Earliest domestication of common millet (Panicum
miliaceum) in East Asia extended to 10,000 years ago
Houyuan Lua,1, Jianping Zhanga, Kam-biu Liub, Naiqin Wua, Yumei Lic, Kunshu Zhoua, Maolin Yed, Tianyu Zhange,
Haijiang Zhange, Xiaoyan Yangf, Licheng Shene, Deke Xua, and Quan Lia
aKey Laboratory of Cenozoic Geology and Environment, Institute of Geology and Geophysics, Chinese Academy of Sciences, Beijing 100029, China;
bDepartment of Oceanography and Coastal Sciences, Louisiana State University, Baton Rouge, LA 70803;cGraduate University of the Chinese Academy of
Sciences, Beijing 100049, China;dInstitute of Archaeology, Chinese Academy of Social Sciences, Beijing 100710, China;eCulture Museum of Cishan, Wuan,
Hebei Province, 036302, China; andfInstitute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China
Edited by Dolores R. Piperno, Smithsonian Tropical Research Institute and National Museum of Natural History, Washington, DC, and approved
March 17, 2009 (received for review January 8, 2009)
The origin of millet from Neolithic China has generally been
accepted, but it remains unknown whether common millet (Pani-
domesticated. Nor do we know the timing of their domestication
and their routes of dispersal. Here, we report the discovery of husk
phytoliths and biomolecular components identifiable solely as
common millet from newly excavated storage pits at the Neolithic
Cishan site, China, dated to between ca. 10,300 and ca. 8,700
calibrated years before present (cal yr BP). After ca. 8,700 cal yr BP,
the grain crops began to contain a small quantity of foxtail millet.
Our research reveals that the common millet was the earliest dry
farming crop in East Asia, which is probably attributed to its
excellent resistance to drought.
Holocene ? origins of agriculture ? phytoliths ? Neolithic ? Cishan
most important and ancient domesticated crops. They were
staple foods in the semiarid regions of East Asia (China, Japan,
Russia, India, and Korea) and even in the entire Eurasian
continent before the popularity of rice and wheat (1–4), and are
still important foods in these regions today (5, 6).
Thirty years ago, the world’s oldest millet remains, dating to
ca. 8,200 calibrated years before present (cal yr BP), were
discovered at the Early Neolithic site of Cishan, northern China.
The site contained ?50,000 kg of grain crops stored in the
storage pits (7–9). Until now, the importance of these findings
has been constrained by limited taxonomic identification with
regard to whether they are from foxtail millet (S. italica) or
common millet (P. miliaceum), because the early reported S.
italica identifications are not all accepted (4, 9–12). This article
presents the phytoliths, biomolecular records, and new radio-
carbon dating from newly excavated grain crop storage pits at the
Cishan site. Large modern reference collections are used to
compare and contrast microfossil morphology and biomolecular
components in different millets and related grass species (13).
The renewed investigations show that common millet agriculture
arose independently in the semiarid regions of China by 10,000
cal yr BP. Our findings contribute to our knowledge of agricul-
tural origins across the globe and have broader implications for
understanding the development of human societies.
The Cishan site (36°34.511? N, 114°06.720? E) is located near
the junction between the Loess Plateau and the North China
Plain at an elevation of 260–270 m above sea level (Fig. 1). The
archaeological site, containing a total of 88 storage pits with
significant quantities (?109 m3) of grain crop remains, was
excavated from 1976 to 1978 (7, 8). Each storage pit included
0.3- to 2-m-thick grain crops, which were well preserved and
found in situ in the 3- to 5-m-deep loess layer (9). All grain
remains have been oxidized to ashes soon after they were
exposed to air. Archaeological excavations also revealed the
remains of houses and numerous millstones (Fig. S1), stone
oxtail millet (Setaria italica) and common millet (or broom-
corn millet; Panicum miliaceum) were among the world’s
shovels, grind rollers, potteries, rich faunal remains, and plant
assemblages including charred fruits of walnut (Juglans regia),
hazel (Corylus heterophylla), and hackberry (Celtis bungeana)
(7–9). Only 214C dates of charcoal from previously excavated
H145 and H48 storage pits yielded uncalibrated ages of 7355 ?
100 yr BP and 7235 ? 105 yr BP, respectively (8). These
remains represent the earliest evidence for the significant use
of dry-farming crop plants in the human diet in East Asia. They
also suggest that by this time agriculture had already been
relatively well developed here.
