix,TableS2), and a list of remains whose identication
or stratigraphic position was ambiguous (SI Appendix,TableS3).
We then correlated the resulting spatiotemporal pattern of archaeo-
logical chickens with human societies and their subsistence strategies.
This correlation allowed us to address the process, circumstances,
and cultural context in which the initial shift in the humanchicken
relationship that led to domestication took place, and the contexts of
their subsequent translocations.
Results
Early Domestic Chickens within the Distribution of Red Junglefowl.
Chicken bones are rare in prehistoric archaeofaunal assemblages
from mainland Southeast Asia. One exception is the Neolithic
moated site of Ban Non Wat (1650 to 1250 BCE) in north-
east Thailand, where >95% of the avifauna has been ascribed
to Gallus. Given the presence of other domesticates, including
pigs and dogs, and the fact that people would have had access
to a highly diverse avifauna in this ecological setting, the
Fig. 1. A map depicting the distribution of both the gray and Ceylon junglefowl species and three subspecies of red junglefowl: G. gallus murghi,G. gallus
spadiceus, and G. gallus jabouillei. The distribution of G. gallus gallus is depicted as the remainder of mainland southeast Asia and Sumatra following the
general distribution in ref. 16. The G. gallus murghi distribution follows that of SI Appendix, Fig. S1, which draws on published maps in ornithological sources
and the Global Biodiversity Information Facility (GBIF) records (119121). For G. gallus spadiceus and G. gallus jabouillei, the GBIF records were augmented by
specimens with genetic data reported by refs. 16 and 122.
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abundance of Gallus remains has been interpreted as evidence
that they may represent a domestic population (18, 19). Although
their domestic status remains unresolved (20), two other factors
lend weight to the initial assessment. First, an exceedingly high
proportion of juvenile bones are present in the assemblage. Sec-
ond, these birds were deposited as grave goods alongside domestic
pigs, dogs, and bovines in human burials at Bronze Age Ban Non
Wat (1038 to 950 BCE) (18) and Ban Na Di (800 to 500
BCE) (21) (SI Appendix,TableS2).
In the Indian subcontinent, the identication of Gallus remains
is problematic due to the co-occurrence of both an endemic red
junglefowl subspecies (Gallus gallus murghi)andthegrayjungle-
fowl (Gallus sonneratii). (Fig. 1 and SI Appendix,Fig.S1). Gallus
remains from forager dumps on the Middle Ganges 4500 to
2000 BCE likely belong to wild birds (22), and domestic chickens
have been claimed to feature in the assemblages associated with
farming communities of the Mature Harappan culture 2600 to
1900 BCE in the Indus River basin (5, 22, 23). Our analysis
demonstrates that key specimens from sites belonging to this
cultural phase including Mohenjo-daro (24) and Harappa (25),
have either been misidentied, or are too large to be condently
categorized as prehistoric domestic birds (SI Appendix,TableS3).
Extensive excavations at Harappa have also failed to identify any
new Gallus bones (26).
Our reappraisal of published chicken bone specimens in
Chalcolithic and Bronze Age sites across India and Pakistan
dating 3500 to 1000 BCE undermines both the species iden-
tication (since they are located in the distribution range of
gray junglefowl) (SI Appendix, Fig. S2) and the domestication
status of these remains (SI Appendix, Table S3). At the site of
Daimabad, India, Gallus is absent from earlier Savalda and
Harappan levels, but prominent in the subsequent early Jorwe
levels dating from 1500 to 1200 BCE (27). Gallus bones have
been reported from several other sites of the Jorwe period
(Nevasa, Inamgaon, Tuljapur Garhi, Walki), as well as contem-
poraneous sites farther south (Southern Neolithic Period III)
(22). This period appears to coincide with historical linguistic
reconstructions for the Dravidian languages in South India,
since each of three linguistic subphyla have distinct etyma, thus
indicating that chickens became widespread after these lan-
guages diverged 1200 BCE (1500 to 500 BCE) (28, 29).
This timing is also consistent with Vedic texts in which chick-
ens are unattested prior to 1200 BCE (30). Collectively, these
lines of evidence suggest that poultry farming in South Asia
was a post-Harappan development.
Early Domestic Chickens Beyond the Distribution of Red Junglefowl.
