Contrasting Patterns in Crop Domestication and Domestication Rates: Recent
Archaeobotanical Insights from the Old World
DORIAN Q FULLER*
Institute of Archaeology, University College London, 31–34 Gordon Square, London WC1H 0PY, UK
Received: 20 September 2006 Revision requested: 8 December 2006 Accepted: 31 January 2007Published electronically: 10 May 2007
†Background Archaeobotany, the study of plant remains from sites of ancient human activity, provides data for
studying the initial evolution of domesticated plants. An important background to this is defining the domestication
syndrome, those traits by which domesticated plants differ from wild relatives. These traits include features that have
been selected under the conditions of cultivation. From archaeological remains the easiest traits to study are seed size
and in cereal crops the loss of natural seed dispersal.
†Scope The rate at which these features evolved and the ordering in which they evolved can now be documented for
a few crops of Asia and Africa. This paper explores this in einkorn wheat (Triticum monococcum) and barley
(Hordeum vulgare) from the Near East, rice (Oryza sativa) from China, mung (Vigna radiata) and urd (Vigna
mungo) beans from India, and pearl millet (Pennisetum glaucum) from west Africa. Brief reference is made to
similar data on lentils (Lens culinaris), peas (Pisum sativum), soybean (Glycine max) and adzuki bean (Vigna
angularis). Available quantitative data from archaeological finds are compiled to explore changes with domestication.
The disjunction in cereals between seed size increase and dispersal is explored, and rates at which these features
evolved are estimated from archaeobotanical data. Contrasts between crops, especially between cereals and pulses,
†Conclusions These data suggest that in domesticated grasses, changes in grain size and shape evolved prior to
non-shattering ears or panicles. Initial grain size increases may have evolved during the first centuries of cultiva-
tion, within perhaps 500–1000 years. Non-shattering infructescences were much slower, becoming fixed about
1000–2000 years later. This suggests a need to reconsider the role of sickle harvesting in domestication. Pulses,
by contrast, do not show evidence for seed size increase in relation to the earliest cultivation, and seed size increase
may be delayed by 2000–4000 years. This implies that conditions that were sufficient to select for larger seed size
in Poaceae were not sufficient in Fabaceae. It is proposed that animal-drawn ploughs (or ards) provided the
selection pressure for larger seeds in legumes. This implies different thresholds of selective pressure, for example
in relation to differing seed ontogenetics and underlying genetic architecture in these families. Pearl millet
(Pennisetum glaucum) may show some similarities to the pulses in terms of a lag-time before truly larger-
grained forms evolved.
Key words: Domestication, cultivation, cereals, pulses, archaeobotany, Triticum, Hordeum, Oryza, Vigna, Pennisetum.
This paper will consider archaeobotanical evidence for the
evolutionary stages of domestication and the rates of evol-
ution of the domestication syndrome for a select number
of the best archaeologically documented crops in the Old
World. The quantitative increase in archaeobotanical data
in recent years indicates that the origins of crop cultivation
was a dynamic and multi-stage evolutionary process that
occurred independently in numerous world regions invol-
ving numerous crops. This evidence results from growth
in archaeobotanical research in recent years. Systematic
sampling for archaeobotanical remains, using the technique
of flotation in the field, began slowly in the 1960s and
1970s, and led to the collection of much larger and more
complete data sets (Pearsall, 2000; Fuller, 2002). In sub-
sequent years, increasing numbers of sites were sampled
and more practitioners of archaeobotany have joined the
field. In the past two decades, refinements in identification,
analysis and modelling have occurred. Taken together these
mean that just in the past few years it has become possible
to offer new insights into plant domestication and the mul-
tiple origins of crops.
As it has been recognized that domestication is a multi-
stage process (e.g. Ford, 1985; Harris, 1989, 1996),
models have summarized the expected stages and their
archaeological visibility. A classic model is that of Harris
(1989), who distinguishes four general stages: (1) wild
plant food procurement (true hunting and gathering), (2)
wild plant food production (the very beginnings of cultiva-
tion), (3) systematic cultivation (of morphologically wild
plants) and finally (4) agriculture based on domesticated
plants. Domestication results from the earlier stages of
wild plant food production and systematic cultivation,
making crops more dependent on humans for survival but
also more productive. Through all of these stages people
put increasing labour effort into a single unit of land and
a single field of crops, in other words this tracks an intensi-
fication of production. But the reward is increased
*For correspondence. E-mail email@example.com
# 2007 The Author(s).
This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://
creativecommons.org/licenses/by-nc/2.0/uk/) which permits unrestricted non-commercial use, distribution, and reproduction in any
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Annals of Botany 100: 903–924, 2007
doi:10.1093/aob/mcm048, available online at www.aob.oxfordjournals.org
productivity of that land, and the ability to produce larger
surpluses, to feed more people, or accumulate as wealth.
An updated version of the Harris chart is given here
(Fig. 1), in which I have modified domestication to indicate
that its is a protracted quantitative process of change and
that those rates may vary for different aspects of that
change. Two points should be highlighted: first, that there
is necessarily a stage of production (cultivation) that pre-
cedes morphological domestication, and archaeobotanists
have been developing means to recognize this. Second,
this transition represents an important change in human
behaviour in terms of the nature of efficiency. While for
foragers time is an important measure of efficiency (trans-
port times, processing time, etc.) (see Kelly, 1995), for
farmers, who have invested labour in smaller parcels of
land for delayed returns, space becomes a crucial metric
for efficiency, i.e. yield per unit area (Hillman and
Davies, 1990: 69–70; Bar-Yosef, 1998). This cultural
change is probably crucial to the change in selective press-
ures on plants that evolve into domesticates.
AGRICULTURE EVOLVING: THE INTERPLAY
OF CULTURAL BEHAVIOUR AND PLANT
The origins and evolution of agriculture involves consider-
ing how human food-procurement behaviours have changed
strategically and how plant populations have evolved in
response. The beginnings of food production represents a
strategic shift in human behaviour, towards the manipu-
lation of the soil environment (clearance, tillage) and
through an influence on the composition of plant popu-
lations in that soil (via preferential seeding and tending of
one or a few species). From the vantage point of present-
day plant populations, it is possible to identify the closest
wild relatives of crops, where and in what habitat they
occur, and how these modern domesticates differ from
their presumed wild ancestors (e.g. Zohary, 1969; Harlan,
1992). The transition from the ancient wild forms to the
modern domesticated forms was an evolutionary process,
and it is necessary to ask whether this can be broken
down into distinct stages, relating to different selective
pressures and human behaviours, and at what rate this
evolutionary process took place. Archaeobotany, the study
of plant remains from sites of ancient human activity,
provides data for studying this process of crop evolution,
which is directly correlated through its stratigraphic
context with the archaeological record of human behaviour.
Archaeobotany, however, has its limits imposed by preser-
vation, and as most remains are preserved by being charred
in a fire, we are usually limited to clear morphological dis-
tinctions and metrical shifts.
Before considering the archaeobotanical data, we must
have in mind a framework for assessing changes. The
‘domestication syndrome’ provides such a framework by
highlighting a set of characters that differ between domesti-
cated crops and their wild ancestors (Harlan et al., 1973;
Hawkes, 1983; Zohary and Hopf, 2000; Gepts, 2004).
These characters can be related to different aspects of culti-
vation in terms of what causes them to evolve. It should be
noted that the domestication syndrome differs for different
kinds of crop-plants. Thus, fruit trees, vines and tubers are
not ‘domesticated’ in the same way as grain crops such as
cereals and pulses, in as much as they tend to be reproduced
vegetatively and to represent the human choice of favourable
mutants (Zohary and Spiegel-Roy, 1975; Hather, 1996;
Kislev et al., 2006). This paper will consider only the dom-
estication of seed-propagated herbaceous annuals.
There has been much discussion about how these domes-
tications evolved. Important contributions come from field
ecological studies of wild progenitors (e.g. Zohary, 1969;
Hillman, 2000), experimental harvesting of wild progenitors
using different methods (Harlan, 1967; Ladzinsky, 1987;
Unger-Hamilton, 1989; Hillman
Anderson, 1992; Lu, 1998; Kislev et al., 2004) and exper-
imental cultivation of wild progenitors (e.g. Oka and
Morishima, 1971; Zohary, 1989; Hillman and Davies,
1990; Willcox, 1992, 1999). Nevertheless there remain
important areas of debate about the nature of the changes
in human practices between collecting and cultivating, and
what motivated those cultural shifts, and resulting adaptive
changes in the plants. While these changes require sowing
of seeds stored from a previous harvest, other selection press-
ures are involved which can be related to harvesting methods,
soil conditions (created through tillage, etc.) and perhaps
crop-processing methods. The domestication syndrome in
grain crops usually includes the following six criteria.
1. Elimination/reduction of natural seed dispersal
For example, non-shattering rachis in cereals, non-
dehiscent pod in pulses. This is often regarded as the
single most important domestication trait (‘domestication’
sensu stricto). It makes a species dependent upon the
human farmer for survival. It means that instead of shed-
ding seeds when they are mature, a plant retains them.
FIG. 1. An evolutionary model from foraging to agriculture, with archae-
obotanical expectations indicated at the bottom (modified from Harris,
1989). The stages of pre-domestication cultivation are shaded. In this
version, domestication is represented as a process of gradual frequency
change, with an earlier, more rapid ‘semi-domestication’ and a later,
slower fixation of full domestication. The gap in time elapsed between
these two can be taken as a minimal estimate of domestication rate (d.r.).
