Archaeol Anthropol Sci (2010) 2:115–131
DOI 10.1007/s12520-010-0035-y
ORIGINAL PAPER
Consilience of genetics and archaeobotany in the entangled
history of rice
Dorian Q. Fuller & Yo-Ichiro Sato & Cristina Castillo & Ling Qin & Alison R. Weisskopf &
Eleanor J. Kingwell-Banham & Jixiang Song & Sung-Mo Ahn & Jacob van Etten
Received: 30 March 2010 / Accepted: 18 May 2010 / Published online: 18 June 2010
# Springer-Verlag 2010
Abstract Major leaps forward in understanding rice both
in genetics and archaeology have taken place in the past
decade or so—with the publication of full draft genomes for
indica and japonica rice, on the one hand, and with the
spread of systematic flotation and increased recovery of
archaeological spikelet bases and other rice remains on
early sites in China, India and Southeast Asia. This paper
will sketch a framework that coherently integrates the
evidence from these burgeoning fields. This framework
implies a reticulate framework in the phylogeny of early
cultivated rice, with multiple starts of cultivation (two is
perhaps not enough) but with the key consolidations of
adaptations that must have been spread through hybrid-
isation and therefore long-distance cultural contacts.
Electronic supplementary material The online version of this article
(doi:10.1007/s12520-010-0035-y) contains supplementary material,
which is available to authorized users.
D. Q. Fuller (*) : C. Castillo : A. R. Weisskopf :
E. J. Kingwell-Banham : J. Song
Institute of Archaeology, University College London,
London, UK
e-mail: d.fuller@ucl.ac.uk
Y.-I. Sato
Research Institute for Humanity and Nature,
Kyoto, Japan
L. Qin
School of Archaeology and Museology, Peking University,
Beijing, China
S.-M. Ahn
Wonkwang University,
Iksan, South Korea
J. van Etten
IE University,
Segovia/Madrid, Spain
Archaeobotanical evidence allows us to document the
gradual evolutionary process of domestication through rice
spikelet bases and grain size change. Separate trends in
grain size change can be identified in India and China. The
earliest centre of rice domestication was in the Yangtze
basin of China, but a largely separate trajectory into rice
cultivation can be traced in the Ganges plains of India.
Intriguingly, contact-induced hybridisation is indicated for the
early development of indica in northern India, ca. 2000 BC.
An updated synthesis of the interwoven patterns of the
spread of various rice varieties throughout Asia and to
Madagascar can be suggested in which rice reached most of
its historical range of important cultivation by the Iron Age.
Keywords Archaeology . Oryza sativa . Domestication .
Dispersal . Neolithic
The distribution of wild rice and genetic diversity
in domesticated rices
Rice is a highly diversified crop, being grown from the
equator to over 40° N, from sea-level ca. 2,700 m in parts
of the Himalayas and in a wide ecological range of
cultivation systems. Although there is much less ecological
variation found within its wild progenitor complex (Oryza
rufipogon and Oryza nivara), these are nonetheless distrib-
uted over a wide geographical range and a spectrum of
ecological niches from permanent to seasonal wetlands. As
the origins of cultivation must have developed in places
where hunter–gatherers were utilising wild populations, the
distribution of the wild progenitor, in the past when
cultivation began, is a key element in identifying the
origins of rice. There are three lines of approach to inferring
this without archaeological evidence, including (1) the
116
modern geographical and ecological distribution of wild
rices, (2) an assessment of the likely effect of past climatic
changes on this distribution, and (3) phylogenetic relation-
ships between various modern wild populations and
domesticated populations. There are complicating factors,
however. Apparent genetic relationships between wild and
cultivated populations may be affected by recent introgres-
sion and hybridisation across lineages, as well as by deeper
shared histories. Thus, a careful comparison of comple-
mentary and contrasting genetic datasets is necessary. As
for geography, the modern distribution of wild rice may
have been impacted not just by climatic change but also
human land-use as suitable habitats for wild rices were
destroyed or modified, for example by becoming cultivated
landscapes. This can be accounted for to some extent by
reference to ancient Chinese historical sources which
indicate the presence of wild rices about 1,000 years ago
in regions where rice does not occur wild today.
The accepted wild progenitor complex for rice is that of
O. rufipogon sensu stricto, predominantly perennial, and
the annual O. nivara (Vaughan et al. 2005, 2008a, b), which
are distributed in South and Southeast Asia and parts of
China (Fig. 1). An understanding of the past distribution of
wild rice must also take into account probable reports from
Tang and Song Dynasty sources (eighth to tenth century
AD; Ho 1977; You 1987). We follow You (1987), in
separating those which are from plausible primary habitats,
such as lakes as opposed to those “wild” rices that sound
Fig. 1 Map of modern and historical distribution of rice’s wild progenitors, O. rufipogon and O. nivara
Archaeol Anthropol Sci (2010) 2:115–131
more like weedy types (“Oryza spontanea”) or late tillers
collected from cultivated areas in times of famine.
During periods of drier climate, such as Pleistocene glacial
periods, we expect that these wild populations would have
been more restricted into wetter tropical refugia, in parts of
Eastern India, Yunnan and mainland Southeast Asia (e.g.
Fig. 2 P). This is consistent with the genetic evidence that the
ancestral haplotypes of O. rufipogon are found in Southeast
Asia and India (Londo et al. 2006). The most recent
geographic expansion and radiation of wild rice populations
would have started ca. 15000 BP (16500 cal. BC) as post-
glacial climatic conditions set in. In addition to the changes
in temperature and rainfall regimes, it should be noted that
this period also saw a dramatic increase in atmospheric CO2
levels from 18 Pa (atmospheric partial pressure) to 27 Pa, by
11000 BP, which is expected to have had a major impact on
the photosynthetic productivity and biomass production of
plants, but especially C3 plants like rice (Sage 1995;
Cunniffe et al. 2008). We expect this period to have
witnessed a major expansion in the geographic range of rice
towards higher latitudes, especially in East Asia where a
northwards expansion was not blocked by the high Hima-
layas. We might also expect some adaptive changes in rice
morphology in response to new climatic and atmospheric
conditions and will return below to how this might confound
the recognition of some domestication traits.
A massive geographical range expansion of wild Oryza
spp. took place between the dry conditions of the glacial
Archaeol Anthropol Sci (2010) 2:115–131
Fig. 2 Map contrasting the expected Pleistocene (P) glacial refugium
distribution of wild rice, based on tropical vegetation zones of Adams
and Faure (1997) and phytolith data from southern Poyang Lake basin
(Zhao and Piperno 2000) and the Holocene (H) maximum rice
expansion associated with conditions of the Early and Middle
Holocene, based on agroclimatic zones of Huke (1982) adjusted for
maximum, the warm and wet conditions of the Terminal
Pleistocene (Fig. 2 P) and early Holocene (Fig. 2 H). Dry
seasons of 7 months or more are likely to preclude
cultivation of traditional late ripening rices and wild rices.
As indicated by Chinese historical sources, the major
adoption of early-ripening rice (which can be grown in 4 to
5 months), generally more tolerant of water shortage, took
place from the eleventh century AD onwards (Ho 1956). For
this reason, the climatic pattern devised by Huke (1982), tied
to unchanging topography, seems a suitable corrective to the
reconstructed vegetation zones of pollen indicators (Adams
and Faure 1997; also, Shi et al. 1993; Ren and Beug 2002).
This indicates that wild rice probably grew throughout the
Yangtze basin and eastern China as far north as Shandong,
and thus, archaeobotanical finds of rice, without clear
domestication traits or indicators of cultivation, could relate
to wild gathering. The potential of inferring cultivation
through archaeological finds of rice beyond the range of the
wild progenitors is thus quite limited, although finds from
northwest Pakistan from 2500 to 2000 BC qualify (Costantini
1987), as does Early Yangshao rice from Nanjiaokou,
northern Henan (Qin and Fuller 2009) and Late Yangshao
rice from Xishanping, Gansu (Li et al. 2007).
Rice is divided into two main sub-species, indica and
japonica (e.g. Oka 1988; Vaughan et al. 2008b), although
117
early Holocene conditions but taking into account topographic
constraints like the Qinling Mountains, as well as vegetation
reconstructions of Adams and Faure (1997) and Yu et al. (2000),
with adjustments in India for more precise vegetation zones (cf.
Asouti and Fuller 2008)
numerous other varieties and sub-varieties are recognised.
Recent genetic classifications (e.g. Garris et al. 2005;
Caicedo et al. 2007; McNally et al. 2009) suggest that
there are five analytically useful groups which can be
recognised: indica, aus, temperate japonica, tropical
japonica (including javanica) and fragrant, although most
discussions of origins have focussed on whether the indica
(including aus) and japonica groups derived from a single
or multiple domestications. There is a now a wide range of
genetic studies, using markers from the chloroplast genome,
neutral variation in the nuclear genome and variation within
functional alleles that have been used to examine this issue.
Chloroplast DNA has suggested two distinct lineages in
both wild and domesticated rice since a 69-bp deletion was
reported in the early 1990s (Chen et al. 1993). Nakamura et
al. (1998) discovered an additional cpDNA polymorphism
in a single sequence repeat (SSR) in the PS-ID region.
Differences with respect to the number of C (cytosine) and
A (adenine) bases arrayed in succession enable not only
differentiation of indica (8C8A) and japonica (6C7A) but
also identify variation amongst japonica (some are 7C6A)
and differentiation of wild rice species (several additional
variants; see also Ishikawa et al. 2002; Takahashi et al.
2008). In addition, chloroplast genome types (both in PS-
ID and 57 K sequence genome types) typical of indica are
118 Archaeol Anthropol Sci (2010) 2:115–131

not represented in modern wild rice samples, suggesting
that the immediate maternal ancestor of indica has either
not been sampled or has gone extinct (Kawakami et al.
2007; Takahashi et al. 2008). More recently, a comparison
of whole chloroplast genomes for single representative
japonica and indica varieties showed a high degree of
divergence with an age estimate for their common ancestor
at 86,000–200,000 years (Tang et al. 2004).