East and Northeast Asia. Previous claims for the presence of
chickens in the Yellow River basin 11,000 to 8,000 y ago (8)
are not supported by either the climatic, palynological, or
archaeofaunal records, all of which suggest the environmental
conditions in the region were not conducive to the thermo-
philic Gallus (1, 31). In addition, a reanalysis of the archaeolog-
ical bones determined the remains did not belong to chickens
(1, 9). In fact, the earliest remains that can condently be
attributed to domestic chickens in East Asia date to the late
Shang Dynasty, which spanned 1350 to 1046 BCE (SI
Appendix, Table S2). In Japan, chickens did not arrive until the
Middle Yayoi 100 BCE to 100 CE, a timing consistent with
their rst appearance in the Korean Peninsula (1).
Central Asia. InCentralAsia,theearliestevidenceofchickens
thus far consists of depictions of cockerels in the Pazyryk kurgans
of the Altai Mountains dated to 500 to 300 BCE (32), and
chicken remains have been identiedinthelatefourthcentury
BCE fortress of Kurgansol (Uzbekistan) built by Alexanders
troops (33). The absence of chickens in this region prior to the
late Iron Age 800 to 600 BCE (34, 35) contradicts the long-
standing hypothesis that poultry farming spread west and north
from the Yellow River basin during the Neolithic across Asia to
Europe (6).
Island Southeast Asia and the Pacific. The earliest condently
dated chicken remains from archaeological sites in the Solomon
Islands, Vanuatu, and Tonga in Near Oceania have been directly
dated to the early rst millennium BCE (36). Their appearance at
these sites so far East suggests they must have been present in
Island Southeast Asia prior to this date. And although historical
linguistic studies have suggested that chickens were rst intro-
duced beyond continental Peninsular East Asia 2000 BCE (37),
the earliest zooarchaeological appearance in the Indonesian Banda
Islandsisonly700 CE (38) (SI Appendix,TableS2). In Remote
Oceania, chickens arrived with people in Hawaii 1200 CE (36),
about 500 y prior to the earliest records of the European introduc-
tion of poultry in Australia (1788 CE) (39) and New Zealand
(1773 CE) (40).
Southwest Asia. The zooarchaeological record in Southwest Asia
is difcult to interpret. Although isolated chicken remains have
been claimed on Bronze Age settlements, radiocarbon dating
has revealed that specimens excavated at Korucutepe, Turkey
(12) and Jerusalem, Israel were intrusive (SI Appendix, Table
S3). The linguistic and iconographic records in this region are
also problematic. The term dar meluh
˘h
˘a(francolins from
meluh
˘h
˘a) mentioned on mid-third millennium BCE cunei-
form tablets, has been interpreted to refer to chickens, since
meluh
˘h
˘ameant Indus Valley (41). Our reassessment of the
zooarchaeological evidence from the Indian subcontinent implies
that domestic fowl rst appeared after the Harappan civilization,
and that these translations may therefore be inaccurate. The birds
Akkadian name, su-la-mu, describes it as dark-colored (42), which
could correspond instead to black francolins (Francolinus francolinus)
(43). Dark plumage could also imply gray junglefowl, a species
common in the Indian subcontinent generally and in the Indus
region specically (44) (SI Appendix,Fig.S1).
Toward the end of the Bronze Age 1,200 BCE, a depiction
of Gallus from the temple area of Ishtar, Iraq (an ancient Meso-
potamian deity) suggests a religious motive for its introduction.
Scattered bone nds across Mesopotamia and the Levant post-
dating the Late Bronze Age collapse (45), however, imply the
adoption of chicken husbandry during Iron Age I (1150 to
965 BCE). Their increasing numbers at Neo-Assyrian, Urar-
tian, Phrygian (SI Appendix, Table S2), and Levantine sites
(46) indicate growing economic interest in the speciesexploita-
tion during the Iron Age II period (965 to 530 BCE). The
zooarchaeological record corroborates an increased visibility of
Gallus in the iconographic record, including a depiction on the
orthostat of Tell Halaf (present day Syria) dated to 900 BCE
(47), and on numerous cylinder seals between 800 to 600
BCE (41, 48). A land of the chickens(probably Media,
located in modern northwest Iran) (49) is mentioned in the
royal inscriptions of Tiglath-Pileser III (745 to 723 BCE), and
the phrase chickens being seen in the cityappears in the sev-
enth century BCE omen series
Summa
alu (50).