Fuller — Patterns in Crop Domestication904
Instead those seeds must be separated by the addition of
human labour (threshing and winnowing), and then the
seeds are dispersed by the farmer. Higher yields can be pro-
duced because the farmer can now wait until all, or most, of
the grains on a plant have matured. This trait can only
evolve under conditions of particular kinds of intensive har-
vesting that favour plants that retain their seeds, followed by
sowing from the harvested seeds. All forms of cultivation
may not select for this trait. While the evolution of this
trait has often been attributed to the early use of sickle
tools (e.g. Wilke et al., 1972; Hillman and Davies, 1990;
Bar-Yosef, 1998), current evidence suggests that this need
not be the case, as discussed below.
2. Reduction in seed dispersal aids
This is connected to the last trait but is selected for in a
different way. Plants often have a range of structures that
aid seed dispersal, including hairs, barbs, awns and even
the general shape of the spikelet in grasses. Thus, domesti-
cated wheat spikelets are less hairy, have shorter or no awns
and are plump, whereas in the wild they are heavily haired,
barbed and aerodynamic in shape (see Hillman and Davies,
1990). All of these tend to be greatly reduced in the dom-
esticated form. This can be considered to have come
about by the removal of natural selection for effective dis-
persal, and once removed metabolic ‘expenditure’ on these
structures is reduced. This trait might evolve under initial
cultivation and be regarded as part of ‘semi-domestication’.
3. Trends towards increasing seed/fruit size
This is likely to be selected for by open environments in
which larger seedlings have advantages, surviving deeper
burial within disturbed soils and thus this trait should be
selected for by tillage and cultivation generally (Harlan
et al., 1973). Larger seeds are strongly correlated with
larger seedlings in many species including cereal and
legume crops (Baskin and Baskin, 2001: 214; Krishnasamy
and Seshu, 1989). Comparative ecology indicates that
larger seeds generally have competitive advantages over
smaller seeds under certain kinds of competition including
deeper burial (Maranon and Grubb, 1993; Westoby et al.,
1996). Although there may be some exceptions, this is
demonstrated from cereal relatives (e.g. Panicum, Lolium,
Avena and Aegilops among grasses; Baskin and Baskin,
2001: 212–213), and in Mediterranean habitats (Maranon
and Grubb, 1993). Experimental cultivation of rice found
some increase in average grain weight within just five gener-
aions (Oka and Morishima, 1971), suggesting this can indeed
evolve quickly. This trait is the key archaeological indicator
of ‘semi-domestication’ in cereals.
4. Loss of germination inhibition
In the wild many seeds will only germinate after certain
conditions have passed, conditions of daylength, tempera-
ture, or after the seed coat is physically damaged. Crops
tend to germinate as soon as they are wet and planted.
This change is often signalled by changes in the seed,
such as thinner seed coats. This is also selected for
simply by cultivation, and sowing from harvested yield,
as those seeds that do not readily germinate will not contrib-
ute to the harvest. This has produced a particularly import-
ant character for the study of New World Chenopodium
domestications (e.g. Smith, 1989; Bruno and Whitehead,
2003). So far it has been less useful for Old World crops.
Although in principle it can be applied to pulses (Butler,
1989; Plitman and Kislev, 1989), seed coats are preserved
rarely in charred archaeological specimens.
5. Synchronous tillering and ripening
This sometimes includes a shift from perennial to annual.
Planting at one time and harvesting at one time will favour
plants that grow in synchronization.
6. More compact growth habit
For example, reduction in branching, e.g. dense spikes or
seed heads (the ‘sunflower effect’; Harlan, 1995: 199), e.g.
from climbing habit to self-standing. Harvesting methods,
like those that select for non-shattering types (no. 1,
above) will also favour plants with single and compact
parts to be harvested.
Of particular importance to the archaeobotanist are those
changes that can be identified in archaeological material.
This is likely to include nos. 1–4, although no. 4 is only
preserved in certain kinds of seeds, and no. 2 may be
difficult to recognize because hairs are often destroyed by
carbonization. For this reason, especially for most cereals,
it is criteria 1 and 3 that archaeologists look at. Grain size
(no. 3) is made complicated because of the potentially
wide range of variation in modern populations, and the
effects of charring (which causes shrinkage and sometimes
distortion). If preserved, remains of the cereal ear rachis or
spikelet base can provide clear evidence for the mode of
shattering (no. 1). In wild types there should be a smooth
scar, indicating normal abscission, whereas in domesticated
(but also in very immature) plants the scar will be rough
because the ear has been broken apart by threshing.
The contrast between shattering and grain size is signifi-
cant. Tough (or non-shattering) rachis ears occur as a rare,
deleterious genetic mutation in most wild grass populations,
which has been demonstrated in wild barley (Kislev, 1997;
Kislev et al., 2004). If wild cereals were harvested simply
by passing through stands and shaking or beating ears to
knock seeds into a basket then the shattering, wild-type
ears would be the ones to predominate in the next year’s
crop. Ethnographically this is the most common method
documented for wild grass and forb seed gathering (e.g.
Harris, 1984; Harlan, 1989). In addition, harvesting exper-
iments in wild wheat and wild Setaria millet indicate that
this is significantly more efficient in terms of return per
unit of labour time (Hillman and Davies, 1990; Lu,
1998). By contrast, if people harvested with a sickle and
cutting the entire ear, or plucking individual ears, or
pulling plants up from the roots, this would tend to disperse
shattering seeds and retain all non-shattering mutants.
Therefore, these could be replanted the following year
Fuller — Patterns in Crop Domestication 905
and over time would come to dominate the population at the
expense of wild, shattering types (Hillman and Davies,
1990). Under ideal circumstances of new soils being
sown, without a seed bank of the wild form, and sickle har-
vesting of mature/near-mature plants, and self-pollination or
sufficient distance from wild stands, the tough rachis geno-
type could evolve very rapidly, with estimates of 20–100
years suggested by Hillman and Davies (1990). As will
be seen below, however, the archaebotanical evidence
appears to indicate a much slower process.
This paper will attempt to arrive at differing modal trajec-
tories in crop domestication and rates of domestication
based on dated archaeological evidence. Of particular signifi-
cance is the distinction between different aspects of the dom-
estication syndrome. First there is the evolution of grain size,
or loss of dispersal appendages, such as hairs and awns,
which can be called ‘semi-domestication’. Second there is
the loss of wild seed dispersal (non-shattering), i.e. full
‘domestication’. The time between these two developments
can be taken to define a domestication rate (‘d.r.’ in
Fig. 1). This provides an archaeological estimate of the
time it took for fully domesticated types to evolve in popu-
lation genetics terms, i.e. to come to dominate cultivated
evidence will be reviewed as it relates to the evolution of
different aspects of the domestication syndrome. A small
selection of crops from the Old World are explored which
represent those with the best quantitative archaebotanical
evidence to date, and which are drawn from across four
different centres of domestication in Asia and Africa.
This paper will begin by exploring a typical cereal model
for domestication, reviewing current evidence for the evol-
ution of the domestication syndrome in some of the main
cereals of the Old World: wheat and barley in the Near
East, and rice in southern China. Although the available
archaeobotanical evidence is quite different, current indi-
cations are that similar evolutionary processes took place
in each case, with evolution of grain shape and size preced-
ing the loss of wild-type seed dispersal.
This paper will also explore a pulse model of domesti-
cation, as the trajectory followed in the evolution of the
full domestication syndrome appears to have differed in
legumes. In particular, there appears to be a long lag-time
between early cultivation and domestication (in terms of
seed dispersal and germination inhibition) and seed size
increase. In pulses, the seed size increase must be con-
sidered a form of cultivar advancement, occurring much
later after the initial development of agriculture (Fuller
and Harvey, 2006: 257). This can be related to changes in
agricultural techniques. This will be explored through the
archaeobotanical evidence for two closely related pulses
domesticated in India, the mungbean (Vigna radiata) and
urdbean (Vigna mungo). A similar model seems to apply
to Near Eastern pulses such as lentils (Lens culinaris) and
peas (Pisum sativum), and possibly east Asian legumes,
adzuki bean (Vigna angularis) and soybean (Glycine
max). Finally, the case of West African pearl millet
(Pennisetum glaucum) is explored as a case in which a
cereal shows some similarities to the pulse domestication
model. A brief overview of the current state of archaeobo-
tanical research in each case region is provided (the Near
East, China, India and sub-Saharan Africa).
