Vegetation History and Archaeobotany
https://doi.org/10.1007/s00334-019-00727-4
ORIGINAL ARTICLE
Between domestication and civilization: the role of agriculture
and arboriculture in the emergence of the first urban societies
Dorian Q. Fuller1,3 · Chris J. Stevens1,2
Received: 10 July 2018 / Accepted: 26 March 2019
© The Author(s) 2019
Abstract
The transition to urbanism has long focused on annual staple crops (cereals and legumes), perhaps at the expense of understanding other changes within agricultural practices that occurred between the end of the initial domestication period and
urbanisation. This paper examines the domestication and role of fruit tree crops within urbanisation in both Western Asia
and China, using a combination of evidence for morphological change and a database that documents both the earliest occurrence of tree fruit crops and their spread beyond their wild range. In Western Asia the domestication of perennial fruit crops
likely occurs between 6500 BC and 3500 BC, although it accompanies a shift in location from that of the earliest domestications within the Fertile Crescent to Mesopotamia, where the earliest urban societies arose. For China, fruit-tree domestication dates between ca 4000 and 2500 BC, commencing after millet domestication and rice domestication in Northern and
Southern China, respectively, but within the period that led up to the urban societies that characterised the Longshan period
in the Yellow River basin and the Liangzhu Culture in the Lower Yangtze. These results place the domestication of major
fruit trees between the end of the domestication of staple annual crops and the rise of urbanism. On this basis it is argued
that arboriculture played a fundamental role within the re-organisation of existing land use, shifting the emphasis from
short-term returns of cereal crops into longer term investment in the developing agricultural landscape in both Western and
East Asia. In this respect perennial tree crops can be placed alongside craft specialisation, such as metallurgy and textiles,
in the formation of urban centres and the shaping the organisational administration that accompanied the rise of urbanism.
Keywords Arboriculture · Near East · Ficus · Olea · Phoenix · Vitis · China · Ziziphus · Amygdalus · Armeniaca
Introduction
Communicated by J. M. Marston.
Electronic supplementary material The online version of this
article (https://doi.org/10.1007/s00334-019-00727-4) contains
supplementary material, which is available to authorized users.
* Chris J. Stevens
c.stevens@ucl.ac.uk
Dorian Q. Fuller
d.fuller@ucl.ac.uk
1
Institute of Archaeology, University of London, 31-34
Gordon Square, London WC1H 0PY, UK
2
School of Archaeology and Museology, Peking University,
Beijing 100871, China
3
School of Archaeology and Museology, Northwest
University, Xi’an, Shaanxi 710069, China
The transformation of hunter-gather societies into the
World’s earliest civilizations has long fascinated scholars.
Gordon Childe (1936) divided this transformation into two
stages: the Neolithic Revolution and the Urban Revolution.
For Childe, domestication was a conscious and rapid process that fuelled the Urban Revolution, allowing settlements
to grow to a previously unprecedented size (Childe 1950).
Agricultural surpluses allowed the creation of “resident specialists who were themselves released from food-production” (Childe 1950, p 8) and able to engage in a range of new
technologies, from specialist metallurgy to large scale textile
production and to writing and administering the organisation
that these new systems required. Hence the social impacts
of domestication and agriculture were realized with the
scaling up that occurred with urbanization, as larger concentrations of populations, including growing numbers of
non-farming specialists and growing trade networks, were
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Vegetation History and Archaeobotany
supported (Marston 2017; Scott 2017). The largest early
agricultural villages throughout the Old World had populations in the hundreds (e.g. Kuijt 2000; Zhang and Hung
2008; Shelach and Teng 2013; Birch-Chapman et al. 2017).
As the Neolithic progressed, settlements in the thousands
arose (e.g. Moore et al. 2000; Shelach 2002; Birch-Chapman
et al. 2017), but it was only through the processes associated
with urbanization that early cities with populations in tens
of thousands emerged (Dumper and Stanley 2007; Liu 2007;
Algaze 2008; Liu and Chen 2012, p 282). However, Neolithic urbanization encompassed not just larger settlements,
but an increased number of smaller associated settlements,
some just 1 ha villages, as illustrated by regional survey data
from Mesopotamia (Algaze 2008) and the northern Chinese
provinces (Wagner et al. 2013).