Early identification assumed these grain crop remains to be
foxtail millet (S. italica). This preliminary identification was
mainly based on a characteristic of very small sizes of grain crop
ash (no charred grains)—often ?2–3 mm in length—that re-
sembled the foxtail millets (Fig. S2), but without any spodogram
evidence for the grain crop remains in the Cishan site (7–10).
These have been reported as the world’s oldest foxtail millet in
the literature (6, 9, 10, 14). However, the millet identification has
been questionable (4, 11, 12, 15), because the macro (ash)
remains were too friable to be observed under the microscope.
Furthermore, very little study has been conducted on the
spodograms or phytoliths of modern millets, so no clear diag-
nostic feature has been used to distinguish foxtail millet from
common millet (12, 16). Previous study has also considered at
some length how the charred dehusked grains of various native
millet species might have been systematically misidentified (3).
Thus, questions remain regarding whether the Cishan grain
crops are from foxtail millet or common millet, or both, because
they cannot be distinguished from the ash vestige (11, 12).
Phytoliths and biomolecular components have provided sub-
stantial empirical evidence demonstrating the considerable an-
tiquity of food production and crop dispersals in many regions
of the world (17–20). Based on our observation and statistics of
the variation of anatomy and silicon structure patterns in the
glumes, lemmas, and paleas occurring in modern cultivated
millets and related grass species collected from different regions
in China (13), we found that 5 key diagnostic characteristics in
phytolith morphology could be used to distinguish between
foxtail millet and common millet (13): (i) a cross-shaped type
phytolith is formed in the lower lemma and glume of S. italica,
Author contributions: H.L. and N.W. designed research; H.L., J.Z., Y.L., K.Z., M.Y., T.Z., H.Z.,
L.S., D.X., and Q.L. performed research; H.L. and Y.L. contributed new reagents/analytic
The authors declare no conflict of interest.
This article is a PNAS Direct Submission.
See Commentary on page 7271.
1To whom correspondence should be addressed. E-mail: firstname.lastname@example.org.
This article contains supporting information online at www.pnas.org/cgi/content/full/
May 5, 2009 ?
vol. 106 ?
no. 18 ?
whereas a bilobe-shaped type is formed in those of P. miliaceum;
(ii) regularly arranged papillae on the surface of the upper
lemma and paleas are peculiar to S. italica; (iii) the epidermal
long cell walls are ?-undulated in S. italica (Fig. 2H), and
?-undulated in P. miliaceum (Fig. 2 B, D, and F); (iv) the endings
structures of epidermal long cells are cross-wavy type in S. italica
and cross-finger type in P. miliaceum; the R value (ratio of the
width of endings interdigitation to the amplitude of undulations)
is higher (0.79 ? 0.12; n ? 3,303) in P. miliaceum than in S. italica
(0.33 ? 0.11; n ? 2,774); and (v) surface ridgy line sculpture of
the upper lemma is peculiar to S. italica. These 5 diagnostic
characteristics used together give the only reliable way of
distinguishing foxtail millet from common millet when only
powder remains are available. In addition, a species-specific
identification of phytoliths is possible for S. italica and P.
miliaceum because they have typically well-defined silica skele-
China. (B) The Cishan site is located on a terrace of the Ming River (Inset). A detailed plan of the west area of Cishan site excavated in 1976–1978, showing the
is presented. (C) A photograph of the newly excavated storage pit CS-V, found on the cliff of the northern terrace. (D) Close-up photograph of the loose layer
of grain crop remains in storage pit CS-III found in situ in the loess layer.
P. miliaceum (B). (C and D) Phytoliths from CS-II storage pit (C), compared with modern ?-II type husk phytoliths from P. miliaceum (D). (E and F) Phytoliths from
BWG (E), compared with modern ?-III type husk phytoliths from P. miliaceum (F). (G and H) Phytoliths from CS-II storage pit (G), compared with modern ?-II type
husk phytoliths from S. italica (H). (I) Bivariate biplot showing coordinates of the 3,303 measurements from epidermal long cells of P. miliaceum and 2,774
measurements from those of S. italica, plotted along axis W (width of endings interdigitation of dendriform epidermal long cells) and axis R (ratios of W to
undulations amplitude of dendriform epidermal long cells), and their classification into 2 groups corresponding to 2 species (P. miliaceum and S. italica) (13).
Also plotted are the fossil samples of husk phytoliths from CS-I-V and BWG, interpreted to be of P. miliaceum origin, dated between ca. 10,300 and ca. 7,500
yr BP. The CS-II, V, and BWG samples contained 0.4–2.83% ?-type husk phytoliths, interpreted to be of S. italica origin, dated to less than ca. 8,700 yr BP.