Archaeobotanical evidence is most often preserved car-
bonized, as the result of charring, and this process affects
the size and shape of seeds. In general, carbonization
leads to shrinkage. Based on a range of experimental studies
(e.g. Lone et al., 1993; Willcox, 2004; Braadbaart and van
Bergen, 2005; Nesbitt, 2006: 21), most carbonized seeds
appear to shrink between 10 and 20%, with a slight bias
towards higher shrinkage in the longest dimension, i.e. a
tendency to become more spherical. Therefore, in comparing
carbonized seeds withmodern seeds some correction factor is
needed. In this paper I use a 10% shrinkage estimate for
cereals and a higher 20% shrinkage estimate for pulses. The
higher estimate for pulses should serve to over-emphasize
potential seedenlargement inpulses.The most important evi-
dence, however, remains comparisons between archaeo-
Note on ages
Throughout this paper ages are referred in calendar years
BC or AD, based on the calibration of radiocarbon ages. For
measured radiocarbon ages back to 11 000 BP calibration is
made with measurements from tree-ring data, while older
dates are calibrated by atmospheric
coral (Reimer et al., 2004). This was done with OxCal
v.3.9 software. For this process and software, see the
Oxford University Radiocarbon Laboratory webpage:
14C estimated from
THE CEREAL MODEL FOR DOMESTICATION:
WHEAT AND BARLEY
Near Eastern agricultural origins: general background
The Near Eastern centre of crop domestication is often
called the ‘fertile crescent’, and, as is well known, a
number of crop progenitors can be found in a restricted
set of adjacent vegetation zones from the Mediterranean
oak woodlands,through the open grassland steppe
(Fig. 2). The wild wheats (Triticum spp.) and barley
(Hordeum vulgare) occur in the slightly drier, more open
parkland steppe with dispersed shrubs, wild almond trees
and oaks, while the wild pulses of south-west Asia (Lens
culinaris, Pisum sativum, Cicer arietinum, Lathyrus
sativus, Vicia spp.) occur in the clearings of nearby wood-
lands and rocky talus slopes (Zohary and Hopf, 2000). That
hunter-gatherers had long been exploiting wild cereals
when available without cultivation is established from
finds at Ohlalo II (21 000–18 500 BC) of wild emmer
(Triticum diococcoides) and barley (Hordeum spontaneum)
together with several other small-seeded grasses, fruits and
acorns (Kislev et al., 1992; Weiss et al., 2004). Climatic
changes at the end of the Pleistocene are regarded as
important for their impact on the availability of these
wild progenitors and human subsistence and are the
Fuller — Patterns in Crop Domestication906
favoured component underpinning explanations for why
Younger Dryas dry (cold) episode from approx. 11 500 to
9800 BC (Bar-Yosef, 1998, 2003; Harris, 1998; Hillman
et al., 2001; Byrd, 2005). Prior to this event the climate
was favourable and dense populations of hunter-gatherers
(the Late Epipalaeolithic phase, 12 500–9700 BC) were
settled in territories that included some possible year-round
settlements. The Younger Dryas brought an end to this,
with most sites being abandoned, and it is argued that a
few groups may have resorted to cultivation during this
period. Village populations reappeared throughout the
area during the Pre-Pottery Neolithic A (PPNA) Period
(9700–8700 BC), and many of them appear to have been
cultivators. Finds of domesticated plants are generally
widespread in the subsequent Pre-Pottery Neolithic B
(PPNB; 8700–6200 BC), and it is by this period that they
began to spread beyond the domestication zone into
central Turkey, Cyprus, Crete and southern Greece.
Domesticated animals first occur about 8200 BC at the
start of the Middle PPNB (Garrard, 2000; Bar-Yosef,
2003; Colledge et al., 2004; Byrd, 2005).
In recent years two important advances have occurred in
cultivation and evidence for different sub-centres of
crop domestication within this region. Evidence for
pre-domestication cultivation has been recognized through
the statistical composition of wild seed assemblages, for
nearly 10 years (Colledge, 1998, 2001, 2002; Harris, 1998;
Willcox, 1999, 2002; Hillman, 2000; Hillman et al., 2001).
As is well known from later agricultural periods, archaeobo-
tanical assemblages are made up predominately of crops and
weeds, together with some gathered fruits and nuts (Jones,
1985). This pattern is already recognized in the PPNA and
in some Late Epipalaeolithic sites (Fig. 2), by samples domi-
nated by wild cereals together with seeds of herbaceous taxa
that flourish in disturbed soils, which are best known today as
arable weeds. This includes taxa such as small legumes
(Trifoliae), Boraginaceae, small-seeded grasses (Hordeum
murinum, Bromus sp., Lolium sp., Lepturus pubescens,
Lololium sp.), Polygonaceae, Silene (Caryophyllaceae),
Galium (Rubiaceae) and
Modelling of the impact of Younger Dryas climate change
on species availability indicates that only cereals and these
weeds show frequency increases against the grain of the
expected climate trends through the sequence at Abu
Hureyra, suggesting their persistence in a new habitat: the
arable field (Hillman et al., 2001). This is also suggested
by the continued association of these same weeds with dom-
esticated cereals in later periods (from the PPNB, 8700–
6200 BC, and onwards).
A number of advances in recent years have made it
increasingly clear that separate histories need to be traced
for different sub-regions, which were separate in as much
as the beginnings of agriculture relied on different combi-
nations of species. Some suggestive evidence comes from
genetics. For example, in barley (Hordeum vulgare) two
variant genes control whether or not the ear shatters. A
recessive mutation in either gene locus leads to the domesti-
cated condition, while the dominant variant at either locus
FIG. 2. Map of south-west Asia, showing the locations of sites with archaeobotanical evidence that contributes to understanding the origins and spread of
agriculture. Sites are differentiated on the basis of whether they provide evidence for pre-domestication cultivation, enlarged grains, mixed or predomi-
nantly domestic-type rachis data. Note that these sites represent a range of periods, and many sites have multiple phases of use, in which case the earliest
phase with significant archaeobotanical data is represented. Shaded areas indicate the general distribution of wild progenitors (based on Zohary and Hopf,
2000, with some refinements from Willcox, 2005). It should be noted that wild emmer (Triticum dicoccoides) occurs over a sub-set of the wild barley
zone, and mainly in the western part of the crescent.
Fuller — Patterns in Crop Domestication907
confers wild-type shattering ears (Zohary and Hopf, 2000:
59–60). The existence of these two variants argues for
two domestications for barley. Recent work on emmer
wheat has identified two different lineages of a gluten
gene which are so different that they are estimated to
have evolved apart 100 000s of years ago, and thus
amongst wild emmer wheat, long before domestication.
Such evidence implies two separate domestications of
emmer (Allaby et al., 1999: 305; Brown, 1999; reviewed
in Jones and Brown, 2000). Another source of evidence
for multiple domestications of the ‘same’ (or similar)
crops comes from refinements in archaeobotanical identifi-
cation criteria. Thus, for example, it is possible on the basis
of grain shape to distinguish einkorn wheat (Triticum
monococcum) with single-grained spikelets (from wild
Triticum boeoticum subsp. aegilopoides) from einkorn
with two-grained spikelets (from wild T. boeoticum
subsp. thaudar or T. urartu). Modern domesticated
einkorn (T. monococcum) is normally only one-grained,
but archaeobotanical evidence indicates the presence of
one of these two grained forms as a wild cereal from the
late Pleistocene in Syria (Hillman, 2000; Willcox, 2002,
2005), and later as a domesticated cereal in Syria, Turkey
and into Neolithic Europe (Kreuz and Boenke, 2002). It
persists in parts of Europe as late as the Iron Age, after
1000 BC (Kreuz and Boenke, 2002; Ko ¨hler-Schneider,
2003), and disappears in its Syrian homeland during the
Chalcolithic, approx. 5000 BC (Van Zeist, 1999; Kreuz
and Boenke, 2002). This implies an additional two-grained
einkorn domestication but this crop went extinct in
prehistory. Similarly, there is now evidence for an extinct
emmer-like wheat, with distinctive glume architecture not
yet paralleled in any studied modern wild or cultivated
landrace but with some similarities to Triticum timopheevi
(Jones et al., 2000; Ko ¨hler-Schneider, 2003). It is known
archaeologically from the eastern half of Europe, Turkey
and Turkmenistan. It persists in parts of Europe as late as
the Bronze Age (approx. 1000 BC) (Ko ¨hler-Schneider,
2003). In addition, early sites in Syria appear to have
cultivated a local form of rye (Secale cf. montanum), but
rye did not become a major crop of the Neolithic Near
East despite occasional later finds (Hillman, 2000: 392),
and was probably a different species from the later
European rye (Secale cereale), domesticated from a field
weed in Late Bronze Age or Iron Age times (approx.
1000 BC) (Ku ¨ster, 2000). Taken together the archaeobota-
nical morphotypes and genetics suggest a minimum of
seven domestications of wheat and barley in the Near
Eastern Fertile Crescent region, and there is no reason to
attribute them all to a single micro-region or a single
process of agricultural origins, but at least two or perhaps
three (Willcox, 2005).
Cereal grain size increase
There is a growing morphometric database for wheat and
barley from the Near East (Colledge, 2001, 2004; Willcox,
2004). This indicates that wheat and barley grains increased
in size starting in the PPNA and earliest PPNB. This is
before clear and widespread evidence for tough rachises
and loss of natural seed dispersal. It is well known that
wild and domesticated cereal grains differ in size and this
has been used to infer the domesticated status of cereals,
already in the PPNA and the earliest PPNB, including
sites from the Jordan Valley (e.g. Fig. 3), the upper
Euphrates in Syria, and the first settlements on Cyprus
(Colledge, 2001, 2004).