A wide range of markers in the nuclear genome show
similar patterns. Whole genome comparisons of represen-
tative accessions for the nuclear genome suggest a similarly
deep divergence on the orders of 100,000 s of years (Vitte
et al. 2004; Ma and Bennetzen 2004), whilst various genes
or gene systems also point to differences such as pSINEs
(Ohtsubo et al. 2004), SSR and STS datasets (Garris et al.
2005; Caicedo et al. 2007), SAM and vTPASE sequence
variation (Londo et al. 2006) and ∼3,000 single nucleotide
polymorphisms assayed across 20 representative landraces
(McNally et al. 2009). Whilst all of these studies agree on
the deep divergence of indica and japonica, they also
suggest that there are some indicators of possible multiple
derivations in both indica and japonica. These studies also
show hybridization between these lineages and most
recently McNally et al. (2009) have provided evidence for
quite frequent hybridisation between the major cultivar
lineages. Much of this hybridisation may be historically
recent, for example taking place in China over the past
800–900 years after early-ripening indica rices were
introduced (cf. Ho 1956), or earlier around (1,500–
2,500 years ago) in India and Persia when early historic
trade connections were well established—the earliest
archaeological rice in the Near East is Parthian (first
century AD; Miller 1981), whilst imported rice is known
from Berenike on the red sea coat of Egypt (ca. 50 BC;
Cappers 2006). Nevertheless, some hybridisation between
early japonica and indica lineages may have taken place
near the start of domestication, even when one lineage was
under primitive pre-domestication cultivation (Fig. 3).
Indeed current evidence for morphological domestication
traits and what is known of associated genes implies as
much (Sang and Ge 2007; Fuller and Qin 2009).
The domestication traits of rice: archaeobotany
and genetic lineages
Domestication refers to the key morphological (and
genetic) adaptations that characterise crops adapted to
cultivation, traits that have often been characterised as
an adaptive ‘syndrome’ of domestication (e.g. Harlan et
al. 1973; Hammer 1984; Fuller 2007; Purugganan and
Fuller 2009; Fuller and Allaby 2010). These domestication
traits evolved in a more or less gradual and piecemeal
process in response to the selection pressures created by
the human activities of cultivation. In other words,
tracking the origins of rice requires us to try to identify
the behaviour of cultivation, then the appearance and rise
to dominance of the domestication traits and finally the
spread of reliance on rice agriculture. New appreciation of
the drawn-out process of the evolution of domestication
traits raises the likelihood that traits were selected
separately in different regions and then brought together
by processes of hybridisation/introgression (see Allaby
2010).

Fig. 3 A simplified diagram of

the main evolutionary pathways
to domesticated Asian rice, with
selected key mutations
indicated, differentiating
wild-type dominant form with
all capital letters and
domesticated type recessive
alleles with lowercase letters.
Mutations: sh4 non-shattering,
Prog1 erect growth, rc white
grain pericarp, sw5 wider grains,
qsh1 further non-shattering, wxy
waxy/glutinous rice (low
amylase), WW wild genepool,
dd domesticated genepool, cult.
cultivated, X major hybridisation
event; crossed arrows indicate
continued gene flow
(introgression)
Archaeol Anthropol Sci (2010) 2:115–131
As with other cereals, non-shattering is a key
domestication trait which is likely to be deleterious in
wild habitats. Geneticists identified six possible quanti-
tative trait loci that appeared to be involved in this trait,
with four that recur across studies (Xiong et al. 1999;
Cai and Morishima 2002; Li et al. 2006a; Lin et al. 2007).
Two of these have subsequently received much attention
prompting sequencing research. These are the recessive
mutation sh4, which is a SNP in a gene on chromosome 4
shared widely across japonica, fragrant, aus and indica
rice (Li et al. 2006b, 2007) and qsh1 which is found only
in some temperate japonica rices (Konishi et al. 2006) and
appears to be derived from a haplotype found in some O.
rufipogon. Most commentators have assumed that the
widespread sh4 was the major mutation involved in initial
rice domestication (e.g. Sang and Ge 2007; Vaughan et al.
2008a; Fuller et al. 2009). Fuller and Qin (2009) argued that
this gene was introduced to proto-indica cultivars in India
after japonica domestication in China, although recently
Zhang et al. (2009a, b) have suggested that this evolved
rapidly after domestication and then became widespread in
cultivated rices through gene flow (hybridisation; Zhang
et al. 2009) therefore inferring that sh4 was not involved
in the initial evolution of non-shattering detected archae-
ologically (e.g. at Tianluoshan). Later in a sub-group of
East Asian rices, qsh1 became fixed during or after the
bottleneck that led to differentiation of temperate japonica
(Zhang et al. 2009). Unfortunately, little is known about
morphological differentiation of the spikelet bases of these
genotypes when harvested and threshed by various
Fig. 4 A compilation of quantified spikelet base data from sites in
China and Thailand, showing the long-term trends to decreasing wild
and increasing domesticated fractions. Sites and samples sizes
indicated. Total sample=3,768 specimens. Data for Tianluoshan and
Liangzhu and separation criteria follow Fuller et al. (2009); data for
Kuahuqiao and Luojiajiao indicate only the wild fraction as immature
119
traditional means, so it is not yet possible to link
archaeological remains to genotypes.
Archaeobotanists are interested in spikelet bases as clear
indicators of whether individual rice specimens are of
domesticated or wild type (e.g. Sato et al. 1991; Thompson
1996, 1997; Sato 2002). There has been some confusion in
the use of this morphological indicator. The criteria
proposed by Zheng et al. (2007), in which there are two
categories, are not entirely consistent with those of
Thompson (1996, 1997), and both present different ‘do-
mestic’ morphotypes. Zheng et al. (2007) examined four
modern populations, two wild and two domesticated;
Thompson examined a few more modern populations.
Based on a more extensive comparative study of wild
material than either of them (159 populations) and taking
into account the likely presence of immature spikelets,
which is indicated in grain morphology as well as empty
waterlogged husks on some early Chinese sites, we now
have a three category division (in addition to a fourth
indeterminate category for poorly preserved specimens;
Fuller and Qin 2008; Fuller et al. 2009). However, amongst
the domestic/non-shattering category, there is still a range
of variation apparent which might relate to different non-
shattering genetic loci (or combinations of loci) as well as
to variation in the cultural practices of harvesting and
threshing tools/techniques. Further research is needed on
this topic.
Nevertheless, the recent realisation that rice spikelet
bases are usually well preserved in the finer fraction of
sieved and flotation samples has allowed us to start
and domesticated forms were not separated (after Zheng et al. 2007);
other data unpublished/currently under study by the authors: Puanqiao
(Zhejiang), Songze-Liangzhu Transition by QL; Sushui (Shanxi)
survey by JS; Chengyao, Ying Valley (Henan) by DF; Khao Sam
Kaeo by CC
120
accumulating spikelet base data in recent years from sites in
China and Thailand (Fig. 4). Rechecking the fine fraction
of previously sorted samples from Mahagara, India (Harvey
et al. 2003; Harvey 2006) has also revealed spikelet bases.
The largest assemblage yet studied is that from Tianluoshan
in the Lower Yangtze region, which is now central to
discussions of the domestication process in the Lower
Yangtze (Fuller et al. 2009; Zhao 2010; Nakamura 2010).
Unfortunately, we lack analogue population studies of the
proportions of spikelet base types found in modern
traditional rice harvests, in which weedy rices and some
immatures should occur. The timing of rice harvests usually
balances loss to incomplete grain maturation with earlier
harvests (e.g. ∼30 days after harvesting) and loss due to
shattering as panicles become ‘over-mature’ with later
harvests (e.g. ∼40 days after flowering; Horiuchi et al.
1971). Some modern agronomic data from China would
suggest that levels of 5–10% immature spikelets might be
expected, affected in part by factors like water and disease
in a given season (cf. Pan 2008), but as indicated by
ethnographic and experimental data in Malaysia (Horiuchi
et al. 1971), these are also affected by harvest time. In
addition, we know little at present about how differing
harvest and processing methods might bias spikelet base
assemblages that enter routine domestic refuse or seasonal
threshing waste.
The other domestication trait often considered in
archaeobotany is grain size (Fig. 5), although any use of
this as an absolute indicator of domestication status on
individual specimens is problematic. It is generally true that
domesticated grain crops have larger (often wider) seeds
than their wild ancestors, which is thought to be selected for
by deeper burial in more disturbed, tilled soils (Harlan et al.
1973; Heiser 1988; Fuller 2007). In rice, however, this trait
is complicated by the great range of variation represented
by modern cultivars and wild populations (Fig. 5a). Also,
much variation in rice grain length can be linked to climatic
variation and ecological adaptations (Oka 1988; Kitano et
al. 1993): More northerly temperate japonica landraces are
short-grained, whilst tropical varieties (the javanica race
rices) are massively long. Complicating this trend is that of
longer-grained upland rices versus shorter-grained lowland
rices in East Asia generally (Nitsuma 1993), whilst some
varieties at high elevations in the tropical mountains (e.g.
Yunnan, Nepal) may be very short-grained. The latitudinal
trend found in modern rices is indeed evident in the data of
Neolithic northern China (e.g. Liu et al. 2007; Qin and
Fuller 2009) and Bronze Age Korea (Ahn 2010), where
grains are generally shorter than those from the Yangtze of
similar date. Seed size is affected by many factors,
including environment, and is polygenically controlled
(Fuller and Allaby 2010). For the most part, these genes
and regulatory networks are poorly understood, but the
Archaeol Anthropol Sci (2010) 2:115–131
many QTL analyses that have been carried out have shown
seed size to be associated with many genes of varying
effect (Gupta et al. 2006). At least three loss-of-function
mutations have been identified in rice that directly influence
grain shape/size (Fan et al. 2006; Song et al. 2007;
Shomura et al. 2008). In addition, mutation to the gene
spd6 appears to increase grain size as well as grain number
(Shan et al. 2009). Two of these result in an increase in the
number of glume cells, thereby giving the grain milk a
larger cavity to fill, resulting in wider grains (Song et al.
2007; Shomura et al. 2008). This shift in husk cell
architecture is important as it offers an explanation of some
of the variation documented amongst and between the
domesticated and wild rice husk phytoliths (Zhao et al.