Scanning microscopic interferometer photographs. (A and B) Phytoliths from CS-I storage pit (A), compared with modern ?-I type husk phytoliths from
www.pnas.org?cgi?doi?10.1073?pnas.0900158106 Lu et al.
tons that are distinguishable from those in Panicum bisulcatum,
Setaria viridis, and Setaria plicata, which have no such demon-
strable patterns (13).
To determine the taxa of foxtail millet and common millet, we
excavated storage pits (CS-I to CS-V) at the Cishan site, and 1
sample (BWG) preserved in a storage bottle from the Culture
Museum of Cishan. These grain crop samples are dated between
ca. 10,300 and ca. 7,500 cal yr BP based on new14C dating
measurements (Fig. 3, Table S1).
All 47 archaeological samples we analyzed contained abun-
dant diagnostic husk phytoliths that can be divided into 2 groups
according to their phytolith assemblages and14C dating results.
The first group, including 27 samples from CS-I, III, and IV, is
dated between ca. 10,300 and ca. 8,700 cal yr BP. All of the husk
phytoliths present are diagnostic of common millet based on
their shapes, ? patterns (Fig. 2 A, C, and E), and average R value
(?0.7) (Fig. 2I). The second group, including 20 samples from
CS-II, V, and BWG, is dated between ca. 8700 and ca. 7,500 cal
yr BP. More than 97% of the husk phytoliths are also diagnostic
of the common millet (97.2% for CS- II, n ? 1,273; 97.5% for
CS-V, n ? 1,000; 99.6% for BWG, n ? 1,000), but a small
quantity (0.4–2.8%) of the husk phytoliths in the second group
can be attributed to S. italica (Fig. 2 G and I).
Fig. 4 shows that the CS-I storage pit contains ?1.5-m-thick
grain crop remains composed of 3 prominent layers of lemma
and palea from common millets alternating with 3 glume plus
reed layers. This storage manner indicates that prehistoric
humans had known how to preserve large volumes of grains in
secure storage. They did so by digging deep storage pits in the
dry loess strata and by covering the floors with thick mats of
millet glumes and reed (Phragmites australis) leaves.
Identification of biomolecular components was mainly based
on our modern reference collection. A recent study has used the
biomolecular components of P. miliaceum, the only miliacin-
exclusive producer reported, as a basis for identification (20).
However, because of the lack of comparable data derived from
S. italica, previous investigators are still not clear how to use
biomarkers to distinguish P. miliaceum from S. italica. In this
study, we examined biomolecular components in 6 samples of
modern Paniceae (see Materials and Methods). The results show
that biomolecular components can be used to distinguish be-
tween P. miliaceum and S. italica based on the presence or
absence of 5 biomarkers—miliacin, ?-amyrin methyl ether
(ME), and 3 pentacyclic triterpene methyl ethers (PTMEs),
although the structures of the 3 PTMEs have yet to be confirmed
(Fig. 5, Fig. S3). The total ion current trace shows the whole
distribution of aromatic hydrocarbons and ethers extracted from
P. miliaceum (Fig. 5A) and S. italica (Fig. 5B) to be in the 54- to
62-min analysis time range. The significant relative abundance of
miliacin (compound 2) (Fig. 5D) was found in both modern
species (89.0 ? 1.64% for P. miliaceum and 33.8 ? 22.2% for S.
italica); the miliacin relative abundances in the aromatic hydro-
carbons and ethers were estimated by measuring the area of the
miliacin peak on the m/z 189 ? 204 ? 218 ion-specific chro-
matogram. However, compounds 1, 4, and 5 are peculiar to P.
miliaceum, and compound 3 is peculiar to S. italica.
Fig. 5 shows that the prominent compound products of the
archaeological samples (BWG, CS-V-03) are compounds 1, 2, 4,
1, 4, and 5 are notably absent from S. italica, which contain
compounds 2 and 3 only. Thus, the distribution pattern of major
biomarkers of BWG and CS-V-03 grain crop remains is similar to
that of modern common millet (Fig. 5A). In 2 archeological
samples, including BWG and CS-V-03, the relative abundance of
miliacin reaches 88.5% and 88.2%, respectively, which is also very
similar to that of common millet. These results provide further
support to our conclusion and suggest that common millet is an
important source of grain crops stored in the storage pits.