This evolutionary shift can be illustrated from internal
archaeobotanical evidence too, i.e. by charting grain
metrics through time, as recently demonstrated with
samples from the site of Jerf el Ahmar on the Upper
Euphrates (Willcox, 2004). Fig. 4A shows the contrast
between the barley grains from the early phase at Jerf el
Ahmar (9500–8800 BC) and the much later Chalcolithic
site of Kosak Shamali (approx. 5000 BC), in which all of
the grains are larger and comparable with the domesticated
size range inferred from modern material. If we look at the
later phase at Jerf el Ahmar (approx. 8500 BC), however, it
can be seen that many of the grains are of the larger size
(Fig. 4B). This implies evolution towards larger grain size
during the occupation of this site, but recovered rachis
remains indicate that ears were still of the wild, shattering
type. A similar pattern is found for the einkorn wheat
grains (Triticum monococcum), which also include some
mixture of rye (Secale cereale) (Fig. 4C, D). Given that
occupation of Jerf el Ahmar lasted less than a millennium
and perhaps only 500 years, we can suggest the rate of evol-
ution of large grains, as occurring in a few centuries and
certainly more rapidly than a millennium. This explains
why large domestic-type grains are already widespread
and predominant on most Near Eastern sites by the start
of the PPNB (approx. 8700 BC). From the even earlier
grain assemblage from Abu Hureyra, where cultivation
has been inferred from the arable weed flora, a few large
and plump rye grains were found, comparable with domesti-
cated grains (Hillman, 2000; Hillman et al., 2001). These
might represent the earliest indications of selection for
grain size increase under cultivation, but possibly a local
This evidence raises the question of how large-grained
varieties evolved. One possibility is that methods of
FIG. 3. Pre-Pottery Neolithic B wheat grain measurements from the
Jordan Valley (after Colledge, 2001). This indicates the predominance of
the larger domesticated-type grains. The gap between the two groups is
comparable with that in modern reference material. These sites are domi-
nated by wild-type barley rachis (see below).
Fuller — Patterns in Crop Domestication 908
processing, such as using sieves after threshing and win-
nowing, served to bias stored cereals towards larger grains
(on traditional crop-processing and the use of sieves, see
Hillman, 1984). This, however, would tend to remove
‘tail grain’, i.e. the smaller grains on an ear, and would
be expected to make very weak selection for genetically
larger-grained plants, as the selected grains would be
from the same genetic population as the discarded small
grains. More likely seems to be the role of tillage. The prac-
tice of preparing land with tillage is likely to have been
necessary to remove competing vegetation from favourable
micro-environments for early cultivation, such as the breaks
of slopes where ground-water tables are higher. Such areas
would naturally be cloaked in herbaceous flora and under-
shrubs such as Tamarix, so tillage would be necessary
(Hillman, 2000: 396; Hillman et al., 2001: 387). Sowing
grains into tilled soil with an underlying water table would
give advantages to those best able to germinate successfully
from greater burial, namely larger-seeded individuals.
Evolution of the tough rachis
The evolution of non-shattering ears was also a gradual
process. Although theoretically it could have happened
very quickly, as demonstrated under ideal experimental
conditions (Hillman and Davies, 1990), the archaeobotanical
evidence indicates a much more gradual evolution of non-
shattering ears. A recent quantitative assessment of the
wheat and barley rachis remains from a few sites
suggested a gradual increase in the proportion of the
domesticated-type ears over the course of the PPNB
(Tanno and Willcox, 2006a). A larger data set has been
compiled here in which evidence for einkorn wheat
spikelet bases and barley rachis segments are considered
(Fig. 5A, B). In general, there is contrast between early
sites which are largely or entirely of the wild-type chaff
remains and later sites dominated by domesticated-type
remains, with some intermediate proportions for sites
chronologically in the middle. Although there are indi-
vidual site exceptions, such as barley rachis from Jordan
valley sites (e.g. Wadi Jilat), these may relate to particular
local circumstances of higher reliance on wild plant foods
(indicated by other data from this site, Colledge, 2001). In
addition, wild cereals, especially barley, may persist as
weeds of cultivation. Nevertheless, at a regional level the
overall trend is clear.
These data can then be transformed to produce a
regression of domestication rate over time (Fig. 6A, B). In
these diagrams individual site proportions are plotted,
both by the minimal figure, based on just the clear non-
FIG. 4. Scatter-plots of archaeological grain measurements showing the increase in grain size under early pre-domestication cultivation (after Willcox,
2004). (A) Barley grain measurements, comparing early Pre-Pottery Neolithic A Jerf el Ahamr with the much later domesticated material from Kosak
Shimali. (B) Comparing early and late Jerf el Ahmar, indicating that shift towards larger grain size had already occurred. (C) Similar comparison of
einkorn grains (probably including some rye grains) at early Jerf el Ahmar and Kosak Shimali. (D) Trend towards larger grain sizes over the course
of Jerf el Ahmar occupation.
Fuller — Patterns in Crop Domestication909
dehiscent type, and a larger figure which includes uncertain
but possible domesticates. The true proportion of domesti-
cates should fall within this range. Sites are plotted
against a median age estimate for each site. In addition,
for barley the averages from each phase (e.g. PPNA,
Early PPNB, Late PPNB) are plotted. The trend from
wild dominance to domesticated dominance is clear but
appears fairly gradual and slow. The rates of evolution do
FIG. 5. Proportions of wild and domesticated barley and einkorn rachis/spikelet remains on early Near Eastern sites, arranged chronologically from left to
right, and grouped into broader phases. (A) Barley rachis types, including domesticated (tough), wild (shattering) and uncertain (but more likely wild). (B)
Proportions of einkorn spikelet forks/glume bases, including domesticated (tough), wild (shattering) and uncertain (but more likely domesticated). Sites,
approximate ages and data sources: Ohalo 2, 21 000–18 500 BC (Kislev et al., 1992); Wadi Hammeh, approx. 12 000 BC (Colledge, 2001); Mureybit,
10 500–9500 BC (Van Zeist and Bakker Heeres, 1986); Iraq-ed-Dubb, approx. 9300 BC (Colledge, 2001); Jerf el Ahmar (early, with two-grained
einkorn), 9700–9300 BC (Willcox, 1999, 2002); Wadi Jilat 7, 8800–8300 BC, Wadi Jilat 13, 7000–6500 BC (Colledge, 2001); Aswad, 8700–8000
BC (Van Zeist and Bakker Heeres, 1985; Tanno and Willcox, 2006a); Azraq 31, 7500–7000 BC (Colledge, 2001); Wadi Fidan A, 7500–7000 BC,
Wadi Fidan C, 7000–6500 BC (Colledge, 2001); El Kowm, 7500–6800 BC (De Moulins, 1997); Catal Hoyuk, 7400–6800 BC (Fairbairn et al.,
2002); Ramad, 7500–6500 BC (Van Zeist and Bakker Heeres, 1985; Tanno and Willcox, 2006a); Magzaliyeh, 7100–6400 BC (Willcox, 2006); Tell
el Kherkh, 8600–8300 BC (Tanno and Willcox 2006a, b); Nevali Cori, 8500–8000 BC (Tanno and Willcox, 2006a); Qaramel, approx. 10000 BC
(Tanno and Willcox, 2006a); Netiv Hagdud, 9500–9000 BC (Kislev, 1997); Cafer Hoyuk, (IX–XIII) 8300–7700 BC, (III–VIII) 7500–7000 BC (De
Moulins, 1997); Kosak Shamali, approx. 5000 BC (Tanno and Willcox, 2006a).
Fuller — Patterns in Crop Domestication910
not come anywhere near the 20–100 years estimated by
Hillman and Davies (1990) on the assumption of sickle har-
vesting morphologically wild near-mature plants, or uproot-
ing of whole plants. This vast difference in rate raises
questions about how we explain the selection for domesti-
cated non-shattering genotypes, an issue to which I return
From these data we can also suggest that domestication
occurred somewhat more quickly in wheat, perhaps
around 1500 years, as opposed to barley which shifts over
a period of 2000 years or slightly more. An earlier domes-
tication date for wheat has been previously noted (e.g.
McCorriston, 2000). A number of factors might be
suggested to account for this. Slightly higher average rates
of cross-pollination are reported in barley, estimated at
about 1–2 % (Allard, 1988; Morrell et al., 2005), as
opposed to wheats, which are less than 1 % (Hillman and
Davies, 1990: 62), although this seems unlikely to be suffi-
cient in itself. Of importance may be the fact that wild
barley (Hordeum spontaneum) is more prone to be a weed
of cultivation, thereby maintaining introgression, whereas
wheat cultivation, which is more water-demanding and
may have been more carefully located, would have been
cut off from the wild progenitor. In addition, the persistence
of wild barley as a weed will affect the composition of
archaeobotanical assemblages and rachis counts may there-
fore be an underestimate of the proportion of domesticated
types among crop populations.
Most remarkable is the difference in evolutionary rate by
comparison with grain size (semi-domestication), of
500–1000 years. The shift in rachises (full domestication)
appears to start mainly after large grain size had already
begun to evolve, and we might therefore suggest a
minimum estimate of 2000 years for the evolution of both
aspects of the domestication syndrome.
CHINESE RICE DOMESTICATION: THE
IMPACT OF UNEVEN RIPENING
Chinese agricultural origins: background
Systematic archaeobotany in China has only recently begun
and evidence for the beginnings of agriculture is still
limited (Fig. 7). In northern China the millets Setaria
italica and Panicum miliaceum were the initial crops. The
earliest well-documented millets are from approx. 6000
BC at Xinglonggou, in Eastern Inner Mongolia, by which
time plump-grained Setaria were already established but
grains of P. miliaceum were near the wild-type in size
and shape (Zhao, 2005). Subsequently, millet cultivation
was established in much of the Yellow River basin by
5500 BC (the Peiligang, Cishan, Beixin and Dadiwan cul-
tures) (Lu, 1999; Crawford, 2005; Crawford et al., 2005).