1998; Harvey 2006)—although there is so much variation
and overlap amongst modern rices to make this an
unreliable separator in many cases (Harvey 2006). In
addition, this implies that ‘domestic’-type phytoliths found
in pre-agricultural Pleistocene cave sites (Zhao 1998, 2010)
could be the product of shifts in climate and latitudinal
vegetation belts in the post-glacial period.
In the archaeobotanical record of India and of the
Yangtze, one finds trends towards larger grain sizes over
time (Fig. 5b, c). However, this is also not the only trend:
Grains from the Yellow River in China tend to be as small
or smaller than those in the Yangtze (Liu et al. 2007),
meaning that this trait cannot be used in a straight forward
way to indicate domestication, but local trends of change
may be considered useful indicators. In India, by contrast,
grain size increase appears to be dampened within the
Ganges core region by contrast to early grains in regions
without wild rice. Whether this indicates a role for recurrent
gene flow from wild populations or different selection
pressures in differing regional cultivation regimes (which
has been argued for pulses: Fuller and Harvey 2006)
requires further research.
Another classic domestication trait is apical dominance,
reflected in reduced lateral branching or tillering that results
in a more erect growth habit. The gene PROG1 which has
attracted particular attention in rice genetics recently
appears to have a large effect on morphology when the
recessive form is expressed (Tan et al. 2008; Jin et al.
2008). The domestic-type prog1 sequence was shared
across all 182 domestic lines tested, with a similar sequence
found in a subset of perennial O. rufipogon from South
China, Taiwan and Thailand (Tan et al. 2008). Whilst this
suggests that this trait is derived from variation that existed
in the wild, wild bearers of the mutation do not express an
erect growth habit indicating the countervailing effect of
other factor(s). Other genes that have been implicated in
rice architecture include Moc1, MAX2 and MAX3, but as
reviewed by Doust (2007), branching pattern is influenced
also by environmental conditions.
Archaeol Anthropol Sci (2010) 2:115–131 121

Fig. 5 Grain size data.

a Modern ranges based on
measured populations in UCL
collection (same as those
reported in Fuller et al. 2007)
and full range of extreme
modern (reported in Vaughan et
al. 2008a). b Grain width of
archaeological site populations
(mean and standard deviation),
plotted against time in the Lower
Yangtze, with trend line through
averages shown. c Grain width
of archaeological site populations
(mean and standard deviation),
plotted against time in northern
India, with trend line through
averages shown. For
archaeological details
on archaeological data,
see Table S1

Chronological and geographical patterns
in the archaeology of rice
Archaeobotanical evidence for the use of wild rice remains
quite limited, whilst the date of the start of cultivation of rice is
controversial in both China and India. The locations of
Pleistocene and Early Holocene reports are shown in
Fig. 6a, with site details in Table S2. A single report of
Pleistocene rice from Xom Trai, Vietnam, at ca. 16000 BP, is
presumably wild (Glover and Higham 1996: 421). Some
Pleistocene rice finds are not directly associated with human
archaeology (i.e. no associated artefacts) and might be
suggested to relate to wild populations, including phytoliths
off of the Shanghai coast (Lu et al. 2002) and perhaps the
infamous rice husks of the palaeosol (peat) deposit at Sorori
in Korea (Lee and Woo 2006; Kim et al. 2008; however, the
latter may represent later Holocene intrusive material: see
critical comments in Ahn 2010). Interestingly, both sites date
to ca. 13000 BP uncalibrated or ca. 13000 cal. BC which is
after a period of rapid global warming and CO2 increase that
should be expected to have extended the wild range of rice
northwards, perhaps even further north than in the Early
Holocene. A few finds of rice from the Late Pleistocene
archaeological occupation sites in China are presumably
122
Fig. 6 The chronological
spread of rice in two maps and
11 ‘thrusts’. Sites plotted in
broad time horizons. Arrows and
numbers refer to the numbered
thrusts referred to in the text.
For details of the sites plotted,
see Table S2
from wild use, such as Yuchanyan, Xianrendong and
Diaotonghuan (Lu 2006; Zhang and Hung 2008). Despite
the suggestion from phytoliths that they document a shift
towards domesticated morphology in husk cells (Zhao 1998,
2010), we would suggest that this might be mimicked by
rice’s natural response to rapid post-glacial climatic change.
What the shift in husk phytoliths suggests is a shift in grain
size, which is just as likely the result of adaptation to climatic
changes at ca. 15000 cal. BC, the date of the phytolith shift
based on the association with early ceramics that are now
well dated to this time (see Kuzmin 2006; Zhang and Hung
2008; Boaretto et al. 2009).
Setting aside Pleistocene finds, we are still faced with
whether Early Holocene finds of rice indicate foraging or
cultivation. In the Lower Yangtze region, the earliest evidence
for rice dates between 10000 and 8000 BC, especially at the
site of Shangshan (Jiang and Liu 2006; Nakamura 2010; Zhao
2010), but evidence documents neither clear cultivation
behaviour nor morphological domestication. Two spikelet
Archaeol Anthropol Sci (2010) 2:115–131
bases could be isolated, which include one unambiguous
wild type and one immature type (cf. Zheng et al. 2007; but
criteria follow Fuller et al. 2009), and thus, there are no
confirmed domesticated types and a uselessly small sample
size. In the middle Yangtze, ceramics with husk temper from
the site of Pengtoushan have direct AMS dates back to
8000–7000 BC, whilst the culturally related site of Bashi-
dang (7000–6000 BC) produced 1,000 s of rice grains,
within an overall broad spectrum assemblage, but without
any good evidence for domestication traits one way or the
other (e.g. HPIACR 2006). It should be noted that the
conventional view assumed that the earliest rice in China
was domesticated indica (e.g. at Hemudu, Bashidang) or
‘intermediate’ between indica and japonica and evolved into
japonica over the course of the Neolithic (e.g. Zhang and
Wang 1998; Zhang 2002). Such a model is contrary to what
would be expected based on current genetic knowledge (e.g.
Sato 2002; Sweeney and McCouch 2007), and morphomet-
ric data on leaf bulliform phytoliths indicate that these are
Archaeol Anthropol Sci (2010) 2:115–131
consistently of the japonica type (or its wild ancestor; e.g.
Sato et al. 1991; Fujiwara 1993; Zheng et al. 2003).
The situation is similar in India, where well-dated early
finds come only from Lahuradewa in the middle Ganges
region, which has ceramics and charred rice remains dating
back to 6000–8000 BC (Tewari et al. 2008). Although the
latest report claims to have ‘domesticated’ rice on the basis
of grain L/W/Th ratios and spikelet bases, the reported
measurements and illustrated spikelet bases actually con-
tradict these claims, with the grain ratios, and absolute
measurements clearly overlapping with O. nivara and O.
rufipogon and one population with Oryza officinalis (for
details, see Fuller 2009). Archaeological grain metrics for
the region suggest that a marked increase in rice grain
breadth occurred after 2000 BC continuing through the first
millennium BC (Fig. 5c). Rice bulliforms from the nearby
Lahuradewa Lake pollen core have been suggested to
include domesticated types, although the original method-
ological study for the criteria used—the number of fan-edge
chips (Lu et al. 2002)—would imply domesticated rices
already introduced into the Lower Yangtze by 13000 BP,
which is so at odds with the archaeobotanical evidence
(from habitation sites, spikelet bases etc.) that this criterion
must be regarded with suspicion. Rather, as already
suggested, the Lower Yangtze sea core phytoliths suggest
the colonisation of the region by wild rice with post-glacial
warming, and this thus undermines fan-edge bulliform
architecture as a domestication trait.
Based on unambiguous morphological domestication
traits, we can recognise the presence of domesticated rice
from 6000 cal. BC in the Lower Yangtze and from ca.
2000 cal. BC in the Ganges. In the Lower Yangtze,
Kuahuqiao (6000–5400 BC) is reported to have a majority
of mature wild-type spikelet bases (Zheng et al. 2007),
whilst insufficient criteria make it unclear how the rest
divide between immature and domesticated types. At
Tianluoshan, it is clearer that a large minority of domesti-
cated types in the earliest samples (ca. 4900 BC) increase
over the subsequent three centuries to outnumber mature
wild types (39:31%), whilst immature types also decrease
(Fuller et al. 2009). By the time of the proto-urban
civilisation of Liangzhu, this ratio had tipped to 68:25 in
favour of domesticates. It is probable that 20–25% wild
types can be derived from weedy forms of rice that have
persisted alongside domesticates. Indeed, evidence from
Late Yangshao Shanxi (Sushui valley survey) and the Ying
Valley (Chengyao) show similar proportions of wild types
and similarly reduced proportions of immature types (5–
10%). In the Ganges, data are only just starting to be
accumulated but so far the few dozen spikelet bases
recovered from Mahagara are almost entirely domesticated,
as are those from early historic sites in the South like Terr
and Paithan. In the latter cases, this may indicate that the
123
cultivation regime (dry cropping during the monsoon?) was
one where weedy rices are less of a problem. By contrast in
Iron Age southern Thailand, represented by Khao Sam
Kaeo, associated with an upland dry-rice weed flora, a high
proportion of wild-type spikelet bases suggest a high
presence of weedy types and probable continuing intro-
gression. Nevertheless, domesticated morphotypes were
present in Neolithic Thailand, by ca. 2000 BC, where 27
spikelet bases recovered from coprolites at Khok Phanom
Di are said to be dominated by domesticated types
(Thompson 1996).
Elsewhere, our record of early rice is limited merely to
data on presence and sometimes quantitative importance in
comparison to other crops. Nevertheless, we can sketch the
broad patterns of the dispersal of rice cultivation. Dispersals
can be divided into 11 distinct thrusts, which are reviewed
below. These thrusts are numbered below and on the arrows
in Figs. 6a, b. The evidential basis of these 11 regional/
chronological units is summarised through a list of sites in
the on-line supplementary material (Table S2).