According to archeobotanical research, the early charred grains
of common millet occurred during the initial stages of various
Early Neolithic sites (Fig. 1), including Dadiwan (ca. 7.8–7.35 cal
kyr BP) (21), Xinglonggou (ca. 8.0–7.5 cal kyr BP) (22), and
vated at Cishan site. Lab no: GZ, Laboratory of Peking University Accelerator
Mass Spectrometry and Key Laboratory of Isotope Geochronology and Geo-
chemistry, Guangzhou, Chinese Academy of Sciences; CNL, Radiocarbon Lab-
oratory of the Institute of Geology and Geophysics, Chinese Academy of
Sciences; ZK, Laboratory of the Institute of Cultural Relics of the Chinese
Bureau of Cultural Relics (8). Red box, calendar 68% range, by CalPal, Univer-
sity of Cologne Radiocarbon Calibration Program Package (www.calpal.de/).
G, grain crops; C, charcoal.
Carbon-14 dates and chronology-corrected dates of samples exca-
layers of lemma and palea phytoliths from common millets alternating with 3
layers of mixed common millet glumes, reed, and panicoid types. Three
radiocarbon dates were obtained from the grain crops. Ages shown are
dendro-corrected calendar years BP. Minor age reversal is attributed to the
intrusion of roots from the modern grasses growing on the cliff foot face.
Lu et al.PNAS ?
May 5, 2009 ?
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Yuezhuang (ca. 7.87 cal kyr BP) (23) in North China, but foxtail
millet was barely present during these stages. Lee et al. (24) have
speculated that the Early Neolithic predominance of broomcorn
over foxtail millet at Xinglonggou and Yuezhuang ca. 6000 cal
B.C. might be a regional phenomenon, implying that broomcorn
millet might have been domesticated earlier than foxtail millet.
Our analytical results of both phytoliths and biomolecular
components have established that the earliest cereal remains
stored in the Cishan Neolithic sites, during ca. 10,300–8,700 cal
yr BP, are not foxtail millet, but only common millet. After 8,700
cal yr BP, the grain crops gradually contained 0.4–2.8% foxtail
millet. Our study also suggests that common millet was used as
a staple food significantly earlier than foxtail millet in northern
China. It provides direct evidence to show that, by 10,000 cal yr
BP, the early people in northern China had developed various
methods of maintenance and multiplication of millet seeds for
olean-18-en-3?-ol ME, M?440, m/z, 425, 393, 257, 218, 204, 189 (100%), 177, 161, 135, 109, 95, 69], PTME-3 [M?440, m/z, 425, 397 (100%), 365, 261, 229, 218,
[M?440, m/z, 425, 408, 393, 257, 221, 203, 189 (100%), 147, 135, 121, 109, 95] (Fig. S3D), respectively. Mass spectra of PTME-2 (miliacin, olean-18-en-3?-ol ME)
are presented in D.
Total ion chromatogram of extracted aromatic hydrocarbons and ethers from common millet (A), foxtail millet (B), and grain crops from BWG and
www.pnas.org?cgi?doi?10.1073?pnas.0900158106Lu et al.
the next generation, and had known how to store crops of staple
food in secure, dry places of storage pits during the Early
Common millet has the lowest water requirement among all
grain crops; it is also a relatively short-season crop, and could
grow well in poor soils (5, 6, 25). The geographical distribution
that foxtail millet is more common in the semiwet eastern areas,
and its optimal growth occurs at mean annual temperature
(MAT) from 8 to 10 °C and mean annual precipitation (MAP)
from 450 to 550 mm. However, common millet is more adapted
to the drier interior areas, and its optimal growing conditions
occur at MAT from 6 to 8 °C and MAP from 350 to 450 mm (5,
6). The origin and dispersal of millet agriculture is a key problem
closely related to the history of human impact on the environ-
ment and transformation of natural vegetation.
Paleoenvironmental data from the Weinan section (26–29)
(Fig. 1) in the southern part of the Loess Plateau between the
Cishan and Dadiwan sites are crucial for understanding the early
stage of the forager–cultivator transition. The early Holocene
was a period of significant environmental change marked by dry
climate conditions as inferred from sediment texture (26, 28),
magnetic susceptibility (26, 28), pollen (27), phytoliths (28), and
mollusk assemblages (29). These proxy records show an envi-
ronmental transition from cold–dry (ca. 11,000–8,700 cal yr BP)
to warm–wet (ca. 8,700–5,500 cal yr BP) conditions. Many
lacustrine and loess records from the Chinese Loess Plateau to
early Holocene (30–34). Under the drier climate conditions, soil
development was slowed, and the soil developed on the under-
lying older and coarser loess of the glacial period was poor in
nutrients (28). This raises the possibility that common millet was
more significant than foxtail millet in the early stages of food
production in North China because it was more adaptable than
foxtail millet to the dry condition prevailing during the early
Holocene. The common millet cultivation may involve complex
selection by natural forces and human activities, although no
clear evidence has been documented in this region for the
transitions from gathering to cultivation and/or from a wild
ancestor to domesticated common millet (1, 2, 5).