Northeast Africa. Zooarchaeology, historical texts, historical lin-
guistics, and iconography suggest that chickens followed multi-
ple routes into Africa. Since the Annals of Pharaoh Tuthmose
III (1479 to 1425 BCE) mention a tribute from Mesopotamia
of four birds that do bear every day(51), it has been suggested
that these birds were chickens and that they were rst introduced
into North Africa via the Nile Valley. This interpretation seems
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unlikely since the zooarchaeological record indicates that chickens
were not present in contemporaneous Mesopotamia. This quota-
tion more likely refers to either geese or ducks (41), especially
since the earliest chickenreported from a contemporaneous
New Kingdom Theban tomb dating to 1550 to 1070 BCE (52)
is a misidentied duck (SI Appendix,TableS3). The determina-
tion of late second millennium BCE Gallus depictions as a domes-
tic chicken on a 19th Dynasty (1292 to 1189 BCE) ostracon and
a 20th Dynasty (1189 to 1077 BCE) silver bowl (53) are also
problematic, since these depictions could represent wild junglefowl
species. It was not until the Achaemenid period, at least 600 y
later (550 to 330 BCE), that chicken remains have been identi-
ed in lower Egypt (54, 55). Under Ptolemaic rule (305 to 30
BCE), fowl husbandry intensied and spread from the Nile delta
upstream to the region of the rst cataract (56, 57).
Direct radiocarbon-dated chicken bones from Pre-Aksumite
northern Ethiopia (800 to 600 BCE) demonstrate that, prior
to their presence in Achaemenid Egypt, chickens had been
translocated, likely across the Red Sea, to the Horn of Africa
(58). South of the Horn, however, the earliest evidence for
Gallus appears more than a millennium later (59), and arrived
alongside the black rat, rice and mungbean that originated
from the Indian subcontinent (60, 61). From coastal southeast-
ern Africa, poultry farming dispersed both into the interior of
Africa (Fig. 2), and to the Comoro Islands and Madagascar by
750 to 900 CE.
The chickens name in Malagasy has no roots in any of the
western Austronesian languages. Instead, linguistic elements in
neighboring Comorian indicate a connection to Bantu, and hence
an introduction from coastal Africa (62), which is supported by
theclosegeneticafnity of Malagasy and East African chickens
(63). Austronesians were present in Madagascar by 500 CE and
may have arrived with chickens. If so, those founder ocks did
not survive, and the birds Austronesian name was lost. In Mauri-
tius, chickens were only introduced in 1598 CE by the Dutch
East India Company (64).
North and West Africa. In North and West Africa, linguistic
studies have suggested that chickens possibly dispersed from
East Africa through the Sahara (65). The zooarchaeological
record, however, demonstrates that chickens were present in
West African Mande-speaking territory prior to their arrival
in the northern Sahel (66, 67) (SI Appendix, Table S2). Given
that fth century BCE Phoenician seafarers (68) and Roman
merchants both established trading posts on the Moroccan
Atlantic coast (69) or somewhat inland (70), it is possible that
chickens were introduced from Berber-speaking Northwest
Africa to West Africa primarily by maritime trade. The subse-
quent dispersal from coastal West Africa upstream along the
drainage basins of Senegal, Gambia, and other rivers could then
explain the mid-rst millennium BCE occurrence of chicken
bones in Mande-speaking territory.
Europe. In Europe, numerous claims have been made for the pres-
ence of chickens in Neolithic and Bronze Age sites (7174). A
recent study reevaluated key specimens used to postulate early
chicken husbandry, but direct radiocarbon dating of the remains
revealed that all the presumed Neolithic to Bronze Age chicken
remains in Bulgaria, Greece, and France were intrusive, and that
of these, the earliest sample dated to only 137 to 327 CE (13).
This study also established the spatiotemporal pattern of the
dispersal of chickens across Europe. Greek seafarers transported
chickens across the northern Mediterranean and the earliest directly
dated chickens in southern Europe have been excavated in Italy
from two Greek colonies dated to 776 to 540 BCE. Phoenician
merchants introduced chickens to the southern Mediterranean,
including the Balearic Islands and the southern Iberian Peninsula
during this period (75) (SI Appendix,TableS2). Trade between
Fig. 2. A map depicting the earliest condently assigned chicken remains across Eurasia, Africa, and Oceania alongside a spatial kriging interpolation of the
timing of the arrival of chickens. The inference was performed using 100 independent, condently assigned, and dated chicken remains listed in SI Appendix,
Table S2. Each data point was placed on a spatial grid with resolution 10 min of a degree. Solid dots represent samples used in the spatial kriging, and hol-
low dots represent additional sample locations not incorporated into the interpolation. The colored shading indicates the inferred age for the introduction
of chickens across the map. Gray areas indicate locations with a SE above the maximum threshold (see SI Appendix for a full description of the methods
used to produce the map).