The spread of rice in 11 thrusts
1. The northern thrust and early temperate japonica
Rice is found in the middle Yellow River, separated by the
Qinling Mountains from the wild rices of the Yangtze
watershed, from about 6,000 years ago, and much more
extensively by 5,000 years ago. For Yangshao or Dawenkou
sites, 16 have apparently reliable reports of rice, whilst for the
Longshan period (2500–1900 BC, including Miaodigou 2),
this tally doubles to 33 (Fig. 6a). Recent systematic sampling
by flotation during excavations and regional surveys has
been extensive and accounts for the majority of this
evidence, including numerous direct AMS dates on rice or
other crop grains. Typically, rice is a minor component
alongside millets. Setaria italica tends to dominate the
archaeobotanical evidence whereas second place is taken
by Panicum miliaceum or rice. This is true of the earliest
finds from Nanjiaokou from Early Yangshao cultural levels
(from 4500 BC, with direct AMS on rice of ca. 3800 BC;
Qin and Fuller 2009). In the subsequent Middle Yangshao
period (3800–3000 BC), Nanjiaokou and several other sites
have produced rice, and it is fairly widespread by 3000 BC.
Geoarchaeological and phytolith evidence from the Yiluo
Valley suggest that rice agriculture in this region was carried
out in wetland fields and was facilitated by a period of active
river alluviation and hillslope erosion, which created suitable
low-lying environments for irrigated rice (Rosen 2008).
Available grain metrics indicate notably short-grained forms
of rice, consistent with a temperate japonica lineage. After a
peak of rice farming (by number of sites) in the Longshan
124
period, rice finds decrease into the Bronze Age with the rise
of the Shang state, which may relate to the climatic
aridification after 2200–2000 BC. Those varieties which
persisted must have been adapted to more northerly
conditions. This reduction in rice land should also be seen
in relation to the introduction of bread wheat and in some
areas barley, at about this time from Central Asia (Li et al.
2007; Lee et al. 2007; Fuller and Zhang 2007). After this
period, wheat and rice took their place alongside the millets
and soybean as the five grains of classical Chinese
civilisation (Chang 1980).
2. An earlier Jomon and Chulmun upland japonica?
There is a hypothesis that rice cultivation was introduced
to Japan already during the late Jomon period prior to
2000 BC and certainly by 1000 BC (e.g. Sato 1996;
D'Andrea 1999). Evidence for this remains limited and
mainly comes from phytoliths from Jomon site sediments
and ceramic fabrics with rice inclusions from Kyushu (e.g.
Fujio 2004; Takahashi 2009; Toyama 2002: Table 2).
Whilst the presence of rice in pre-Yayoi cultural contexts
seems clear, for sites in Kyushu (e.g. Uenoihara, Ooishi,
Fukuda, Minami Mizote), some commentators put the date
of these finds only marginally earlier than 1000 BC (e.g.
Keally 2004). The earliest direct AMS dates on grains come
from terminal Jomon Kazahari far to the north in Aomori
prefecture, with dates of ca. 900–700 BC. The presence of
rice in this era may also be indicated in South Korea,
although data are limited and ambiguous (Ahn 2010). The
database for South Korea is quite extensive (56 sites) by
comparison to its land area, but the vast majority of
evidence dates to after 1000 BC, and all earlier finds are
rather controversial. Unambiguous rice finds date to after
2000 BC—based on the earliest direct rice AMS date at
Oun 1, although the wide error margin could be consistent
with 1500–1450 BC (Crawford and Lee 2003). This early
rice may have been grown in dry field systems, whilst the
later Mumun is generally associated with paddy field
agriculture (Ahn 2010), like the Yayoi in Japan (Imamura
1996; Kuraku 2002; Takahashi 2009).
This Jomon rice has been interpreted as a dry field crop,
differentiated from the Yayoi paddy field rice, because the
Jomon sites were located in upland areas. Whether this
implies photo-period insensitive, early-ripening varieties is
not clear. It has been suggested that these rices would have
represented an early and primitive form of tropical japonica
cultivars, distinct from the temperate japonica paddy rice
introduced in the Yayoi period (Sato 1996; cf. Takamiya
2001: 216). Ancient DNA analyses of Yayoi and later rice
remains suggest a mixture of temperate and tropical
japonica types which would be consistent with an
introduced Yayoi temperate paddy rice over a substratum
of earlier tropical japonica (Sato 1996; Tanaka et al. 2010).
Archaeol Anthropol Sci (2010) 2:115–131
If dates for rice in Japan prove to be truly earlier than those
in Korea, then diffusion directly from the Lower Yangtze/
Shandong coast area would seem to be indicated. However,
uncertainties over Chulmun rice reports in Korea (see Ahn
2010) mean that there may have been an earlier, dry-
cropped, form of rice cultivation that spread from the
Shandong peninsula to Korea and Japan earlier than the
more obvious paddy field diffusion wave of the Mumun
and Yayoi period. Archaeological evidence for upland rice
in China at this period is so far limited, although the weed
flora of Liangchenzheng in Shandong (2500–1800 BC)
could be consistent with rain-fed cultivation (Crawford et
al. 2005). In addition to systematic archaeobotany and
direct dating in Korea and Japan from this era, further work
aimed at understanding the ecology of rice, e.g. through
weed flora, is needed in China.
3. To Taiwan and beyond: a southerly quasi-temperate
japonica
Rice appears in Taiwan along with foxtail millet in the early
third millennium BC (Tsang 2005), presumably with the
spread of agriculture from mainland China. Hard data are
limited, with five Neolithic sites having reported rice and
none of them with a full, systematic archaeobotanical study.
The interesting co-occurrence of rice and millet together with
cultural practices, like front tooth removal, and apparently
shared early vocabulary may suggest a derivation from the
southern Shandong coast/northern Jiangsu (Sagart 2008)
rather than the Lower Yangtze area (postulated by Bellwood
1997, 2005), as millet and rice are unknown together in
Lower Yangtze archaeobotany. Although quantitative data
are unavailable, Tsang’s (2005) photograph suggests a short-
grained rice morphotype, like early grains from Shandong
and the Yellow River basin.
From Taiwan, rice is expected to have spread further
south into island Southeast Asia, although hard evidence is
scarce (Paz 2003; Barton and Paz 2007), with one Neolithic
site in the Philippines, three in Borneo, two in Bali and one
in Sulawesi. Half of these are based on husk impressions in
pottery only. Though some of these may be as early as
2000 BC, there are few morphological details available.
This may indicate that the Austronesian-associated ceramic
Neolithic of the islands although linked with some rice
agriculture in its early stages was mainly driven by other
forms of subsistence (Bellwood 1997).
4. Proto-indica origins and spread
Although rice use is established by around 7500 BC at
the site of Lahuradewa (Tewari et al. 2006, 2008; Saxena et
al. 2006), whether this rice was cultivated or gathered wild
remains controversial (see reviews in Fuller 2006; Fuller
and Qin 2009). Later evidence from around 2500–2000 BC
is stronger which suggests that rice was an established
Archaeol Anthropol Sci (2010) 2:115–131
cultivar and staple food and for some sites the only crop in
this region. On this basis, archaeologists working in India
contend that rice was separately brought into cultivation in
the Ganges region, although good evidence for the process
of morphological domestication is unclear. Spikelet base
remains have not been reported from India, although recent
work by one of us (DF) has recovered domesticated
spikelet bases from Mahagara from between 1800 and
1600 BC. If it is assumed that domestication was controlled
by the sh4 non-shattering mutation, then this evidence
provides the minimum age by which the non-shattering trait
was introduced into ancestral indica. Alternatively, we
could suggest that a different locus was involved in early
non-shattering in indica and sh4 was introduced later but
became universal as it was in some ways more effective.
What is clear from the evidence at present is the end of the
process of agricultural origins in this region, as rice/other
cereal/pulse-cultivating sedentary villages with domestic
livestock become widely established from 2000 BC (Fuller
2006). Crop, livestock and ceramic forms point towards links
to the upper Indus/Ravi River (Harappan) region to the west.
This may indicate that an intensive and expansive agricul-
tural economy focussed on rice did not develop prior to the
input of other crops, including winter crops adopted from the
West. One reason for this may be that domesticated japonica
rice was needed to introduce some key domestication alleles,
including prog1 (erect growth), sh4 (non-shattering) and rc
(white pericarp; Kovach et al. 2007; Jin et al. 2008; Tan et al.
2008). As indicated in Fig. 4c, grain size increase (thicken-
ing) is indicated only after 2000 BC through the Chalcolithic
and Iron Age periods.
5. Tropical japonica from South China to Southeast Asia
Despite a tally of 67 sites/phases in mainland Southeast
Asia with reported rice (from Neolithic through Iron Age
times), 66% consist only of the casual report of rice-
tempered pottery, with only 17 sites in Thailand having had
systematic sampling. Southern China, including Guangxi,
Guangdong and Hong Kong, is still data poor (Lu 2009):
Six sites plus two more from southern Hunan have reports
of rice, with little published in any detail (Fig. 6b). This
means that we can sketch some broad chronological
patterns (see Rispoli 2008; Zhang and Hung 2008), but
only for a few sites in Thailand can we attempt to say
something more about the local significance of rice in the
diet, or cropping ecology. Broadly speaking, in mainland
Southeast Asia, rice appears by 2000 BC, especially
associated with the lower reaches of the Red River
(Vietnam), the Mekong River (Cambodia) and Chao Phraya
(Thailand). Interior finds up river are at present consistently
later (perhaps mainly from 1500 BC, see e.g. Higham and
Higham 2009; cf. Vincent 2003). This argues against the
Higham hypothesis (e.g. Higham 2003) of a dispersal down
125
the spoke-like rivers from Yunnan, but rather a more
coastally focussed, or perhaps on the lower mountain slope
zones, zonal dispersal from southern China (Guangdong).
Recent assessment of ceramic complexes by Rispoli (2008)
similarly points to links from the middle Yangtze to
Guangdong and Guangxi (see also Lu 2009, Zhang and
Hung 2010) and onwards to Vietnam and mainland
Southeast Asia. This revised view would fit with a process
that saw the development of true tropical japonicas of the
lowlands and lower slopes, with an ecology based on
rainfall or natural flooding (as suggested by White 1995).