Rice from the south was added to this agricultural system
only in the third millennium BC, with a few rice finds
from Late Yangshao contexts (3000–2500 BC) and many
more from the Longshan period (2500–2000 BC). In
recent years, the orthodoxy has been that rice agriculture
began early, perhaps at the start of the Holocene or late
Pleistocene, in the Middle Yangtze, perhaps amongst sea-
sonally inhabited cave-sites (e.g. Lu, 1999; Crawford,
2005). Critical re-assessment, however, suggests that rice
may have been independently domesticated in both the
middle and the lower Yangzte river areas, and that domes-
tication was later than has been presumed (Fuller et al.,
Domestication as morphometric and maturity shift
It is now well established that Oryza sativa was domesti-
cated more than once, with distinct origins of tropical mon-
soonal indica and marshland, sub-tropical japonica.
Genetic studies are now numerous in support of these sep-
arate origins (e.g. Chen et al., 1993; Cheng et al., 2003;
Londo et al., 2006). Archaeological evidence also favours
distinct centres of early rice cultivation in the Middle
Ganges valley, India (Fuller, 2002, 2006), and the Middle
or Lower Yangtze, China (Lu, 1999). This situation falsifies
the older suggestions regarding Chinese Neolithic rice as
FIG. 6. Domestication rates in barley and einkorn modelled from archae-
obotanical data (based on Fig. 5). Proportion of domesticated type for each
site is plotted by a box against a median estimate of site age. A margin of
error is indicated by the line which connects the sum of domesticated and
uncertain types (indicated by a cross or x). Trend lines are shown based on
the lower estimate. (A) Barley domestication rate model, on which period
averages are also plotted for the PPNA, Early PPNB and Late PPNB, in
which the diamond indicates the proportion of domesticated types and
the circle the sum of domesticated and uncertain types. (B) Einkorn dom-
estication rate model; the much later Kosak Shamali has been excluded.
Fuller — Patterns in Crop Domestication911
predominantly indica, or an ancient ‘intermediate type’ that
evolved into both indica and japonica. In light of this, I and
collaborators have reconsidered published sources on early
Chinese rice, and begun new archaeobotanical investi-
gations (Fuller et al., 2007). Similar research has been
recently pursued on Indian rice (Harvey, 2006). Previous
studies essentially began with the assumption that archaeo-
logical rice was a crop, and only asked instead whether it
was indica, japonica or something ‘intermediate’ (e.g.
Oka, 1988; Zhang, 2002). In fact, the morphometric data
of early rice often fit well within the range of wild
species, including Oryza rufipogon but also sometimes
Oryza spp. not in the sativa-complex. On this basis we
have suggested that another wild species might sometimes
have been utilized as a hunter-gatherer resource (Fig. 8A,
O. officinalis might be suggested (cf. Vaughan, 1994),
and this species produces substantial grain numbers per
plant, although further consideration of feasibility must
take into account potential stand sizes (generally small),
as well as potential processing techniques that would be
required for dehusking this thick-hulled species. Other
Oryza species or a particularly small-grained race of
O. rufipogon should also be considered.
Much of the grain, however, is notably long and skinny,
which could also be a trait of immature rice (Fig. 8B).
Consideration of rice development, with its extended
period of unevenripening
Hoshikawa, 1993), and expectations from optimal foraging
(more than2 weeks,
theory and ethnographic expectations about harvesting lead
us to expect early users of wild rices to have targeted imma-
ture plants to maximize their recovery of near-mature
grains. Indeed in order to maximize grain yields wild rice
should be targeted in the first 3 or 4 d after the very first
grains mature, but at this time slightly more than 40 % of
the grains are still immature by more than 6 d. Within a
week potential yields are vastly reduced by something on
the order of 70–80 % of mature grains (Fig. 9). As with eth-
nographically documented hunter-gatherers, we would
therefore expect wild rice to be harvested when immature
(e.g. the Bagundji wild Panicum gatherers of Australia:
Allen, 1974; cf. Harris, 1984).
Archaeologically, the morphometric data available from
Kuahuqiao and Longqiuzhuang (mainly of the earlier
Majiabang period, which equates with the Hemudu
period) suggests that grain assemblages are dominated by
immature grains (Fig. 8B). It must be noted that this
assumes the mature grains would have been in the range
of modern domesticates, but they do appear somewhat
plumper than expected for mature O. rufipogon. This in
fact is plausible if we consider the likelihood that early pre-
domestication cultivation had already begun to select for
larger grains. It must be noted, however, that grain size
evolution in rice has been complex and not always
towards larger grain sizes. Many short-grained japonica
races may have shorter, but often still plumper, grains
than those in the wild progenitor. It is, nevertheless, clear
that grain shape has changed under domestication and
FIG. 7. Map of East Asia indicating early millet and rice sites, with inset of Yangtze region, showing archaeological sites mentioned in this article: 1,
Hemudu; 2, Tianluoshan; 3, Kuahuqiao; 4, Shangshan; 5, Liangzhu; 6, Majiabang area, including Nanzhuangqiao, Luojiajiao and Pu’anqiao; 7,
Nanhebang; 8, Maqiao; 9, Songze; 10, Xujiawan; 11, Chuodun; 12, Weidun; 13, Longnan and Caoxieshan; 14, Qiucheng; 15, Longqiuzhuang; 16,
Sanxingcun; 17, Lingjiatan; 18, Jiahu; 19, Bashidang; 20, Pengtoushan.
Fuller — Patterns in Crop Domestication912
may also be affected by the proportions of mature and
immature grains represented in a harvest.
In addition, the reduction in hairs on the awns of rice
recovered from Hemudu (Sato, 2002) implies relaxation
of natural selection for seed dispersal aids, which would
be expected under cultivation. The Hemudu culture is
rightly famous for it numerous wood and bone spades,
tools which imply soil manipulation and probably planting.
This is in contrast to Kuahuqiao, which produced just two
poorly made proto-type spades. The slight difference in
grain thickness between Kuahuqiao (6000–5400 BC) and
the Majiabang period Longqiuzhuang (5000–4000 BC)
could therefore suggest selection for larger grains under
early cultivation, and we might therefore suggest the begin-
nings of cultivation between the end of the Kuahuqiao
phase (approx. 5400 BC)and the earlyHemudu/
Majaiabang phase (from 5000 BC). This would imply a
rate of evolution on a par with the grain-size increase in
wheat and barley. Thus, the evidence from Hemudu and
the earlier Majiabang period (from early to mid fifth mil-
lennium BC) both suggest pre-domestication cultivation,
which may have started already at the end of or after the
Kuahuqiao period. The rice at this stage can be regarded
as semi-domesticated, like that of PPNA to early PPNB
wheat and barley.
A clear contrast is seen with assemblages from around
4000 BC. The latest assemblage from Longqiuzhuang
(early Songze period, just after approx. 4000 BC) has
grains which are significantly longer, plumper (2.5–3 mm)
and most likely fully mature. In addition, signifi-
cantly plumper grains have been recovered from Chuodun
(late Majiabang, just beforeapprox. 4000 BC). This suggests
FIG. 8. (A) Scatter plot of length and width of grains measured in modern populations (15 grains measured from 72 populations; Fuller et al., 2007). (B)
Grain measurements from selected Neolithic sites, showing maximum and minimum measured ranges with solid lines and statistical standard deviations
with dashed lines (as reported). Note that grains from Kuahuqiao, Bashidang and the lower (Majiabang period) levels (8–6) at Longqiuzhuang fall largely
or entirely in the expected immature grain proportions, while the latest grains from Longqiuzhuang, Songze period (level 4), indicate a clear shift towards
longer and fatter grains that can be regarded as fully mature, and thus domesticated. Chouden (Late Majiabang) also indicates a shift towards mature
japonica-type grains, but suggests local population differences from the domesticated rice at Longqiuzhuang. The small grains from Jiahu are suggestive
of wild rice not in the sativa complex, such as O. officinalis. Sources: Kuahuqiao (Zheng et al., 2004b), Longqiuzhuang (Huang and Zhang, 2000), Jiahu
(Henan Provincial Institute of Archaeology, 1999), Chuodun (Tang, 2003) and Middle Yangzte Bashidang (Pei, 1998).
Fuller — Patterns in Crop Domestication913
an important morphological shift in archaeological rice
occurred in the Lower Yangtze region during the later fifth
millennium BC. This shift seems most likely to be due to
a shift towards harvesting of mature panicles as opposed
to immature panicles, rather than an evolutionary develop-
ment in grain shape. Such a shift would imply that it
became feasible to allow grains to mature on the plant
without loss of the grains, or in other words that
tough, domesticated-type rachises had evolved to dominate
the rice populations being harvested. This hypothesis is
currently being tested by myself and collaborators on large
assemblages of preserved rice spikelet bases from the fifth
millennium BC site of Tian Luo Shan. Additional
evidence comes from measurements on bulliform phyto-
liths, silica bodies from the leaves of rice (Fig. 10). These
Majiabang and Songze periods, i.e. before and after 4000
BC. Modern studies suggest that the size of these phytoliths
relates to plant maturity and tillering stage (Zheng et al.,
These data suggest a rate of evolution from the beginning
of cultivation to full domestication on the order of 1000–
1500 years, similar in magnitude but slightly faster than
that in the Near East. An important factor in this is that
Late Majiabang period sites are known that preserve early
paddy field systems. These consist of networks of shallow
largersizes between the
pools, and ditches, as at Caioxieshan and Chuodun (Zou
et al., 2000; Gu, 2003). This development in cultivation
techniques would imply greater separation of sown popu-
lations from wild stands, and a reduction therefore in cross-
pollination with free-growing wild rice, which has been
estimated at 10–50% (Oka and Morishima, 1967). This
separation into distinct field systems may be highly signifi-
cant, as, theoretically, out-breeding species should evolve
more slowly towards adaptations to cultivation (Allard,
Also important is evidence for climatic change, which is
likely to have been reducing populations of wild rice.