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western Mediterranean craft centers and early Celtic communities
promoted the chickens northward expansion. By the late sixth/
early fth century BCE, chickens were present in the Upper
Rhine and Danube River basins (76) (Fig. 2), and in southeast
England (13). By that time, chickens had also been introduced to
the coastal Black Sea region through Greek trade networks, sug-
gesting that the chickens initial dispersal in Europe was primarily
through maritime routes.
Chicken farming became rmly established in western conti-
nental Europe during the late Iron Age and under Roman cul-
tural inuence in the Low Countries (SI Appendix, Table S2).
Chickens arrived in coastal southwest Sweden at the onset of
the rst millennium CE or slightly earlier, before dispersing
during the Migration period (400 to 550 CE). The chickens
rst appearance in the eastern Baltic region dates to the late
Pre-Roman Iron Age, where these remains may have been asso-
ciated with a burial (77). The presence of this bird may represent
a trade import, however, since chickens were clearly exploited
across the Baltic States and in Finland by the late Iron Age (600
to 800 CE) (Fig. 2 and SI Appendix,TableS2). Throughout
much of early medieval Europe, poultry farming likely beneted
from the spread of Christianity (78), and chickens were also asso-
ciated with early monastic contexts in Ireland (SI Appendix,TableS2).
Discussion
Assessing the Spatiotemporal Pattern of Chicken Domestication.
Two initial hypotheses proposed separate temporal and geographic
origins of domestic chickens. Zeuner (5) argued that domestic
chickens were present in the Indus Valley during the mature
Harappan period (2600 to 1900 BCE) and subsequently intro-
duced to Mesopotamia. Based upon the presumption that archae-
ological bird remains dated to the sixth millennium BCE in
Neolithic northern China were chickens, West and Zhou (6)
claimed that chicken domestication must have taken place in
Southeast Asia prior to this before being translocated into China,
and then farther west following a northern route. A more recent
study concluded that red junglefowl were domesticated in the
Yellow River basin shortly after the onset of the Holocene (8) (SI
Appendix,TableS1).
With respect to South Asia, the claim that chickens were pre-
sent within the Indus Valley Civilization was based upon two
bone remains from Harappa (25) and four from Mohenjo-daro
(24), as well as an incomplete hengurine from Mohenjo-
daro (23, 79). Our reanalysis of the two Harappan bones shows
that one is morphologically inconsistent with Gallus, and the
others taxonomic classication is ambiguous. The taxonomic
classication of three of the four fragmentary bird bones from
Mohenjo-daro is also questionable. In addition, all of these
bones, including a completely preserved femur, pertain to indi-
viduals that signicantly exceed the size of prehistoric chickens.
Finally, all four specimens have been collected in upper strata.
Given the propensity for chicken bones to move between strati-
graphic boundaries (13), its possible these remains are recent
intrusions. These lines of evidence call into question the
assumption that poultry farming was present in Bronze Age
Mohenjo-daro (SI Appendix, Table S3).
Excavations conducted in other Indus Valley Civilization sites
and contemporaneous settlements produced additional chicken
bones, especially in Saraushtra (SI Appendix,Fig.S2). These speci-
mens were classied as domestic fowl based on the absence of
modern wild jungle fowl populations in the region (22) and the
aforementioned incorrect claim for fowl husbandry in the Indus
Valley. Although currently located beyond the present-day natural
distribution of both red and gray junglefowl (Fig. 1 and SI
Appendix,Fig.S1), this region shares similar ecological characteris-
tics and borders the region where gray junglefowl is extant (80). It
is therefore likely that the natural range of gray junglefowl extended
into the Indus River basin during the Mid-Holocene, and that
these remains derive instead from local wild populations. This
conclusion is supported by the presence of other fauna present
in zooarchaeological assemblages, or in Harappan art (including
Indian Hog Deer: Axis porcinus;SwampDeer:Cervus duvauceli;
and the Indian Rhinoceros: Rhinoceros unicornis), indicating rel-
atively larger distributions of several fauna in the past (81). The
combined weight of this evidence suggests that, contrary to the
long-standing hypothesis, chickens were not domesticated in
the Indus Valley.