Modern genetics suggests that this trajectory saw the early
development of true glutinous (waxy; cf. Olsen and
Purugganan 2002) and fragrant (badh2.1) varieties of rice
(cf. Kovach et al. 2009). Also of note is the evidence that
foxtail millet was a major crop within this trajectory dating
to the Neolithic and Bronze Age in mainland Southeast
Asia (Weber et al. 2010), tracing its roots to northern China
(Lee et al. 2007), via the middle Yangtze, where it is
reported from Chengtoushan at ca. 4500 BC (Nasu et al.
2007), via Gantouyan near the Guangxi–Vietnam border
with both rice and foxtail millet by 3000–2000 BC (Lu
2009). Economically and ecologically, rice may not have
been any more significant than foxtail millet, but diversi-
fication of its cooking qualities suggests that it had
important cultural or aesthetic significance.
6. Carry on up the Yangtze
Rice cultivation also presumably spread up the Yangtze
River into the Sichuan basin and to Yunnan (Fig. 6b). Rice
is known in the Chegdu plain from Jinsha during the
Baodun period, 2500–1800 BC (Zhao, Zhijun, personal
communication.) In Yunnan, rice is reported from Neolithic
sites that date to the third millennium BC, such as at
Baiyangcun, ca. 2100 BC (An 1999). However, in both
provinces, archaeobotanical sampling has been rare. It is
also worth noting that, in both areas, millet(s) occur as well
as rice and the evidence from Karuo in the mountains of
Qamdo, eastern Tibet puts foxtail millet there by 3000 BC
(An 1999) and might indicate that foxtail millet cultivation
in mountain areas was established centuries to a millennium
earlier than rice cultivation.
7. The introduction of japonica genes and the making
of indica
Recent genetic evidence has confirmed the model of Sato
(1996) that indica is of hybrid origin including nuclear genes
from domesticated japonica, although alternative scenarios
may be conceived for the initial meeting of proto-indica,
either as wild populations or as a cultivar under pre-
domestication cultivation (Sang and Ge 2007; Vaughan et
al. 2008b; Zhang et al. 2009; Fuller and Qin 2009). Whilst
direct dispersal from the upper Yangtze via Assam is
126
proposed by some and would fit with hypotheses that derive
rice cultivation from the dispersal of farmers in the
Austroasiatic language family (e.g. Higham 2003; Bellwood
2005), current archaeobotanical evidence for rice does not
provide support for the model. As noted above, rice use, if
not cultivation, has a millennia long history in the Ganges,
and rice, of unclear domestication status, seems to become
widespread in the upper Ganges and upper Indus region by
2500 BC. It is only at this same time horizon that rice
cultivation appears to have spread to the upper Yangtze in
both Sichuan and Yunnan, leaving a large geographical gap
to be covered in no time at all. In addition, historical
linguistic data from Indian Austroasiatic (Munda) languages
and archaeological evidence from eastern India point towards
an even later arrival (ca. 2000 BC), after the dispersal of
Indian pulse and western animal domesticates (Fuller 2003).
On the other hand, evidence for several cultivars
originating in China occurs widely in Late Harappan times
in northwest India and Pakistan (Fuller 2006: 36; Fuller and
Qin 2009). The earliest plausible japonica rice grains with
short and wide grains is reported from the Swat valley and
Baluchistan (see Costantini 1979, 1987), from ca. 1900 BC.
In addition, there is a list of other crops of Chinese origin
that occur for the first time in South Asia in the northwest,
including P. miliaceum (see Jarrige 1985; Fuller and Boivin
2009), apricots, peaches and Cannabis. Also found are the
distinctive stone harvesting knives of a form that must have
been adopted from China into Northern Pakistan and
Kashmir (Sharma 1982: Stacul 1992). It can therefore be
suggested that domesticated japonica followed other crops
and tools westward and then south to India, via a precursor
of the Silk Road. Once japonica rice was available
alongside local, unimproved proto-indica cultivars, hybrid-
isation and back-crossing could rapidly improve crops in
the active selection regimes of farmers.
By contrast, if we set rice aside, the earliest plausible
introductions from the northeast to the Ganges are later,
including the fibre crop ramie (Boehmeria nivea), a citrus
fruit (Citrus medica sensu lato) and mango, which occur all
after ca. 1400 BC (on mango and citron, see Asouti and
Fuller 2008: 91, 114–115; on ramie, see Saraswat et al. 1994:
287).
8. The dispersal of rice through South Asia
The sub-continental record for rice (Fig. 6) indicates the
diffusion of this species from the Ganges region and/or
Eastern India (Orissa) starting in the late third millennium
and an important presence of rice cultivation on the
peninsula only in the first millennium BC (Fuller 2002).
Outside of a Ganges core, rice occurs first in parts of the
greater Indus valley and perhaps in parts of Gujarat and
Rajasthan, although this may have also involved introduced
japonica rice or hybrids thereof. It is not clear if any of this
Archaeol Anthropol Sci (2010) 2:115–131
early rice cultivation was based on paddy field systems or
was merely supported by seasonal floods or monsoon rains.
With development of paddy-like systems in the Ganges,
however, a major southwards push along the east coast to
Tamil Nadu occurred in Late Iron Age times (Fuller and
Qin 2009). In this same general period, indica rice moved
probably by overseas trade to Southeast Asia and Sri
Lanka.
9. The spread of paddy field cultivation through Korea
and Japan
Setting aside the problematic report of Oryzoid remains
at Late Pleistocene Sorori and debates over the reliability of
Chulmun finds of around 2000 BC (see Ahn 2010), there is
no doubt that rice was widely cultivated in Korea by the
end of the second millennium BC. Based on material
culture evidence, rice, including wetfield cultivation, is
inferred to spread from the Shandong Peninsula to the
Liaoning peninsula and to Korea (Norton 2007; Ahn 2010).
Most early rice dates are associated with the Early Mumun
period which starts from ca. 1200 BC. Mumun sites have
also yielded apparent paddy field systems, although dry
fields are also reported, and it seems likely that both rain-
fed and irrigated rice were part of the agricultural repertoire
of Mumun Korea (Ahn 2010).
It is broadly accepted that the introduction of paddy field
rice farming into Japan came from Korea at the transition to
the Yayoi period, between 1000 and 500 BC. The presence of
rice paddy agriculture in northern Kyushu, western Japan
during the early first millennium BC is well documented (e.g.
Imamura 1996; Kuraku 2002; Hudson 2003; Takahashi
2009). The database from Japan is extensive and cannot be
fully reviewed here, although Takahashi (2009) maps 29
sites with early paddy field evidence. Many of these have
included geoarchaeology, phytolith, diatom and insect
studies (e.g. Mori 2002; Kuraku 2002).
10. Further Asian spreads and crossings in northwestern
Southeast Asia
There was necessarily further dispersal from several of
the above areas: westwards from Indo-China, south/west-
wards from Yunnan and eastwards from the Ganges
towards a ‘melting-pot’ ecologically and culturally in
Bangladesh, Assam and western Burma. This mixing and
diversification process may account for the highly hybri-
dised but often locally divergent nature of aus rices, which
recent genetic data suggest to have higher levels of genomic
hybridisation than other indica rice or japonica (McNally et
al. 2009). In much of modern Southeast Asia, tropical
japonica cultivation is focussed in upland areas, whereas
indica is grown in the lowlands. The origins of this pattern
are unclear, but it is of interest that evidence from Metal
Age Khao Sam Kaeo (400–0 BC) on the Malay Peninsula
Archaeol Anthropol Sci (2010) 2:115–131
is suggestive of dry, upland cultivation (Castillo, unpub-
lished data). By contrast, the expansion of urban polities
across the plains of central Thailand in the early centuries
AD culminating the Dvaravati period with dense distribu-
tions of moated settlements is attributed to intensive,
lowland (and presumably paddy field) cultivation (Mudar
1999), which we might expect to represent indica.
11. Westward dispersals: Persia and Madagascar
During the early centuries AD, rice was dispersed further
westward. From India, rice moved towards western Asia
through Persia. The only datum for placing rice in Western
Asia is Parthian evidence (first century AD) from Susa in
Iran (Miller 1981). This northwestern dispersal presumably
included some of the aromatic rices, which have their
origins mainly from tropical japonica rices (Kovach et al.
2009), although some hybridisations with indica would
have occurred (Jain et al. 2004). Whether or not more
typical indica rices accompanied this spread is unknown.
Meanwhile, from Island Southeast Asia, rice was carried by
Austronesian ancestors of the Malagasy to Madagascar. The
date of the initial settlement of Madagascar by Asian
immigrants remains controversial, with archaeological
evidence for human presence as old as the first century
AD, whilst linguistics arguments suggest a fifth century AD
or later date (Dewar and Wright 1993). Linguistics
normally derives this migration from Southwest Borneo.
Modern rices in Madagascar include both tropical japonica
and indica types, and it is plausible that by the early
centuries AD, both lineages were present in Southeast Asia
(as indicated above for #9).
Discussion: towards a framework of ecological
trajectories and cultural preferences in early rice
Asian rices have had a complex history. Genetic research
and archaeobotany have the tools to provide an increasing
resolution to this history. Genetics provides access to
identifying some of the key genes in the evolution of
domesticated rice, where they are distributed and are likely
to originate, as well as the phylogenetic and reticulate
relationships of rice lineages. Hitherto, most genetically
based models of rice origins, with one or two origins, and
straight forward outward dispersals have been too simple to
account for the data now available: deep neutral genome
divergence, large numbers of shared selected alleles and
widespread evidence for hybridisation between lineages.
Similarly, archaeologists have been too quick to pigeonhole
rice as domesticated or wild, indica or japonica, and have
taken less account of the evolutionary dynamics and
processes by which domestication traits emerged at the
level of populations in response to human activities, or how
127
the modern varietal groupings of indica and japonica are
themselves emergent outcomes of evolutionary processes.
This review has attempted to sketch an integrated synthesis,
although gaps in the evidence have had to be filled with
conjecture. Nevertheless, a series of working hypotheses
have emerged which may structure further research.