The aim of this paper is to examine the scaling up of
the agricultural systems that emerged following domestication, to those that supported some of the world’s earliest
urban societies within Western Asia after 4000 BC and China
after 2500 BC. The centrality of staple cereal crops to underpinning early state formation has often been emphasized
(Steensberg 1989; Miller and Wetterstrom 2000; Algaze
2008; Scott 2017), fostering the development of writing,
administrative structures, and increasingly hierarchical
social systems (Steensberg 1989; Scott 2017). However,
other factors potentially contributing to the process of agricultural change that led to urbanism have been potentially
overlooked. In particular, playing a potentially major role in
this development was the domestication of perennial crops:
tree-fruits and vines (Miller 1991; Miller and Wetterstrom
2000; Weiss 2015). As Miller and Wetterstrom (2000, p
1,126) conclude “orchard crops began to make a noticeable
contribution to the diet” at ca 3500 BC in Western Asia.
Here we consider the role of several major perennial fruitproducing plants, the origins of arboriculture as such, as part
of the process of agricultural change that led to urbanism
within three of the original centres of ancient civilization:
in Western Asia, the middle to lower Yellow River Valley in
northern China, and the Yangtze River Delta in eastern China.
We consider the location and timing of their domestication
through both the existing literature and the first appearance of
domesticates within the archaeobotanical record. In addition
we consider the general changes in agriculture associated with
crop diversity as seen through the database alongside that for
the role of annual and perennial crops in general.
Comparing arboriculture, cereal
domestication and urbanism
Recent work has shown that many of the assumptions and
theories concerning the domestication of plants and animals are incorrect and domestication is better understood
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as a process of gradual evolution rather than rapid revolution. Domestication has been documented for an increasing number of species, utilizing chronologically sequenced
and quantified records of morphological change through
the examination of preserved remains of plants and animals
themselves (Fuller et al. 2014; Larson and Fuller 2014;
Zeder 2016; Allaby et al. 2017). Initial Neolithic domestications can be seen as protracted, unconscious evolutionary transformations, starting when the wild progenitors of
domesticated cereals and pulses were first taken into cultivation. These actions, grouped collectively under the term
pre-domestication cultivation, comprised clearance, tillage,
and sowing, and gave rise to morphological changes that
spread through cultivated populations over the course of
a few millennia (domestication) (Fuller 2007; Fuller et al.
2014; Allaby et al. 2017), but that would have been invisible
within a single human generation.
The investigation within this paper into the domestication
of a number of major perennial fruit-yielding trees, shrubs
and vines (here, collectively referred to as arboreal domesticates) is conducted through a similar lens to that applied
to annual crops. We then compare timelines of morphological change and the geographical spread in arboriculture
for individual species, alongside evidence for the relative
exploitation of various cereal and pulse crops, and human
population proxies.
The major domestication traits for fruit-trees in part relate
to fruit size and edibility; including reduced astringency,
seedlessness, and generally less stringy, softer, more palatable fruit (Janick 2005); and in part to modes of pollination,
length of juvenile phases, and cloning (Miller and Gross
2011). Most significantly for some species, stone size and
shape also changed, which can potentially be tracked from
archaeobotanical material (e.g. Terral et al. 2010; Liphschitz
et al. 2013; Pagnoux et al. 2015; Dighton et al. 2017; Fuller
2018).
Today many fruit trees and vines are grown through either
grafting a cutting onto an established rootstock or planting
cuttings directly. Such practices produce clones, whereas the
planting of stones or seeds will produce genetic variants of
both parents. In this light, given that morphological changes,
as seen for cereals, are evident in the arboreal domesticates
for which we have data, then early cultivation must have
included the planting of stones and seeds. The patterns of
change seen in stone size and shape support a protracted
gradual morphological evolution for arboreal domesticates
(see Fuller 2018), similar to that seen for the domestication of annual crops (see Fuller et al. 2014). Such evidence
corroborates an initial emphasis during the domestication
process on sexual reproduction (see Goldschmidt 2013;
Weiss 2015) through the planting of stones/seeds, while
simultaneously implying a slow rise in the importance of
perennial crops. This theory is contrary to the inference of
Vegetation History and Archaeobotany
Zohary et al. (2012) that perennial domestication was a rapid
and conscious process in comparison to annual cereals and
pulses, and that it involved primarily vegetative propagation. However, selection processes, in terms of human action
and its interaction with fruit tree genetics, are still poorly
understood.