Regarding northern China, subsequent reevaluations of the
galliform remains (1, 9), as well as photographs and drawings,
demonstrate that they are pheasant bones (2, 82). In addition,
high-resolution climate and precipitation records from temper-
ate Holocene East Asia, and the habitat requirements of the
vertebrate taxa associated with the pheasants, suggest that the
subtropical forest habitat conducive to thermophilic red jungle-
fowl did not extend into northern China during the Holocene
climatic optimum (1). Finally, mitochondrial analyses of mod-
ern breeds support a late dispersal scenario of chickens into
northern China (83). Our analysis thus supports a much later
arrival in this region consistent with the rst appearance of
chickens in Japan in the early rst millennium CE, and in
Mongolia during the early second millennium CE.
Our combined reanalyses of zooarchaeological, linguistic,
genetic, and iconographic evidence suggests the following sce-
nario. The rst chickens were likely derived from a population
of the subspecies G. gallus spadiceus, whose current range spans
southwestern China, northern Thailand, and Myanmar (16)
(Fig.1). The rst unambiguous chicken bones in the archaeo-
logical record are present within the faunal assemblage at
Neolithic Ban Non Wat in central Thailand, and date to
1650 to 1250 BCE. Once incorporated into human societies,
chickens dispersed into and beyond the range of other Gallus
subspecies and species. The evidence presented here demon-
strates that chickens did not appear in archaeological contexts
within Central China, South Asia, or Mesopotamia until the
late second millennium BCE, just before their initial presence
in Melanesia. By 700 BCE, chickens had arrived in Ethiopia
and Mediterranean Europe (Fig. 2). This western dispersal was
substantially more rapid than the establishment of chicken
populations in temperate regions present in higher latitudes
(Fig. 2). Overall, our analyses indicate a temporal origin and
spread of chickens that substantially postdates many of those
suggested by previous studies (6, 8).
A Hypothesis for the Process of Chicken Domestication.
Within the native range of red junglefowl, many Southeast Asian
languages refer to chickens as bamboo fowl,given how readily
they take advantage of cyclical bamboo mass owering and seed-
ing events (10). Red junglefowl are also known to consume rice
grains (Oryza sp.) (44, 84), a staple dietary component of South-
east Asian domestic chickens (85). Rice and millet (Setaria
italica) were cultivated by mixed foraging and cereal producing
Neolithic communities in mainland Southeast Asia (8688). An
analysis of the material culture (including pottery decoration)
associated with these agriculturalists suggests that rice and millet
cultivating communities dispersed from the Yangtze valley (8992)
into southern China, where they arrived by the mid-third millen-
nium BCE (90, 93). From there, they continued into peninsular
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Southeast Asia, where people following distinct dispersal path-
ways emphasized either rice or millet (94).
Land reclamation for cereal cultivation led to the replace-
ment of primary forest by secondary vegetation, a habitat more
suitable for red junglefowl. Outside of bamboo thickets, red
junglefowl are known to thrive in slash-and-burn agricultural
systems (10). The novel presence of cultivated elds, fallow
elds (necessary for either millet or rainfed rice), cereal harvest
residues, remainders of human food preparation and consump-
tion, invertebrates associated with keeping pigs and cattle, and
other aspects of the human niche may have attracted red jun-
glefowl to human settlements and their immediate catchment.
The long-term abundance of cereals within the human niche
would have led to dramatic shifts in selective pressure that
lasted multiple generations, including a relaxation of selection
against larger clutch size, as well as increased selection against
territoriality in cocks (10). These conditions likely also facili-
tated larger bird population densities near farming communi-
ties, followed by subsequent accommodationof birds within
the village. The availability of cultivated cereals may have there-
fore catalyzed a shift in the relationship between people and red
junglefowl consistent with the commensal pathway (95, 96).
The current archaeobotanical evidence indicates that sites
with rice cultivation appeared within the distribution of G. gallus
spadiceus [the subspecies recently identied as the most likely pro-
genitor of chickens (16)] from about 2000 BCE in two regions:
southern Yunnan and northeast Thailand (Fig. 3 and SI Appendix,
Table S4). Although cereal farming was also present at this time
within the inferred distributions of Gallus gallus jabouillei and
G. gallus gallus, there is as yet no evidence for the early presence of
domestic chicken populations. Although there is conrmation of
early rice farming near coastal areas and in low-lying wetlands (86,
94), the zooarchaeological records in Neolithic Vietnam, for
example, show evidence for the hunting of wetland birds, but lack
remains of Gallus (97). In more interior zones, early rainfed rice/
millet cultivation likely spurred a tighter relationship between peo-
ple and G. gallus spadiceus, as attested by the earliest conrmed
chickenbonesatBanNonWatandNonNokTha(SI Appendix,
Table S2)(98).Becausethericewasrainfed,itwouldhave
required more land area and fallow cycles relative to later, more
productive irrigated rice (87), and these conditions would have
created large areas of secondary thicket vegetation. In the northern
region of the G. gallus spadiceus distribution, similar processes
were likely possible, but detailed avifaunal studies at prehistoric
archaeological sites in Yunnan have not yet been carried out.