References
Adams JM, Faure H (1997) Preliminary vegetation maps of the world
since the last glacial maximum: an aid to archaeological
understanding. J Archaeol Sci 24:623–647
Ahn S (2010) The emergence of rice agriculture in Korea: archae-
obotanical perspectives. Archaeol Anthropol Sci 2(2).
doi:10.1007/s12520-010-0029-9
Allaby R (2010) Integrating the processes in the evolutionary system
of domestication. J Exp Bot 61:935–944. doi:10.1093/jxb/erp382
An Z (1999) The origin and eastward spread or rice-planting culture of
China. Wenwu 2:63–70, In Chinese
Asouti E, Fuller DQ (2008) Trees and woodlands of South India:
archaeological perspectives. Left Coast, Walnut Creel, Retrieved
from http://www.amazon.co.uk/Trees-Woodlands-South-India-Ar
chaeological/dp/1598742310
Barton H, Paz V (2007) Subterranean diets in the tropical rain forests
of Sarawak, Malaysia. In: Denham T, Iriarte J, Vrydaghs L (eds)
Rethinking agriculture. Archaeological and ethnoarchaeological
perspectives. Left Coast, Walnut Creek, pp 50–77
Bellwood P (1997) Prehistory of the Indo-Malaysian Archipelago
(Second rev.). University of Hawaii Press, Honolulu
Bellwood P (2005) First farmers: the origins of agricultural societies.
Blackwell, Oxford
Boaretto E, Wu X, Yuan J, Bar-Yosef O, Chu V, Pan Y, Liu K, Cohen
D, Jiao T, Li S, Gu H, Goldberg P, Weiner S (2009) Radiocarbon
dating of charcoal and bone collagen associated with early
pottery at Yuchanyan Cave, Hunan Province, China. Proc Natl
Acad Sci 106(24):9595–9600
Cai HW, Morishima H (2002) QTL clusters reflect character
associations in wild and cultivated rice. Theor Appl Genet
104:1217–1228
Caicedo AL, Williamson SH, Hernandez RD, Boyko A, Fiedel-Alon
A, York TL, Polato NR, Olsen KM, Nielsen R, McCouch SR,
Bustamante CD, Purugganan MD (2007) Genome-wide patterns
of nucleotide polymorphism in domesticated rice. PLoS Genetics
3(9):e163. doi:10.1371/journal.pgen.0030163
Cappers R (2006) Roman foodprints at Berenike: archaeobotanical
evidence of subsistence and trade in the Eastern Desert of Egypt.
Costen Institute of Archaeology, UCLA, Los Angeles
Chang K (1980) Shang civilization. Yale University Press, New
Haven
Chen W, Nakamura I, Sato Y, Nakai H (1993) Distribution of deletion
type in cpDNA of cultivated and wild rice. Jpn J Genet 68:597–603
Costantini L (1979) Plant remains at Pirak. In: Jarrige J, Santoni M
(eds) Fouilles de Pirak, vol. 1. Diffusion de Boccard, Paris, pp
326–333
Costantini L (1987) Appendix B. Vegetal remains. In: Stacul G (ed)
Prehistoric and protohistoric Swat, Pakistan. Instituto Italiano per
il Medio ed Estremo Orientale, Rome, pp 155–165
Crawford GW, Lee G (2003) Agricultural origins in the Korean
Peninsula. Antiquity 77(295):87–95
Crawford G, Underhill A, Zhao Z, Lee G, Feinman G, Nicholas L,
Luan F, Yu H, Fang H, Cai F (2005) Late Neolithic plant remains
128
from Northern China: preliminary results from Liangchengzhen,
Shandong. Curr Anthropol 46(2):309–317
Cunniffe J, Osborne CP, Ripley BS, Charles M, Jones G (2008)
Response of wild C4 crop progenitors to subambient CO2
highlights a possible role in the origin of agriculture. Glob
Chang Biol 14:576–587
D'Andrea AC (1999) The dispersal of domesticated plants into north-
eastern Japan. In: Gosden C, Hather JG (eds) The prehistory of
food. Appetites for change. Routledge, London, pp 166–183
Dewar RE, Wright HT (1993) The culture history of Madagascar. J
World Prehist 7(4):417–466
Doust AN (2007) Architectural evolution and its implications for
domestication in grasses. Ann Bot 100(5):941–950
Fan C, Xing Y, Mao H, Lu T, Han B, Xu C, Li X, Zhang Q (2006)
GS3, a major QTL for grain length and weight and minor QTL
for grain width and thickness in rice, encodes a putative
transmembrane protein. Theor Appl Genet 112:1164–1171
Fujio S (2004) The beginning of agriculture in the Japanese islands.
In: Society of Archaeological Studies (ed) Cultural diversity and
the archaeology of the 21st century. Society of Archaeological
Studies, Okayama, pp 62–73
Fujiwara H (1993) Research into the history of rice cultivation using
plant opal analysis. In: Pearsall DM, Piperno DR (eds) Current
research in phytolith analysis: applications in archaeology
and palaeoecology. MASCA Research Papers in Science and
Archaeology, volume 10. University of Pennsylvania, Philadelphia,
pp 160–174
Fuller DQ (2002) Fifty years of archaeobotanical studies in India:
laying a solid foundation. In: Settar S, Korisettar R, Settar S,
Korisettar R (eds) Indian archaeology in retrospect, volume III.
Archaeology and interactive disciplines. Manohar, New Delhi, pp
247–364
Fuller DQ (2003) An agricultural perspective on Dravidian historical
linguistics: archaeological crop packages, livestock and Dravid-
ian crop vocabulary. In: Bellwood P, Renfrew C (eds) Examining
the farming/language dispersal hypothesis. Mc Donald Institute
for Archaeological Research, Cambridge, pp 191–214
Fuller DQ (2006) Agricultural origins and frontiers in South Asia: a
working synthesis. J World Prehist 20(1):1–86
Fuller DQ (2007) Contrasting patterns in crop domestication and
domestication rates: recent archaeobotanical insights from the
Old World. Ann Bot 100(5):903–924
Fuller DQ (2009) Indian archaeobotany watch: Lahuradewa 2008. 25
June 2009 Web-log, retrieved from the internet: http://archae
obotanist.blogspot.com/2009/06/indian-archaeobotany-watch-
lahuradewa.html
Fuller DQ, Allaby RG (2010) Seed dispersal and crop domestication:
shattering, germination and seasonality in evolution under cultiva-
tion. In: Østergaard L (ed) Fruit development and seed dispersal,
annual plant reviews 38. Wiley-Blackwell, Oxford, pp 238–295
Fuller DQ, Boivin NL (2009) Crops, cattle and commensals across the
Indian Ocean: current and potential archaeobiological evidence.
Étud Océan Indien 42–43:13–46
Fuller DQ, Harvey EL (2006) The archaeobotany of Indian pulses:
identification, processing and evidence for cultivation. Env
Archaeol 11(2):219–246. doi:10.1179/174963106x123232
Fuller D, Qin L (2008) Immature rice and its archaeobotanical
recognition: a reply to Pan. Antiquity 82(316), on-line. Retrieved
from http://antiquity.ac.uk/ProjGall/fuller2/index.html
Fuller D, Qin L (2009) Water management and labour in the origins
and dispersal of Asian rice. World Archaeol 41(1):88–111
Fuller DQ, Zhang H (2007) A preliminary report of the survey
archaeobotany of the upper Ying Valley (Henan Province) [in
Chinese and English]. In: University and Henan Provincial
Institute of Archaeology (ed) Dengfeng wangchenggang yizhi
de faxian yu yanjiu (2002–2005) [Archaeological Discovery and
Archaeol Anthropol Sci (2010) 2:115–131
Research at the Wangchenggang Site in Dengfeng (2002–2005)]
[in Chinese]. Great Elephant, Zhengzhou, pp 916–958
Fuller DQ, Harvey E, Qin L (2007) Presumed domestication? Evidence
for wild rice cultivation and domestication in the fifth millennium
BC of the lower Yangtze region. Antiquity 81:316–331
Fuller DQ, Qin L, Zheng Y, Zhao Z, Chen X, Hosoya LA, Sun G (2009)
The domestication process and domestication rate in rice: spikelet
bases from the Lower Yangtze. Science 323(5921):1607–1610
Garris AJ, Tai TH, Coburn J, Kresovich S, McCouch SR (2005)
Genetic structure and diversity in Oryza sativa L. Genetics
169:1631–1638
Glover IC, Higham CF (1996) New evidence for early rice cultivation
in South, Southeast and East Asia. In: Harris DR (ed) The origins
and spread of agriculture and pastoralism in Eurasia. UCL,
London, pp 413–441
Gupta PK, Rustgi S, Kumar N (2006) Genetic and molecular basis of
grain size and grain number and its relevance to grain productivity in
higher plants. Genome 571:565–571. doi:10.1139/G06-063
Hammer K (1984) Das Domestikationssyndrome. Kulterpflanze
32:11–34
Harlan JR, De Wet JM, Price EG (1973) Comparative evolution of
cereals. Evolution 27:311–325
Harvey EL (2006) Early agricultural communities in Northern and
Eastern India: an archaeobotanical investigation. Unpublished
Ph.D. dissertation, University College London
Harvey EL, Fuller D, Pal JN, Gupta MC (2003) Early agriculture of
the Neolithic Vindhyas (North Central India). In: Franke-Vogt U,
Weisshaar H-J (eds) South Asian archaeology, Proceedings of the
seventeenth international conference of the European Association
of South Asian Archaeologists (7–11 July 2003, Bonn). Deutches
Archaeologisches Institut, Achen, pp 329–334
Heiser CB (1988) Aspects of unconscious selection and the evolution
of domesticated plants. Euphytica 37:77–81
Higham C (2003) Languages and farming dispersals: Austroasiatic
languages and rice cultivation. In: Bellwood P, Renfrew C (eds)
Examining the farming/language dispersal hypothesis. Mc Donald
Institute for Archaeological Research, Cambridge, pp 223–232
Higham C, Higham T (2009) A new chronological framework for
prehistoric Southeast Asia, based on a Bayesian model from Ban
Non Wat. Antiquity 83(319):125–144
Ho P-T (1956) Early-ripening rice in Chinese history. Econ Hist Rev 9
(2):200–218
Ho P-T (1977) The indigenous origins of Chinese agriculture. In: Reed
CA (ed) Origins of agriculture. Mouton, The Hague, pp 413–484
Horiuchi T, Samy SJ, Phang CC (1971) Grain loss during hand
harvesting in the rice cultivation in Kedah, West Malaysia. Tonan
Ajia Kinkyu (Southeast Asian Studies) 9(2):220–226
Hudson M (2003) Agriculture and language change in the Japanese
Islands. In: Bellwood P, Renfrew AC (eds) Examining the
farming/language dispersal hypothesis. McDonald Institute for
Archaeological Research, Cambridge, pp 311–318
HPIAC [=Hunan Provincial Institute of Archaeology and Cultural
Relics] (2006) Pengtoushan and Bashidang [in Chinese].