Globally the mid Holocene, after 6000 BC, and especially
between 5000 and 3000 BC, was a period of cooling, as
FIG. 9. (A) Graph indicating the expected relative frequency of grains
reaching maturity during eight stages, of 2 d each, on an individual rice
plant (based on anthesis data of a modern japonica cultivar, from
Hoshikawa, 1993). (B) A graph converting this data into potential grain
yields to a hunter-gather if this were a morphologically wild plant. This
indicates the proportion of grains more than 6 d immature which would
differ in grain proportions from mature grains. Although this represents
an individual plant it must be assumed that a population of wild rice has
individuals that begin this process at different times over a period of
weeks. With cultivation there should be a tendency for plants to become
FIG. 10. Size increase in Lower Yangzi rice phytoliths. (A) Measured
horizontal length and vertical length of rice bulliform phytoliths from
Majiabang period samples (M); (B) measurements from samples of the
subsequent Songze (S) and Liangzhu (L) phases. The dashed oval rep-
resents the distribution of the Majiabang measurements. Data re-plotted
from Zheng et al. (1994, 2004a, b) and Wang and Ding (2000).
Fuller — Patterns in Crop Domestication914
indicated for example by declining methane levels from
Greenland ice cores (Blunier et al., 1995). This is linked
to declines in monsoon rainfall in East Africa (Gasse,
2000), South Asia (Madella and Fuller, 2006) and East
Asia (Wang et al., 1999). This process would be expected
to have impacted South China, pushing more tropical taxa
such as wild rice further south. Also significant is pollen
core evidence from the Lower Yangzte region that indicates
a marked decline in broad-leaved trees, including oaks,
after 5200 BC (Yu et al., 2000; Yi et al., 2003; Tao
et al., 2006).
This is significant because evidence from Kuahuqiao,
Hemudu and Tian Luo Shan suggests that it was acorns
(Quercus, Lithocarpus, Cyclobalanopsis) that were the
dietary staples of these cultures. These were stored in vast
quantities in storage pits, along with water chestnuts
(Trapa bispinosa) (Zhejiang
Archaeology, 2003; 2004; my unpubl. data). As compara-
tive cases indicate, e.g. ethnographic California or the
archaeology of the Jomon culture of Japan, tree nuts, and
particularly acorns, make ideal staple food sources (e.g.
Heizer and Elsasser, 1980: 82–144; Takahashi and
Hosoya, 2002; Barlow and Heck, 2002). Therefore, the
decline in nut-bearing trees may have promoted increased
reliance on, and cultivation of, rice (Fuller et al., 2007).
These climatic factors may also explain the apparent
difference in the domestication rate between Near Eastern
cereals and East Asian rice. In the Early Holocene, the
climate was warmer and wetter and therefore conducive to
the expansion and persistence of wild wheat and barley
stands in the Near East. By contrast, in Mid-Holocene
China wild rice would have been in decline and on its
way to local extinction. This may have promoted a some-
what faster rate of domestication.
THE PULSE DOMESTICATION MODEL: SEED
SIZE AS EPIPHENOMENON
The domestication syndrome for pulses is essentially the
same as that for cereals, with increases in seed size,
reduced pod-shattering and importantly loss of germination
inhibition (Plitman and Kislev, 1989; Smartt, 1990; Zohary
and Hopf, 2000). This need not imply, however, that the
rates of domestication and the order in which these traits
evolved are necessarily the same. In fact, evidence suggests
a difference from cereals. This can be illustrated with
evidence for the early evolution of the Indian mungbean
(Vigna radiata) and urdbean (Vigna mungo) under
South Asia has received less attention from archaeolo-
gists studying early agriculture, although a large number
of minor cereals (millets), pulses, cucurbitaceae crops and
indica-type rices were domesticated there. Archaeology
together with the modern distributions of wild progenitors
suggest as least three, and perhaps five, distinct centres of
plant domestication in India, including South India,
Gujarat and the Ganges Plain (for a comprehensive
review, see Fuller, 2006; also Fuller, 2002). These
mid-Holocene, between 4000 and 2000 BC, and occurred
amongst local populations of hunter-gatherers in monsoonal
India, although some influences from the west due to the
spread of livestock cannot be ruled out. In the north-west,
early agriculture was based on the dispersal of wheat,
barley and winter pulse cultivation from the Near East,
together with domesticated animals. Despite the long
delay before agriculture developed in other parts of
India the earliest phases suggest local packages of crops
into which wheat, barley or Near Eastern pulses were
The current distribution of wild mung and urd, taken
together with their archaeological record, suggests three
areas of domestication (Fig. 11). Vigna radiata was
probably brought under cultivation in the north-west,
perhaps in the Himalayan foothills in the Punjab region,
and in the far south, as early finds in both regions are
widely separated. Early finds of urdbean come from
Gujarat and the northern Peninsula where wild populations
of this species persist (Vigna mungo var. sylvestris) without
associated wild mungbean (Vigna radiata var. sublobata
sensu stricto) (Fullerand
Tomooka et al., 2003). Early finds in South India of
V. radiata occur in the driest savannah environments of
the peninsula together with large quantities of horsegram
(Macrotyloma uniflorum) and small millets (Brachiaria
ramosa, Setaria verticillata), and in some cases associated
with introduced domesticates (Fuller et al., 2004). The
association with other crops, their large quantity and the
difference from the habitat of the wild progenitor all
imply cultivation of these early mungbeans (on the past
environmental conditions at these sites, see Fuller and
These early Neolithic pulses, however, do not appear to
be different in size from their wild progenitor (Fig. 12A,
B). Even assuming the upper end of 20 % shrinkage,
measured specimens from southern Neolithic sites fall
almost entirely within the size range represented by
modern wild populations (Fuller and Harvey, 2006). By
contrast, those from Early Historic to Early Medieval
periods fall in the range of modern domesticated popu-
lations (Fig. 13A). These data suggest on peninsular India
selection for increasing pulse seed size was slight through
the second millennium BC but had occurred by the first mil-
lennium AD. The evidence from late Chalcolithic Tuljapur
Garhi (late second millennium BC to early first millennium
BC) shows a wide size
domesticated-type, although the average falls in the latter
range. This site might represent evidence for the actual
process of size increase, suggesting that this occurred
most markedly during the late second millennium BC
through the Iron Age.
These data raise the question of what change in the
environmental conditions, most likely in terms of agricul-
tural practices, selected for seed enlargement in these
pulses. Presumably domestication in terms of dispersal
and germination was selected for early and rapidly, necess-
ary to make any pulse a worthwhile crop (Ladzinsky, 1987,
1993; Zohary, 1989; Blumler, 1991). If it was being routi-
nely harvested, either by pod-plucking or plant uprooting,
and sown from stores we would expect domesticated
Harvey,2006; see also
range from wild-type to
Fuller — Patterns in Crop Domestication 915
forms to have evolved with less dehiscent pods and loss of
dormancy. But size increase was not part of the initial dom-
estication syndrome. What explains the delay? I would
suggest that changes in agricultural techniques, such as
deeper tillage (with ards), created a selective advantage
for larger seeded genotypes (Fuller and Harvey, 2006:
257). In this regard the shift during the Iron Age (first mil-
lennium BC) is significant as this is the period when ard
tillage began on the Indian peninsula, and perhaps in the
latest second millennium BC (cf. Shinde, 1987; Fuller
et al., 2001).
Measurements from northern India suggest a similar
pattern in the middle Ganges and Orissa, but suggest a
contrast with early pulses from the north-west (Fig. 13D).
What can be seen in the plot is that Neolithic seeds fall
largely in the expected primitive size range, while later
sites such as late Chalcolithic Narhan and later Iron Age
Interestingly, two early sites, Balu and Kunal from the
later third millennium BC, also have large Vigna seeds.
Both of these sites, however, are within the Eastern Indus
Civilization cultural zone, where we expect deep tillage
with ards to have been the norm (Allchin and Allchin,
1997: 170; Miller, 2003). It might be suggested that in
the Indus zone enlarged seeds have already been selected
for by tillage in the third millennium, whereas further
havelarge Vigna grains.
FIG. 11. A map of the wild progenitors of Vigna radiata and V. mungo in India in relation to the moist deciduous forests and the region with extensive
wild rice populations (based on Tomooka et al., 2003; Fuller and Harvey, 2006). Archaeobotanical finds of the Vigna pulses are indicated which include
secure species-level identifications. Sites are numbered: 1, Semthan; 2, Hund; 3, Balu; 4, Kunal; 5, Burthana Tigrana; 6, Mitithal; 7, Hulas; 8, Hulaskhera;
9, Charda; 10, Imlidh-Kurd; 11, Narhan; 12, Khairadih; 13, Malhar; 14, Senuwar; 15, Tokwa; 16, Mahagara; 17, Koldihwa; 18, Balathal; 19, Babar Kot;
20, Rojdi (two phases); 21, Oriyo Timbo; 22, Kaothe; 23, Tuljapur Garhi; 24, Paithan; 25, Apegaon; 26, Bhokardan; 27, Nevasa; 28, Inamgaon; 29, Terr;
30, Golabai Sassan; 31, Piklihal; 32, Hallur; 33, Tekkalakota, Kurugodu, Sanganakallu and Hiregudda; 34, Hattibelagallu; 35, Sanyasula Cave; 36,
Veerapuram; 37, Rupanagudi; 38, Ramapuram, Hanumantaraopeta and Peddamudiyam; 39, Kodumanal; 40, Perur. For details of primary sources, see
Fuller and Harvey (2006).