Our research indicates that the earliest significant common
millet cultivation system was established in the semiarid regions
of China by 10,000 cal yr BP, and that the relatively dry condition
in the early Holocene may have been favorable for the domes-
tication of common millet over foxtail millet. Our study shows
that common millet appeared as a staple crop in northern China
?10,000 years ago, suggesting that common millet might have
been domesticated independently in this area and later spread to
Russia, India, the Middle East, and Europe. Nevertheless, like
floodplains of the Lower Tigris and Euphrates was a key factor
in the emergence of civilization, the spread of common millet to
the more productive regions of the Yellow River and its tribu-
development of social complexity in the Chinese civilization.
Materials and Methods
Phytolith Extraction from Archaeological Material. The Cishan site is located on
the north terrace of the Ming River (Fig. 1). The geography of this region
consists of alluvial terraces covered by loess deposits. Five storage pits (CS-I to
sediments (4–5 m thick for CS-I to CS-IV, 11 m thick for CS-V) that consist of
alternating layers of loess and soil. Each of the storage pits (CS-I to CS-V)
5-m-deep loess layer (Table S1).
1 sample (BWG) from the Culture Museum of Cishan were analyzed. Of these,
samples were prepared according to the procedure slightly modified from
Piperno (35, 36) and Lu et al. (37). It consists of sodium pyrophosphate
(Na4P2O7) deflocculation, treatment with 30% hydrogen peroxide (H2O2) and
cold 15% hydrochloric acid (HCl), zinc bromide (ZnBr2; density, 2.35 g/cm3)
heavy liquid separation, and mounting on a slide with Canada balsam. Phy-
with phase-contrast and microscopic interferometer at 400? magnification.
Millet Molecule Extractions from Modern Plants and Archaeological Material.
We examined modern millet molecules from 6 domesticated samples of
Paniceae, including 3 samples of Setaria italica L. Beauv. (foxtail millet) and 3
samples of Panicum miliaceum L. (common millet). Two archaeological sam-
ples of grain crop remains, including 1 sample from CS-V storage pit and 1
sample (BWG) in a storage bottle from the Culture Museum of Cishan, were
One gram of each of dried sample (smaller than 100 mesh) was ultrasoni-
cally extracted for 3 times with a mixture of acetone and pentane at 1:1
(vol/vol). After filtration, the mixture was eluted with n-hexane to give
aliphatic hydrocarbons (N). Using dichloromethane in n-hexane passing
through the column resulted in satisfactory separation of aromatic hydrocar-
chromatography coupled with mass spectrometry (GC/MS) .
GC-MS was conducted by using an Agilent 6890 N GC-5973 N MSD mass
a fused silica chemically bonded capillary column (J&W DB-5; 0.25 mm in
diameter, 30 m long, 0.25 ?m film thickness). Each sample was injected onto
the column at 280 °C in the splitless mode. After a 1-min isothermal hold at
50 °C,thecolumntemperaturewasincreasedby30 °C/minto120 °C;andthen
3 °C/min to 290 °C, successively, with a 30-min isothermal hold at 290 °C. The
flow rate of the helium carrier gas was 1.2 mL/min (40 cm/sec).
Compounds were identified by their retention time within the gas chro-
matograph, their fragmentation pattern within the mass spectrometer, and
libraries (NIST02L). The miliacin relative abundances were estimated by mea-
suring the area of the miliacin peak on the m/z 189 ? 204 ? 218 ion-specific
chromatogram. Note that this value may vary under different GC/MS condi-
tions, and there will be a small change in the abundance of this ion.
ACKNOWLEDGMENTS. We are grateful to Dolores R. Piperno for critically
reading the original manuscript and her helpful comments in improving this
manuscript. We also greatly appreciate the valuable comments from two
anonymous reviewers. We are grateful to Jacob Je ´re ´my and A. Gerasimenko
for their help on biomolecular analyses, and D. Q. Fuller and L. Qin for their
discussions on phytolith analyses. We thank C. D. Shen, Z. Y. Gu, and B. Xu for
their help in radiocarbon age measurement. This work was supported by the
Chinese Academy of Sciences (100 Talents Program, kzcx2-yw-117), the Na-
tional Natural Science Foundation of China (40771216; 40325002), Chinese
Civilization Origin projects (2006BAK21B02), and the U.S. National Science
Foundation (BCS-0623514, ATM-0402475).
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