Wenwu, Beijing
Huke RE (1982) Agroclimatic and dry-season maps of the South, and
Southeast, and East Asia. International Rice Research Institute,
Los Banos
Imamura K (1996) Prehistoric Japan. New perspectives on insular East
Asia. UCL, London
Ishikawa R, Sato Y-I, Tang T, Nakamura I (2002) Different maternal
origins of Japanese lowland and upland rice populations. Theor
Appl Genet 104:976–980
Jain S, Jain RK, McCouch SR (2004) Genetic analysis of Indian
aromatic and quality rice (Oryza sativa L.) germplasm using
panels of fluorescently-labeled microsatellite markers. Theor
Appl Genet 109:965–977
Archaeol Anthropol Sci (2010) 2:115–131
Jarrige J (1985) Continuity and change in the North Kachi Plain
(Baluchistan, Pakistan) at the beginning of the second millenni-
um B.C. In: Schotmans J, Taddei M (eds) South Asian
archaeology 1983. Instituto Universitario Orientale, Dipartimento
di Studi Asiatici, Naples, pp 35–68
Jiang L, Liu L (2006) New evidence for the origins of sedentism and
rice domestication in the Lower Yangzi River, China. Antiquity
80:355–361
Jin J, Huang W, Gao J, Yang J, Shi M, Zhu M, Luo D, Lin H (2008)
Genetic control of rice plant architecture under domestication.
Nat Genet 40:1365–1369
Kawakami S, Ebana K, Nishikawa T, Sato Y, Vaughan DA, Kadowaki
K (2007) Genetic variation in the chloroplast genomes suggest
multiple domestication of cultivated Asian rice (Oryza sativa L.).
Genome 50:180–187
Keally CT (2004) Bad science and the distortion of history:
radiocarbon dating in Japanese Archaeology. Sophia Int Rev
26. Accessed on-line (30 March 2010), http://www.t-net.ne.jp/
∼keally/Reports/sir2004.html
Kim J, Yang D, Nahm W, Yi S, Hong S, Yun H et al (2008) Last
Glacial and Holocene fluvial wetland sedimentary stratigraphy:
comparison between Soro-ri and Jangheung-ri archeological
sites, Korea. Quatern Int 176-177:135–142, Retrieved from
http://dx.doi.org/10.1016/j.quaint.2007.05.013
Kitano H, Futsuhara Y, Satoh H (1993) Morphological variations in
rice cultivars. In: Matsuo T, Hoshikawa K (eds) Science of the
rice plant. Volume One. Morphology. Food and Agriculture
Policy Research Center, Tokyo, pp 79–88
Konishi S, Izawa T, Lin SY, Ebana K, Fukuta Y, Sasaki T, Yano M
(2006) An SNP caused loss of seed shattering during rice
domestication. Science 312(5778):1392–1396
Kovach MJ, Sweeney MT, McCouch SR (2007) New insights into
the history of rice domestication. Trends Genet 23:578–
587
Kovach MJ, Calingacion MN, Fitzgerald MA, McCouch SR (2009)
The origin and evolution of fragrance in rice (Oryza sativa L.).
Proc Natl Acad Sci USA 106(34):14444–14449
Kuraku Y (2002) The origin and development of rice cultivation in
Japan. In: Yasuda Y (ed) The origins of pottery and agriculture.
Lustre, Delhi, pp 312–317
Kuzmin YV (2006) Chronology of the earliest pottery in East Asia:
progress and pitfalls. Antiquity 80(308):362–371
Lee Y, Woo J (2006) La culture du Paleolithique dans la region de
Jungwon en Coree. Anthropologie 110(2):175–186, Retrieved
from http://www.sciencedirect.com/science/article/B6X0X-
4JTRTBR-2/2/94d21c38ae6e2e732a883b8498039a8f
Lee G-A, Crawford G, Liu L, Chen X (2007) Plants and people from
the early Neolithic to the Shang periods in North China. Proc
Natl Acad Sci 104(3):1087–1092
Li C, Zhou A, Sang T (2006a) Genetic analysis of the rice
domestication syndrome with the wild annual species, Oryza
nivara. New Phytol 170(1):185–193
Li C, Zhou A, Sang T (2006b) Rice domestication by reducing
shattering. Science 311:1936–1939
Li X, Dodson J, Zhou X, Zhang H, Masutomoto R (2007) Early
cultivated wheat and broadening of agriculture in Neolithic
China. Holocene 17:555–560
Lin Z, Griffith ME, Li X, Zhu Z, Tan L, Fu Y et al (2007) Origin of
seed shattering in rice (Oryza sativa L.). Planta 226:11–20
Liu L, Lee G, Jiang L, Zhang J (2007) Evidence for the early
beginning (c. 9000 cal. BP) of rice domestication in China: a
response. Holocene 17:1059–1068
Londo JP, Chiang Y, Hung K, Chiang T, Schaal BA (2006)
Phylogeography of Asian wild rice, Oryza rufipogon, reveals
multiple independent domestications of cultivated rice, Oryza
sativa. Proc Natl Acad Sci USA 103:9578–9583
129
Lu TLD (2006) The occurrence of cereal cultivation in China. Asian
Perspect 45(2):129–158
Lu TLD (2009) Prehistoric coexistence: the expansion of farming
society from the Yangzi River to Western South China. In: Ikeya
K, Ogawa H, Mitchell P (eds) Interactions between hunter–
gatherers and farmers: from prehistory to present. National
Museum of Ethnology, Osaka, pp 47–52
Lu H, Liu Z, Wu N, Berné S, Saito Y, Liu B, Wang L (2002) Rice
domestication and climatic change: phytolith evidence from East
China. Boreas: An International Journal of Quaternary Research
31(4):378–385
Ma J, Bennetzen JL (2004) Rapid recent growth and divergence of rice
nuclear genomes. Proc Natl Acad Sci USA 101:12404–12410
McNally KL, Childs KL, Bohnert R, Davidson RM, Zhao K, Ulat VJ,
Zeller G, Clark RM, Hoen DR, Bureau TE, Stokowski R,
Ballinger DG, Frazer KA, Cox DR, Padhukasahasram B,
Bustamante CD, Weigel D, Mackill DJ, Bruskiewich RM, Ratsch
G, Buell CR, Leung H, Leach JE (2009) Genomewide SNP
variation reveals relationships among landraces and modern
varieties of rice. Proc Natl Acad Sci USA 106(30):12273–12278
Miller NF (1981) Plant remains from Ville Royale II, Susa. Cah DAFI
12:137–142
Mori Y (2002) The origin and development of rice paddy cultivation
in Japan based on evidence of insect and diatom fossils. In:
Yasuda Y (ed) The origins of pottery and agriculture. Lustre,
Delhi, pp 273–296
Mudar KM (1999) How many Dvaravati Kingdoms? Locational
analysis of first millennium A.D. Moated settlements in Central
Thailand. J Anthropol Archaeol 18:1–28
Nakamura S (2010) The origin of rice cultivation in the lower Yangtze
Region, China. Archaeol Anthropol Sci 2 (this issue)
Nakamura I, Urairong H, Kameya N, Fukuta Y, Chitrkon S, Sato YI
(1998) Six different plastid subtypes were found in the O. sativa–
O. rufipogon complex. Rice Genet Newslett 15:80–82
Nasu H, Momohara A, Yasuda Y, He J (2007) The occurrence and
identification of Setaria italica (L.) P. Beauv. (foxtail millet)
grains from the Chengtoushan site (ca. 5800 cal B.P.) in central
China, with reference to the domestication centre in Asia. Veg
Hist Archaeobot 16(6):481–494
Nitsuma Y (1993) Upland rice. In: Matsuo T, Hoshikawa K (eds)
Science of the rice plant, vol. 1. Morphology. Food and
Agriculture Policy Research Center, Tokyo, pp 70–76
Norton CJ (2007) Sedentism, territorial circumscription, and the
increased use of plant domesticates across Neolithic–Bronze
Age Korea. Asian Perspect 46(1):133–165
Ohtsubo H, Cheng C, Ohsawa I, Tsichimoto S, Ohtsubo E (2004) Rice
retroposon p-SINE1 and origin of cultivated rice. Breed Sci 54:1–11
Oka H (1988) Origins of cultivated rice. Elsevier, Amsterdam
Olsen KM, Purugganan MD (2002) Molecular evidence on the origin
and evolution of glutinous rice. Genetics 162:941–950
Pan Y (2008) Immature wild rice harvesting at Kuahuqiao, China?