In regions north of Yunnan, wet rice agriculture, characterized
by small-scale, intensive wetland elds in the Yangtze (87, 99), is
present as early as 4000 to 3000 BCE, and these wet paddyeld
systems spread throughout central China during the Neolithic.
Chickens, however, were absent in the Neolithic (100) and only
appear 1000 BCE (Fig. 3). Relative to wet rice, dry rice and
millet cultivation that predominate in the tropical south is
Fig. 3. A map depicting the distribution of dated archaeological rice nds taken from the revised Rice Archaeological Database compiled by D.Q.F. and
colleagues, RAD 2.0 (90), with newly added archaeological records and cleaned reports with associated dates that appear too early based on current under-
standing of archaeological chronology. This is especially the case in mainland Southeast Asia where most of the arrival of cereal agriculture is now thought
to be 2500 BC for northern Vietnam and southernmost China only and 2000 BC for the rest of the region (94, 123).
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characterized by more extensive mosaics of eld and fallow, a
nichemoresuitedtowildandcommensalredjunglefowl.
A Combined ChickenRice Dispersal Across Asia and Africa.
Following the integration of chickens and human agricultural
societies, the correlated spatiotemporal patterns of rice and
chicken dispersal across Asia is striking. In South Asia for exam-
ple, sedentism became widespread in the Ganges plains in the
second millennium BCE (22, 101), and domesticated rice,
wheat, barley, and other grain crops began to dominate the
economy (102, 103). At this time, agriculture in the Deccan
focused on small millets, beans, and occasionally wheat and
barley (22, 101). It is during this period, perhaps from the later
second millennium BCE, that the subsistence context would
have been ideal for chickens or commensal jungle fowl, and
this timing corresponds with the arrival of bird remains unam-
biguously identied as chickens in the Indian subcontinent.
While there is evidence for proto-indica rice management by
huntershergatherers in the middle Ganges plains predating
2000 BCE (102, 104, 105), proto-indica rice was managed in
natural, seasonal wetlands that were unlikely to attract Gallus in
large numbers. This is illustrated by a dearth of Gallus remains
and a low level of both cereal production and livestock hus-
bandry in the archaeological record (22). The domesticated
indica rice introduced 1600 to 1500 BCE in the Upper and
Middle Ganges (104, 105) was typically rainfed in more exten-
sive systems with periods of fallow (102, 106), and thus more
attractive to fowl. Recent genomic evidence derived from mod-
ern populations suggested that although modern domestic
chickens in South Asia possess signatures associated with the
local subspecies G. gallus murghi, these ancestral afnities are
the result not of a local, independent domestication process,
but the result of admixture with introduced domestic chickens
derived from G. gallus spadiceus (16).
In Iron Age Mesopotamia, rice and millet cultivation may
also have been linked to the initiation and intensication of
poultry farming. For example, the cultivation of Chinese millets
(Panicum miliaceum,Setaria italica) began in the later second
millennium BCE (107109) and intensied after 1000 BCE
(110, 111), precisely when poultry husbandry becomes visible
archaeologically (Fig. 2 and SI Appendix, Table S2). Middle
Assyrian texts also conrm that irrigated rice became estab-
lished in Syria by 1100 BCE, and references to this practice
increased from the eighth century BCE (112). Thus, alongside
the diversication of grain crops, chickens may represent an
additional element of the broadening of Near Eastern subsis-
tence practices after the late Bronze Age collapse (45).
There is also a correlation in Africa between the appearance
of chickens and rice agriculture. The translocation of chickens
to coastal Southeast Africa and the Indian Ocean islands in the
eighth/ninth centuries CE coincides with the introduction of
Asian crops, such as rice, tree cotton, and mung bean (113). In
addition, archaeobotanical evidence in the Niger Basin illus-
trates the prominent role of cereal diversication, including
more widespread rice cultivation and increasing urbanism
300 to 900 CE (114), a temporal window that coincides with
the rst appearance of chickens.