Seed size increase during domestication is a general trait
recorded for many species, including cereals and vegetables
(Kluyver et al. 2013, 2017). In seed-grown annuals, such
changes can be associated with the creation of a level playing
field through cultivation, potentially including burial (Harlan
et al. 1973; Fuller 2007; Gegas et al. 2010; Fuller and Stevens 2017). How these explanations translate to increased
size seen in fruit stones, pips etc. is less clear. In most fruits,
increased seed length is achieved with minimal increase in
overall embryo volume. Therefore, Fuller (2018) suggests
that the lengthening of perennial fruit stones increases the
total fruit flesh with minimal additional investment in seed
volume, and thus infers some conscious element of selection for more edible matter in fruits which in turn increases
seed length and the length:width ratio. Alternatively, as with
cereals, if a number of stones, pips, or seeds were planted at
the same time, then those with larger (more elongate) seeds
that produce stronger seedlings might be selected through
the removal of smaller, less well-established seedlings. However, experimental observations of correlations of seed/stone
size and shape and seedling characteristics in fruits are still
needed to test these theories.
Despite these issues, the metrical data (Fuller 2018),
combined with database evidence for both the earliest
appearance and increased presence of fruit-tree crops, lead
to the hypothesis that their domestication and incorporation into pre-existing patterns of agricultural land-use was
a contributing factor in urbanization in Western Asia and
China. While each region contained centres of domestication and subsequent urbanization (Maisels 1998), these
processes are more spatially distinct in Mesopotamia than
was likely the case in the Yellow River or Yangtze River
(Fig. 1). Hence the Liangzhu Culture broadly matches the
same area in the Lower Yangtze for which at least one trajectory of rice domestication has been mapped (Fuller et al.
2009, 2016). Likewise the Longshan Culture arose in the
middle to lower Yellow River Valley (Henan, Shandong) and
the Guanzhong Plain (Lower Wei Valley), which comprise
at least three of the five possible centres for millet domestication (Stevens and Fuller 2017). Both China and West
Asia have rich archaeobotanical records that document the
processes of domestication of annual grain crops, including
cereals and pulses. For both regions the process of cereal
domestication spans around 2,000–3,000 years. In Western Asia the process of domestication for cereal and pulse
crops terminates around 7500–7000 BC (Fuller et al. 2018),
whereas for China the completion of cereal domestication
appears to have occurred largely around 4500–4000 BC (cf.
Stevens and Fuller 2017).
Fig. 1 Map of early centres of domestication compared to centres of
urbanization for the Near East, Northern China and the Yangtze Valley. This plots sites with early cultivation evidence (10000–6000 BC
for the Near East; 8000–4000 BC for North and South China, data
from ESM Tables S1, S2, S3) with a selection of early urban sites,
over 40 ha (from the fourth millennium BC for Mesopotamia; from
2500 to 1900 BC for China). Early urban centres in relation to zones of
agricultural origins in West Asia and China. Selected primary urban
centres, > 40 ha: (1) Tell Brak, (2) Tel Delhim and Tell al-Hayyad,
(3) Uruk (Warka), (4) Eridu, (5) Taosi, (6) Erlitou, (7) Wangchenggang, (8) Shijiahe, (9) Liangchengzhen and Yaowangcheng, (10)
Liangzhu
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Vegetation History and Archaeobotany
For China, the beginnings of urban society appear to grow
directly from the end of the initial domestication period with
perhaps only less than two millennia separating the end of
cereal domestication (ca 4000 BC) from the initiation of
urbanisation (ca 2500 BC) (Liu and Chen 2012). Conversely,
the founding crops of West Asian civilizations, wheat and
barley, were not domesticated where the first evidence for
urbanization is found. Rather they spread from the core areas
of the northern and southern Levant and/or eastern Fertile
Crescent into the middle to lower Euphrates valley after ca
7000 BC, with urbanisation occurring around three millennia
later, after ca 4000 BC, in the middle and lower Euphrates
and Tigris valleys (Maisels 1998; Algaze 2008).
Materials and methods
The assessment of the early cultivation and exchange of
selected perennial fruits is based on a database compilation
of crop archaeological occurrences, the Old World Crops
Archaeobotanical Database (OWCAD), developed by the
ComPAg (Comparative Pathways to Agriculture) project
at University College London (UCL). The database covers
Asia, Africa, and more selectively Europe, from the terminal Pleistocene to historic times. It provides an index of the
primary literature from which other information, such as
quantitative data on occurrence or metrics, can be derived.
In total, the database includes ~ 2,300 archaeological sites
with archaeobotanical data, some broken down into multiple
phases. These data have previously been used to chart the
spread of cereals between East and West Asia (Stevens et al.
2016), and within East and Southeast Asia (Stevens and
Fuller 2017). Here we utilize a subset of data representing
Western Asia and China, and associated peripheral regions.