Fuller — Patterns in Crop Domestication916
east this process did no occur until the late second millen-
nium BC. This of course raises the questions of whether the
Vigna pulses introduced into the middle Ganges zone had
diffused before emergence of large-seeded forms, or had
diffused from the south rather than the Harappan zone, or
whether the absence of positive selection through tillage
could have led to a reversion to smaller seed size.
This model provides an explanation for the archaeologi-
cal difficulty in identifying the beginnings of pulse domes-
tication. Pods are not normally preserved, and germination
characters are cryptic in rarely preserved seed coats (Butler,
1989). Only in peas does a clear distinction exist between
smooth-testa domesticates (Pisum sativum spp. sativum)
and tuberculate-testa wild peas (Pisum sativum ssp.
elatius) (Butler, 1989; Zohary and Hopf, 2000). Early
finds of pulses co-occur with cereal crops and weed assem-
blages, suggesting that they too were crops, but no measur-
able change in seed size is noted (Garrard, 2000; Zohary
and Hopf, 2000; Hillman et al., 2001; Tanno and
Willcox, 2006b; Weiss et al., 2006). Even millennia after
the beginnings of cereal cultivation and the establishment
of domesticated cereals, when agriculture had dispersed to
Europe, finds of peas and lentils (e.g. from Neolithic and
Early Brone Age Greece, Bulgaria, Germany and the
Netherlands) are little different from their pre-agricultural
Near Eastern ancestors in terms of average and minimum
sizes, although there is a slight tendency for larger size
maxima (e.g. Van Zeist and Bottema, 1971; Bakels, 1978;
Hopf, 1973; Housely, 1981; Hansen, 1991). Perhaps the
earliest sites with notably large lentils and peas are some
Near Eastern pottery Neolithic sites (approx. 6000–5000
BC), such as Erbaba in Turkey and Tepe Sabz in Iran
(Helbaek, 1969; Van Zeist and Buitenhuis, 1983), but
such sites are the minority, with consistently enlarged
pulse seeds encountered only in the Late Bronze Age,
Iron Age and Roman periods. Taken together this suggests
that there was an even longer delay in serious size increase
in Near Eastern pulses than in Indian Vigna, something on
the order 2000–4000 years, or even longer in some regions.
It also raises the possibility that some size increases in par-
ticular regions and periods were contingent on local circum-
stances of selection. It may also be noted that limited data
on early East Asian soybeans (Glycine max) and adzuki
beans (Vigna angularis) indicate that as late as the third
millennium BC they are still largely within the wild size
range, with enlarged seed sizes from the later Bronze Age
FIG. 12. Metrical data for Indian Vigna pulses indicating no size increase
with domestication. (A) Modern length–width measurements in Vigna
radiata, V. mungo and their wild progenitors. The separation between
them has been adjusted for a high estimate of 20% shrinkage with carbon-
ization and shown on the other plots as a dashed line. (B) Neolithic Vigna
finds from South India (before 1400 BC) (data from Fuller and Harvey.
FIG. 13. (A) Late Chalcolithic (1400–900 BC), Iron Age (900–200 BC)
and Early Historic (200 BC – 400 AD) Vigna finds from South India, indi-
cating size increase. (B) Archaeological Vigna measurements from north-
ern India, including Ganges valley Neolithic, Iron Age and Harappan
Bronze Age civilization finds, suggesting a correlation between larger
size and plough agriculture (data from Fuller and Harvey, 2006).
Fuller — Patterns in Crop Domestication 917
or Iron Age (Crawford and Lee, 2003; Crawford et al.,
2005), periods in which ploughs were established.
A PULSE-LIKE CEREAL? THE CASE OF
AFRICAN PEARL MILLET
A final interesting case is provided by pearl millet
(Pennisetum glaucum), an important domesticate of West
Africa (D’Andrea and Casey, 2002; Fuller, 2003; Zach
and Klee, 2003). Archaeobotanical research in Africa
remains very limited especially given the size of the conti-
nent. Most studies have focused on Egypt, but there is a
sub-Saharan West Africa. As outlined by Harlan (1971)
there are a wide range of crops with varying wild progenitor
distributions in Africa (Fig. 14). In terms of major cereal
domestications the northern savannahs and the Sahel zone
south of the Sahara are important. Sorghum bicolor ssp.
bicolor is likely to have originated from the eastern part
of the savannahs, such as between Lake Chad and the
Western Sudan, or north-west Ethiopia (Harlan and
Stemler, 1976). By contrast, pearl millet has been linked
to one, or more likely two, domestications in the Sahel
zone west of Lake Chad (Tostain, 1992; Fuller, 2003;
Neumann, 2003). An important factor in where and when
agriculture developed in Africa is climate change. As
wetter conditions of the early (10 000–6200 BC) and mid
Holocene (5900–2200 BC) caused a northward shift in
vegetation zones and much of the Sahara was savanna-like.
This environment supported populations of pottery-making
hunter-gatherers, who utilized a range of wild millet-
grasses, including wild Sorghum bicolor in the east
(Egypt’s Western Desert), and these groups adopted live-
stock early, certainly by 6000–5000 BC (Marshall and
Hildebrand, 2002; Fuller, 2005). As conditions dried, after
3500 BC, and populations with their herds were forced
south, some appear to have taken up cultivation. Evidence
for the actual transition to agriculture remains elusive but
in the early second millennium BC widely dispersed popu-
lations in West Africa were cultivating morphologically
domesticated pearl millet (D’Andrea and Casey, 2002).
That pearl millet had dispersed to India by approx. 1700
BC (Fuller, 2003) suggests that this process began by the
third millennium BC and saw rapid dispersal of cropping
across the northern savannas.
Pearl millet domestication is inferred from two sets of
evidence. First, there was loss of natural seed shedding,
which is linked to the shift from sessile involucres to devel-
opment of a non-dehiscent peduncle (Poncet et al., 2000).
This shift is already evident from ceramic impressions of
pearl millet chaff by 1700–1500 BC in Mauretania
(Amblard and Pernes, 1989; MacDonald et al., 2003), and
FIG. 14. A synoptic geography of early agricultural developments and precursors in Africa. Shown are the modern distributions of wild Sorghum bicolor
and Pennisetum glaucum with genetic connections to the domesticates (after Harlan, 1971; Tostian, 1992). The previously wetter conditions imply a
northward shift in the Sahara–Sahel transition (see Gasse, 2000; Marshall and Hildebrand, 2002). Early Holocene ceramic-using forager sites based
on Jesse (2003), and mid-Holocene pastoral sites based on Jousse (2004). Early evidence for wild sorghum gathering is indicated (based on Stemler,
1990; Barakat and Fahmy, 1999; Wasylikowa and Dahlberg, 1999). The spread of Near Eastern crops is indicated in the Nile Valley vis-a `-vis the pre-
ceramic Neolithic distribution in the Eastern Mediterranean. Sites with early pearl millet are numbered: 1, Dhar Tichitt sites (cited in D’Andrea and
Casey, 2002); 2, Dhar Oualata sites (Amblard and Pernes, 1989); 3, Djiganyai (MacDonald et al., 2003); 4, Winde Koroji (MacDonald, 1996); 5,
Karkarichinkat (cited in D’Andrea and Casey, 2002); 6, Ti-n-Akof (cited in D’Andrea and Casey, 2002); 7, Oursi (cited in D’Andrea and Casey,
2002); 8, Birimi (D’Andrea et al., 2001); 9, Ganjigana (Klee et al., 2004); 10, Kursakata (Zach and Klee, 2003). Historical sites with pearl millet metrical
data: 11, Arondo (cited in Zach and Klee, 2003); 12, Jarma (Pelling, 2005); 13, Qasr Ibrim (Steele and Bunting, 1982).
Fuller — Patterns in Crop Domestication 918
slightly later in Nigeria (Klee et al., 2000, 2004). Pearl
millet shows a subtle but clear change in grain shape,
becoming apically thicker and more club-shaped than its
wild counterpart, i.e. an increased thickness/breadth ratio
(Brunken et al., 1977; D’Andrea et al., 2001; Zach and
Klee, 2003). However, a major increase in seed size
appears delayed (D’Andrea et al., 2001: 346). Figure 15A
shows the expected range of domesticated and wild forms
based on reported modern populations, adjusted for 10 %
Of note is that early West Africa populations, from the
second and first millennia BC, have their averages firmly
in the wild size range, although there are long tails of vari-
ation that extend into the larger size range (e.g. at Birimi).
One of the earliest finds of pearl millet from India comes
from Surkotada, Gujarat, approx. 1700 BC, which can be
seen to fall with these early domesticated African popu-
lations. By contrast, a rather later, Gangetic population
from Narhan (where ard tillage is established, and where
larger Vigna pulses were present, see Fig. 13D) is
markedly larger, suggesting selection for larger-grained
pearl millet. In Africa larger grained populations appear
in the first millennium AD, represented by finds from
Nubia and Libya, as well as Medieval Senegal. However,
the continued small-grained populations in Early Historic
South India (Nevasa) and apparent reversion in later
Medieval Libya suggests that there may be factors that
work against gigantism in pearl millet, and in the absence
reinforcing selection populations may tend towards the
smaller size ranges.