Conclusion
Our reassessment of archaeological, iconographic and textual evi-
dence indicates rst that the origins and dispersal of domestic
chickens both within and beyond the native range of red jungle-
fowl was, relative to many farm animals, a relatively late
phenomenon (Fig. 2). Our conservative approach suggests that
chickens were incorporated into human societies as domestic birds
by 1500 BCE in peninsular Southeast Asia, and that they then
rapidly spread south into Island Southeast Asia and west across
South Asia and Mesopotamia to Europe and Africa. Additional
archaeological investigations are necessary to test this chronology,
and new excavations may reveal not only an earlier association
between humans and red junglefowl, but also earlier human-
mediated translocations of chickens across the globe.
The attractiveness of rice and millet agriculture to red jungle-
fowl may have facilitated both the initiation of the domestica-
tion process, and the subsequent dispersal of domestic chickens.
Initially, the more extensive cultivation systems that created
mosaics of secondary tropical scrub (fallows) alongside cereal
elds may have been important, whereas later, more intensive
rice agriculture would have provided grain surpluses to support
larger populations of domestic birds. The year-long availability
of grains and associated foodstuffs may thus have served to draw
these arboreal birds into a human niche. Archaeologically, the rst
appearance in peninsular Southeast Asia of both rice and unam-
biguous domestic chickens is also consistent with the recent con-
clusion that domestic chickens were derived from the subspecies
G. gallus spadiceus (16).
Grain production within a human niche has been shown to
also attract other birds. For example, a recent genetic and isotopic
study (115) of pheasant (Phasianus colchicus)boneremains(origi-
nally presumed to be chickens) from the site of Dadiwan in north-
ern China demonstrated that the birds subsisted on cultivated
millet, thus suggesting that this grain attracted different gallina-
ceous birds into village settings. By providing predictable feeding
opportunities across the region for distinct bird species (including
red junglefowl), grain cultivation may have acted as a magnet for
seed-eating birds, thereby kickstarting a shift in the commensal
relationship with people that eventually led to domestication.
Future efforts to integrate newly excavated sites and the applica-
tion of direct radiocarbon dating will modify the picture presented
here, while providing insights into the climatic, economic, and
socio-cultural characteristics that inuenced the dispersal of chick-
ens across the globe. From a biological perspective, although rice
and millet cultivation may have facilitated chicken husbandry in
subtropical regions, it was the speciesadaptability and granivorous
diet that led to its adoption by agriculturalists cultivating barley,
wheat, and other cereals in arid and temperate environments. In
addition, the delayed integration of fowl into human settlements
at higher latitudes in Eurasia (Fig. 2 and SI Appendix,TableS2)
suggests that the generally thermophilic species adapted more
slowly to colder climates.
Culturally, the presence of early iconography and chicken bones
in Mesopotamian and western European palatial architecture,
noble households, elite burials, sanctuaries, and other divine con-
texts, suggests that chickens were initially associated with elites in
numerous locations (116). A recent study (13) demonstrated that
early chickens in European archaeological contexts appear as com-
plete skeletons. Cut marks on individual chicken bones found as
refuse are not present until several centuries later, suggesting that
the rst chickens were initially revered rather than consumed.
Since the regular keeping of chickens for food did not take place
until centuries after their arrival in Europe, this also undermines
the hypothesis that meat consumption drove the chickenstransi-
tion to a domestic bird.
To test the scenario outlined above, it is necessary to care-
fully document and directly date additional Gallus nds. In
addition, the establishment of osteological identication criteria
separating wild junglefowl from their domestic relatives, and the
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simultaneous generation and analysis of morphological, isotopic,
and ancient genetic datasets is essential to rene the origins and
dispersal of the worlds most ubiquitous domestic bird.
Materials and Methods
Evaluation of Archaeological Chicken Remains. To critically assess and
establish the early appearance of domestic chicken remains across Eurasia, Africa,
and Oceania, this study analyzed the remains described in faunal databases and
publications. We did so by revisiting and critiquing the species identication,
the determination of domestic status, the integrity of the sites stratigraphy, and
the overall zooarchaeological and cultural context of the sites. This conservative
approach allowed us to generate two tables: one which includes the earliest
recorded archaeological remains that we could condently assign as domestic
chickens (SI Appendix, Table S2), and a list of remains whose identication,
dating, and stratigraphic position was ambiguous (SI Appendix,TableS3)
(117, 118). This latter table also described the rationale for the exclusion of the
individual remains from SI Appendix,TableS2. The challenges of compiling this
global dataset included issues related to preservation, identication, recovery,
and dating. More detailed descriptions of each of these topics and how they
inuencedthe compilation of the tables are included in SI Appendix.