Antiquity 82. Online: http://www.antiquity.ac.uk/ProjGall/pan/in
Paz V (2003) Island Southeast Asia: spread or friction zone? In:
Bellwood P, Renfrew AC (eds) Examining the farming/language
dispersal hypothesis. McDonald Institute for Archaeological
Research, Cambridge, pp 275–285
Purugganan MD, Fuller DQ (2009) The nature of selection during
plant domestication. Nature 457(7231):843–848. doi:10.1038/
nature07895
Qin L, Fuller (Fu Daolian) DQ (2009) Appendix 3. The Nanjiaokou
site 2007 excavated Early to Mid Yangshao plant remains. In:
Henan Provincial Institute of Cultural Relics and Archaeology
(ed) Nanjiaokou site in Sanmenxia. Science, Beijing, pp 427–
435, In Chinese
Ren G, Beug H (2002) Mapping Holocene pollen data and vegetation
of China. Quatern Sci Rev 21:1395–1422
130
Rispoli F (2008) The incised and impressed pottery of Mainland
Southeast Asia: following the paths of Neolithization. East West
57:235–304
Rosen A (2008) The impact of environmental change and human land
use on alluvial valleys on the Loess Plateau of China during the
Mid-Holocene. Geomorphology 101:298–307
Sato Y, Tang SX, Yang LU, Tang LH (1991) Wild-rice seeds in the
oldest rice remains. Rice Genetics Newsletter 8:76
Sagart L (2008) The expansion of Setaria farmers in East Asia: a
linguistic and archaeological model. In: Sanchez-Mazas A,
Blench RM, Ross M, Lin M, Pejros I (eds) Human migrations
in continental East Asia and Taiwan: matching archaeology,
linguistics and genetics. Taylor & Francis, London, pp 133–157
Sage RF (1995) Was low atmospheric CO2 during the Pleistocene a
limiting factor for the origin of agriculture? Glob Chang Biol
1:93–106
Sang T, Ge S (2007) The puzzle of rice domestication. J Integr Plant
Biol 49(6):760–768
Saraswat KS, Sharma NK, Saini DC (1994) Plant economy ay ancient
Narhan (ca. 1,300 B.C.–300/400 A.D.). In: Singh P (ed)
Excavations at Narhan (1984–1989). Banaras Hindu University,
Varanasi, pp 255–346
Sato Y (1996) DNA-ga Kataru Inasaku Bunmaei Kingento Tenkai
[The origins of rice culture inferred from DNA analysis] (in
Japanese). NHK Books, Tokyo
Sato Y (2002) Origin of rice cultivation in the Yangtze River Basin.
In: Yasuda Y (ed) The origins of pottery and agriculture. Lustre,
New Delhi, pp 143–150
Saxena A, Prasad V, Singh IB, Chauhan MS, Hassan R (2006) On the
Holocene record of phytoliths of wild and cultivated rice from
Ganga Plain: evidence for rice-based agriculture. Curr Sci 90
(11):1547–1552
Shan J, Zhu M, Shi M, Gao J, Lin H (2009) Fine mapping and
candidate gene analysis of spd6, responsible for small panicle
and dwarfness in wild rice (Oryza rufipogon Griff.). Theor Appl
Genet 119(5):827–836
Sharma AK (1982) Excavations at Gufkral, 1981. Puratattva 11:19–25
Shi Y, Kong Z, Wang S, Tang L, Wang F, Yao T et al (1993) Mid-
Holocene climates and environments in China. Glob Planet
Change 7:219–233
Shomura A, Izawa T, Ebana K, Ebitani T, Kanegae H, Konishi S, Yano M
(2008) Deletion in a gene associated with grain size increased yields
during rice domestication. Nat Genet 40:1023–1028
Song X, Huang W, Shi M, Zhu M-Z, Lin H (2007) A QTL for rice
grain width and weight encodes a previously unknown RING-
type E3 ubiquitin ligase. Nat Genet 39:623–630
Stacul G (1992) Swat, Pirak, and connected problems (mid-2nd
millennium B.C.). In: Jarrige C, Gerry JP, Meadow RH (eds)
South Asian archaeology 1989. Prehistory, Madison, pp 267–270
Sweeney MT, McCouch SR (2007) The complex history of the
domestication of rice. Ann Bot 100:951–957
Takahashi R (2009) Symbiotic relations between paddy-field rice
cultivators and hunter-gatherer-fishers in Japanese prehistory:
archaeological considerations on the transition from the Jomon
Age to the Yayoi Age. In: Ikeyea K, Ogawa H, Mithcell P (eds)
Interactios between hunter-gatherers and farmers: from prehistory
to present. Senri Ethnological Studies 73. National Museum of
Ethnology, Osaka, pp 71–98
Takahashi H, Sato Y, Nakamura I (2008) Evolutionary analysis of two
plastid DNA sequences in cultivated and wild species of Oryza.
Breed Sci 233:225–233
Takamiya H (2001) Introductory routes of rice to Japan: an examination
of the Southern Hypothesis. Asian Perspect 40(2):209–226
Tan L, Li X, Liu F, Sun X, Li C, Zhu Z et al (2008) Control of a key
transition from prostrate to erect growth in rice domestication.
Nat Genet 40:1360–1364
Archaeol Anthropol Sci (2010) 2:115–131
Tanaka K, Ishikawa R, Honda T (2010) Rice archaeological remains and
the possibility of DNA archaeology: examples from Yayoi and Heian
periods of Northern Japan. Archaeol Anthropol Sci (this volume)
Tang JB, Xia HA, Cao ML, Zhang XQ, Zheng WY, Hu SN et al
(2004) A comparison of rice chloroplast genomes. Plant Physiol
135:412–420
Tewari R, Srivastava RK, Singh KK, Saraswat KS, Singh IB, Chauhan
MS et al (2006) Second preliminary report of the excavations at
Lahuradewa. District Sant Kabir Nagar, U. P.: 2002–2003–2004
& 2005–06. Pragdhara 16:35–68
Tewari R, Srivastava RK, Saraswat KS, Singh IB, Singh KK (2008)
Early farming at Lahuradewa. Pragdhara 18:347–373
Thompson GB (1996) The excavations of Khok Phanom Di, a
prehistoric site in Central Thailand. Volume IV. Subsistence and
environment: the botanical evidence. The biological remains part
III. The Society of Antiquaries of London, London
Thompson GB (1997) Archaeobotanical indicators of rice domestica-
tion—a critical evaluation of diagnostic criteria. In: Ciarla R,
Rispoli F (eds) South-East Asian archaeology 1992. Instituto
Italiano per il Medio ed Estremo Orientale, Rome, pp 159–174
Toyama S (2002) The origin and spread of rice cultivation as seen
from rice remains. In: Yasuda Y (ed) The origins of pottery and
agriculture. Lustre, Delhi, pp 263–272
Tsang C (2005) Recent discoveries at a Tapenkeng culture site in
Taiwan: implications for the problem of Austronesian origins. In:
Sagart L, Blench R, Sanchez-Mazas A (eds) The peopling of East
Asia: putting together archaeology, linguistics and genetics.
Routledge-Curzon, London, pp 63–74
Vaughan DA, Kadowaki K, Kaga A, Tomooka N (2005) On the
phylogeny and biogeography of the genus Oryza. Breed Sci
55:113–122
Vaughan DA, Lu B, Tomooka N (2008a) The evolving story of rice
evolution. Plant Sci 174(4):394–408
Vaughan DA, Lu B, Tomooka N (2008b) Was Asian rice (Oryza
sativa) domesticated more than once? Rice 1:16–24
Vincent B (2003) Rice in pottery: new evidence for early rice
cultivation in Thailand. Indo-Pacific Prehistory Association
Bulletin 23:51–58
Vitte C, Ishii T, Lamy F, Brar D, Panaud O (2004) Genomic
paleontology provides evidence for two distinct origins of Asian
rice (Oryza sativa L.). Mol Genet Genomics 272:504–511
Weber S, Lehman H, Barela T, Hawks S, Harriman D (2010) Rice or
millets: early farming strategies in prehistoric central Thailand.
Archaeol Anthropol Sci 2 (this volume). doi:10.1007/s12520-
010-0030-3
White JC (1995) Modeling the development of early rice agriculture:
ethnoecological perspectives from northeast Thailand. Asian
Perspect 34:37–68
Xiong LZ, Liu KD, Dai XK, Xu CG, Zhang Q (1999) Identification of
genetic factors controlling domestication-related traits of rice
using an F2 population of a cross between Oryza sativa and O.
rufipogon. Theor Appl Genet 98(2):243–251. doi:10.1007/
s001220051064
You X (1987) The wild rice in Chinese ancient records
(Zhongguo Gushu Zhong Jizai de Yeshengdao Tantao) [in
Chinese]. Gujin Nongye [Ancient and Modern Agriculture]
1987(1):1–6
Yu SY, Zhu C, Song J, Qu WZ (2000) Role of climate in the rise and fall
of Neolithic cultures on the Yangtze Delta. Boreas 29:157–165
Zhang W (2002) The bi-peak tubercle of rice, the character of ancient
rice and the origin of cultivated rice. In: Yasuda Y (ed) The
origins of pottery and agriculture. Lustre, Delhi, pp 205–216
Zhang C, Hung H-C (2008) The Neolithic of southern China—origin,
development and dispersal. Asian Perspect 47(2):299–329
Zhang C, Hung H (2010) The emergence of agriculture in southern
China. Antiquity 84(323):11–25
Archaeol Anthropol Sci (2010) 2:115–131
Zhang J, Wang X (1998) Notes on the recent discovery of ancient
cultivated rice at Jiahu, Henan Province: a new theory
concerning the origin of Oryza japonica in China. Antiquity
72:897–901
Zhang LB, Zhu Q, Wu ZQ, Ross-Ibarra J, Gaut B, Ge S, Sang T
(2009) Selection on grain shattering genes and rates of rice
domestication. New Phytol 184:708–720
Zhao Z (1998) The Middle Yangtze region in China is one place
where rice was domesticated: phytolith evidence from the
Diaotonghuan Cave, Northern Jiangxi. Antiquity 72:885–897
Zhao Z (2010) New data and new issues for the study of origin of
rice agriculture in China. Archaeol Anthropol Sci 2 (this
volume)
131
Zhao Z, Piperno D (2000) Late Pleistocene/Holocene environments in
the Middle Yangtze River Valley of the Taihu Lake in China and
rice (Oryza sativa L.) domestication: the phytoliths evidence.
Geoarchaeology 15:203–222
Zhao Z, Pearsall DM, Benfer RA, Piperno DR (1998) Distinguishing
rice (Oryza sativa Poaceae) from wild Oryza species through
phytolith analysis II: finalised method. Econ Bot 52:134–145
Zheng Y, Dong Y, Matsui A, Udatsu E, Fuijiwara H (2003) Molecular
genetic basis of determining subspecies of ancient rice using
shape of phytoliths. J Archaeol Sci 30:1215–1221
Zheng Y, Sun G, Chen X (2007) Characteristics of the short rachillae
of rice from archaeological sites dating to 7000 years ago. Chin
Sci Bull 52:1654–1660