This raises questions about the selection pressures
involved in large-grained Pennisetum. As both Libya and
South India lack wild populations, this cannot be attributed
to cross-pollination with wild types. There may be some
constraints particular to this crop, as one experiment indi-
cates that optimal germination occurred under higher temp-
eratures that resulted in lower average grain weights
(Mohamed et al., 1985). In addition, pearl millet involucres
are polymorphic in grain count with the vast majority pro-
ducing two grains, a large minority with one larger grain,
and a further minority producing 3–9 grains, which are
FIG. 15. Scatter plots of pearl millet (Pennisetum glaucum) grain width vs. thickness. (A) Modern population averages and minima of domesticated
populations reduced by 10% to account for shrinkage, compared with modern wild population averages with maxima, reduced by 10%. Dashed line indi-
cates expected separation between wild and domesticated forms. Sources: Brunken et al. (1977) and Zach and Klee (2003). (B) Plots of archaeological site
averages and ranges. Early West Africa averages (Birimi, 1700–1500 BCE; Kursakata, 1500–800 BCE) fall in the wild zone although ranges extend into
the larger domesticated zone. The earliest finds in India (Surkotada, approx. 1700 BCE) are close to these as are Early Historic (200 BCE–300 CE)
Nevasa in southern India. North Indian Narhan (1400–800 BCE) shows a marked shift towards larger sizes comparable with modern domesticates, as
does early medieval Qasr Ibrim (Egypt, approx. 450 CE: this find is preserved by dessication and has been reduced to be comparable with carbonized
material). Jarma in south-west Libya may show an apparent shift towards somewhat larger grains during the early first millennium CE, comparable with
the size found in medieval Senegal at Arundo. Later Medieval Jarma has shifted back towards to near wild size range. Sources: Kajale (1977), Steele and
Bunting (1982), Chanchala (1995), Saraswat et al. (1994), D’Andrea et al. (2001) and Zach and Klee (2003) (Jarma data from Ruth Pelling, personal
Fuller — Patterns in Crop Domestication919
necessarily smaller (Godbole, 1925). Thus, selection for
higher grain counts, and more reliable germination, might
conflict with selection for larger seed sizes. Nevertheless
as a working hypothesis, I would propose that, as with
pulses, there is a deeper burial threshold that selected for
gigantism in pearl millet at some times and in some
locations. In that regard it might be noted that the larger
grain populations in Libya and Nubia, like that in
Gangetic India, are associated with more intensive plough
cultures, whereas ards were not present in West Africa
and may have declined in post-Garamantean Libya. Thus,
we can hypothesize that large-grained varieties evolved
under plough systems and then dispersed back to West
Africa at a later date. If so, this would imply separate
events of grain enlargement in India and north-eastern
Africa. While initial cultivation must have selected for non-
shattering, and slight changes in grain weight and shape
(the club shape), serious gigantism may have required a
stronger selection pressure and therefore evolved later: a
millennium or more later in India, and two millennia later
Archaeological evidence indicates that the entire domesti-
cation syndrome did not suddenly appear when people
began to cultivate plants. Rather, different aspects of the
syndrome evolved in response to the new ecological con-
ditions of early cultivation. What these data suggest is
that in domesticated grasses, changes in grain size and
shattering ears or panicles (‘domestication’ sensu stricto).
While initial grain size increases may have evolved
during the first centuries of cultivation, within perhaps
500–1000 years, non-shattering was much slower, becom-
ing fixed about 1000–2000 years subsequently. Pulses by
contrast do not show evidence for seed size increase in
relation to the earliest cultivation, but seed size increase
may be delayed by 2000–4000 years. This implies that con-
ditions that were sufficient to select for larger seed-size in
Poaceae were not sufficient in Fabaceae. This implies
different thresholds of selective pressure in relation to dif-
fering seed ontogenetics and underlying genetic architec-
ture in these families. Pearl millet (Pennisetum glaucum)
may show some similarities to the pulses in terms of a
lag-time before truly larger-grained forms evolved. These
results may aid in predicting when and where certain crop
domestications are likely to have occurred based on count-
ing backwards from the earliest known domestic finds.
Thus, for example, we would predict that pearl millet culti-
vation began by 3200–2700 BC. These results also raise
questions about taxonomically linked differences in evol-
ution under the selection forces of cultivation.
evolved priorto non-
Reconsidering sickles and cereal domestication
There has been a tendency to assume that harvesting with
a sickle was the selective force that led to domestication, i.e.
non-shattering (as discussed above). The archaeological
evidence,however, does notsupport this in any
documented case. In China, as discussed already, rice
grains begin to plump and increase in size but domesti-
cation is indicated by the shift towards predominantly
mature-grained harvests (and inferred non-shattering),
during the fifth millennium BC, and by approx. 4000 BC.
In this region there are no clear archaeological sickles
until after 3500 BC, the Later Songze period (approx.
3500 BC), after which they become widespread in the
Liangzhu culture (3300–2200 BC). These sickles may be
a cultural borrowing from millet cultivators in central
China, where such tools were in use since at least 5000
BC (cf. Chang, 1986). Even in central and northern
China, the earliest sickles occur at sites that already have
millet cultivation, and earliest documented domestic
millets from Xinglonggou (near Chifeng, China), before
6000 BC (Zhao, 2005), come from a culture without
sickles. In China, sickles consistently represent a technol-
ogy development after domesticated plants are fully
In the Near East sickles were in use prior to agriculture
and must now be argued to be transferred to agriculture
relatively late, after domestication. Preserved sickles, and
more commonly lithic sickle blades, are known from
Natufian contexts (13 000–10 500 BC), in a period for
which there is no evidence for domesticates, and non-
shattering domesticates continued to be absent through
the PPNA (through 8800 BC) (see Fig. 3). Microscopic
studies of ‘sickle gloss’ have been used to suggest they
were cereal-harvesting (Unger-Hamilton, 1989; Anderson,
1992), but we cannot rule out harvesting of sedges
(Cyperaceae) and reeds (Phragmites) as materials for bask-
etry or thatching. As suggested by Sauer (1958), the early
Natufian sickles were prototype saws, designed for raw
material gathering rather than seed collecting. As indicated
by the archaeobotanical evidence reviewed above, the rate
of evolution of tough rachis einkorn and barley is far too
slow to be accounted for by a model of strong selective
pressure that would be expected if sickling was carried
out regularly, as modelled by Hillman and Davies (1990).
Thus, it would appear that early cultivators continued to
employ the time-efficient harvesting methods associated
with hunter-gatherers. Once cultivated, and populations
had noticeably large proportions (majorities) of non-
shattering types, then the transfer of the sickle technology
to agriculture may have been seen as an obvious enhance-
ment. In evolutionary terms the sickle is thus an ‘exapta-
tion’ (sensu Gould and Vrba, 1982), in that it developed
for some other purpose, and was later transferred to
crop-harvesting of already domesticated crops.
I would propose alternative explanations for the selection
of domesticated-type crops that can account for the slow
creep towards domestication. As others have noted, the har-
vesting of cereals when green, i.e. immature, regardless of
technique, will not select for domesticated types (Hillman
and Davies, 1990; Willcox, 1999). Harvesting green,
however, may not provide full returns from a given stand
of crops, as additional seeds (including late tillers) may
form and approach maturation subsequent to the harvest.
For the early farmers, who have invested significant
labour into a restricted unit of land, it becomes important
Fuller — Patterns in Crop Domestication 920
to maximize returns from that unit of land (as noted by
Hillman and Davies, 1990: 69; Bar-Yosef, 1998). This
may encourage multiple episodes of harvest. Later harvest-
ings, whether by plucking or beating, will encounter dom-
esticated genotypes in a higher frequency than earlier
harvests. If, as an aspect of random variation, some
farming households choose to store the late harvest as
seed for sowing the following year, those fields so sown
will start an increase in the domesticated type. Other house-
holds, however, may store for sowing their earlier harvests.
Therefore, taken at the level of a human community, or on a
regional scale, there might be only a very small proportion
of sown crop that had some selection for the domesticated
type. Such a model might therefore account for significantly
longer periods involved in the fixation of non-shattering
types in cultivated populations. By contrast, every farmer
and every sown population would be under selective
pressure to germinate rapidly, leading to seed size increase
and loss of germination inhibitors. Similarly, natural selec-
tion for dispersal aids such as awns will be uniformly
domestication’ traits to evolve more rapidly.
Domestication as an interdisciplinary study of evolution
Domestication in plants is not one thing, nor has it been
one uniform process. While there are recurrent parallels,
due to the same selective pressures of cultivation, different
domestication traits have evolved at different rates and these
have varied markedly across families, such as between
cereals and legumes. Further archaeobotanical research
will help to pin down the actual rates at which different
domesticates evolved, and needs to be expanded to
address a larger range of species. The archaeological
record also provides insights into what people are doing
during this evolutionary process in terms of their technol-
ogies and ecological adaptations. Understanding past
domestications is an exciting area of interdisciplinary inves-
tigation, between archaeologists and plant scientists, which
may offer insights relevant to future directions in the evol-
ution of crops under human manipulation.
These ideas have benefited from discussions with Gordon
Hillman, Sue Colledge and David Harris. I must thank
Emma Harvey, Ling Qin, Mervyn Jupe, Ruth Pelling and
Sue Colledge for assistance in compiling some of the data
sets. I thank David Harris, Sue Colledge, Mary Ann
Murray and Chris Stevens for their comments on a draft
of this paper, and the input of the peer-reviewers. I thank
the OECD for financial support to participate in the work-
Gigantism’. Funding to pay the Open Access publication
charges for this article was provided by the OECD.
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