Data Availability. Data have been deposited in GitHub (https://github.com/
ekirving/chickens). All other study data are included in the main text and
SI Appendix.
ACKNOWLEDGMENTS. We thank Thierry Argant, Guy Bar-Oz, Laszlo Bartosie-
wicz, Cornelia Becker, Norbert Benecke, Zbigniew Boche
nski, K
evin Bouchit
e,
Hijlke Buitenhuis, Bea De Cupere, Hui Deng, Keith Dobney, Silvia Eccher, Masaki
Eda, Gerhard Forstenpointner, Alfred Galik, Anne Birgitte Gotfredsen, Martin
Heide, Charles Higham, Thomas Higham, Friedhelm Hoffmann, Hitomi Hongo,
Liora Horwitz, Chiori Kitagawa, Stephan Kroll, Ren
eKysel
y, Peter Lape, Yves Lig-
nereux, Mark Maltby, Kristiina Mannermaa, Richard H. Meadow, Arturo Morales,
Elisabeth von der Osten-Sacken, Lee Perry-Gal, Eva Rannam
ae, Lidar Sapir-Hen,
Barbara Stopp, Umberto Tecchiati, Richard Thomas, Katerina Trantalidou, Wim
Van Neer, Yiru Wang, and Chong Yu for constructive criticism and guidance.
G.L., L.F., and O.L. (2017-2018) were supported by a European Research Council
(ERC) starting grant (ERC-2013-StG-337574-UNDEAD), and L.F. and G.L. were
supported by the ERC starting grant (ERC-2019-StG-853272-PALAEOFARM). G.L.,
O.L. (2014-2017), N.S., and J.B. were supported by the Arts and Humanities
Research Council (grant AH/L006979/1). E.K.I.-P. was supported by the Lundbeck
Foundation (grant R302-2018-2155) and the Novo Nordisk Foundation (grant
NNF18SA0035006). O.L. was supported by the European Unions Horizon 2020
research and innovation programme under the Marie Sklodowska-Curie grant
agreement no. 895107.
Author afliations:
a
ArchaeoBioCenter and Institute of Palaeoanatomy, Domestication
Research, and the History of Veterinary Medicine, Ludwig Maximilian University Munich,
80539 Munich, Germany;
b
Bavarian Natural History Collections, State Collection of
Palaeoanatomy Munich (SPM), 80333 Munich, Germany;
c
Centre for Anthropobiology and
Genomics of Toulouse, CNRS UMR 5288, Universit
e de Toulouse, Universit
ePaulSabatier,
31000 Toulouse, France;
d
Palaeogenomics & Bio-Archaeology Research Network, School
of Archaeology, University of Oxford, Oxford OX1 3TG, United Kingdom;
e
Instituto
Nacional de Antropolog
ıa y Pensamiento Latinoamericano, Ciudad Aut
onoma de Buenos
Aires, C1426BJN, Argentina;
f
Lundbeck Foundation GeoGenetics Centre, GLOBE Institute,
University of Copenhagen, 1165 Copenhagen, Denmark;
g
Department of Archaeology and
Anthropology, Bournemouth University, Poole BH12 5BB, United Kingdom;
h
School of
History, Archaeology, and Religion, Cardiff University, Cardiff CF10 3AT, United
Kingdom;
i
Department of Archaeology, University of Exeter, Exeter EX4 4PY, United
Kingdom;
j
Arch
eozoologie, Arch
eobotanique: Soci
et
es, Pratiques et Environnements,
Bases de Donn
ees sur la Biodiversit
e,
Ecologie, Environnement, et Soci
et
es, Mus
eum
National d'Histoire Naturelle, CNRS, 75005 Paris, France;
k
Arch
eologie des Soci
et
es
M
editerran
eennes, UMR 5140, Universit
ePaul-Val
ery, CNRS, LabEx Archimede IA-ANR11-
LABX-0032-01, 34090 Montpellier, France;
l
State Ofce for Cultural Heritage Baden-
Wuerttemberg, 78467 Constance, Germany;
m
Palaeogenomics Group, Institute of
Palaeoanatomy, Domestication Research, and the History of Veterinary Medicine, Ludwig
Maximilian University, 80539 Munich, Germany;
n
School of Biological and Behavioural
Sciences, Queen Mary University of London, London E1 4DQ, United Kingdom;
o
Institute
of Archaeology, University College London, London WC1H 0PY, United Kingdom; and
p
School of Cultural Heritage, Northwest University, 710069 Xian, China
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