The Western Asia data includes 489 sites/phases (based on
median calibrated radiocarbon ages) (ESM Table S1). The
Chinese sites dating predominately to between 7000 and 1 BC
are divided into two sub-regions: a northern Chinese region
centred on the Yellow River Basin (Henan, Inner Mongolia,
Jilin, Liaoning, Qinghai, Shaanxi, Shandong, Shanxi and
Zinjiang), comprising 434 sites/phases (ESM Table S2),
and a Southern Chinese region, corresponding mainly to the
Yangtze basin (Guangdong, Guangxi, Guizhou, Hong Kong,
Hubei, Hunan, Jiangsu, Jiangxi, Shanghai, Sichuan, Tibet,
Yunnan and Zhejiang) and comprising 115 sites/phases
(ESM Table S3). In preparing maps we have included the
additional occurrences of fruit taxa outside these regions,
including in Egypt, Aegean Europe, Western Central Asia,
and South Asia, and the sites plotted on each of the maps of
early fruit occurrence are also provided in supplementary
tables (ESM Tables S4–S10).
These data provide a basis for assessing the distribution of fruits in space (via maps) and time, and in respect
13
to their inferred wild native ranges. Temporal patterns are
considered in terms of ubiquity across sites in time slices,
which allow comparison of general trends in the occurrence
of these species (following Miller 1988; Weber 1991).
As a baseline comparison we show the same statistics for
cereals and major pulses. Morphological change in West
Asian crops and Chinese rice, and soybean, have been published elsewhere (Fuller et al. 2014; Allaby et al. 2017) and
are summarized from those sources in this paper. Datasets
tracking the morphological change in some of the fruit species considered here, namely peach, date and olive, have also
been published elsewhere (Fuller 2018).
Results: West Asia—the domestication
and dispersal of fig, grape, olive, and date
The geographical distribution of archaeobotanical records
for remains of all four species indicates that their earliest
exploitation began within the areas of their plausible wild
distributions in the pre-Pottery Neolithic, first occurring outside these ranges after the fifth millennium BC (the Chalcolithic), and particularly from the fourth millennium BC. The
rising importance of these species overall is evidenced from
regional ubiquity data with archaeobotanical occurrences
of stones/pips increasing in the fourth millennium BC, and
is especially marked for the grape and olive, both of which
increase subsequently in the third millennium BC (Fig. 2).
Dates are never particularly frequent, but this is likely due to
their limited native range and cultivation falling to the south
of Mesopotamia. These patterns in fruit occurrence need to
be considered alongside a chronology of urbanization, which
in the first part of the fourth millennium BC (4000–3600 BC)
saw the emergence of half a dozen sites from 25 to 80 ha,
with the massive growth of Warka to ~ 250 ha by 3100 BC
(Algaze 2008). With respect to annual crops, two trends are
worth noting. The first is that some of the fall-off seen within
annual pulses is because as these crops spread out from West
Asia the package broke apart, with chickpea, for example,
being a rare component in Iran (cf. Stevens et al. 2016). The
second is that there is a genuine decline in hulled wheats,
particularly einkorn, during the lead into urbanism with a
slight but notable increase in free-threshing wheat, a pattern
that was noted in an earlier study by Miller (1991).
Fig (Ficus carica)
The earliest evidence of consumption of fig comes from
Pre-Pottery Neolithic A sites located in the Jordan Valley
stretching north into the Upper Euphrates, and it is plausible
that cultivation had already commenced at this time. Domestication of this species likely comprised two pathways: one
involving wild recessive mutations, the other parthenocarpy
Vegetation History and Archaeobotany
Fig. 2 Percentage of West
Asian sites divided from 10000
to 8000 BC and into 1 millennium time blocks from 8000
to 1000 BC with (top) remains
of fig, date, olive and grape,
(centre) remains of leguminous
crops—pea, lentil, chickpea
and (bottom) remains of cereal
crops—einkorn, emmer, freethreshing wheat, barley. Note
the decline of hulled wheats to
free-threshing wheats and the
increase in grape and olive
(fruit development without fertilization). In the former,
recessive genes confer elongated carpels, with a long style,
to female flowers, allowing larger fleshy fruits to develop,
whilst restricting the ability of pollinating wasps to lay their
eggs. These produce large sweet fruits with crunchy “seeds”
(so-called Smyrna figs). Within this domestication process,
female plants would need to have been cultivated alongside
hermaphrodite plants (caprifigs) with both male and shortstyle female flowers, expressing the dominant genotypes and
harbouring egg-laying fig wasps. Within the second pathway, parthenocarpic mutants produce seedless fruits without
being fertilized (common figs or Adriatic figs). Kislev et al.
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Vegetation History and Archaeobotany
(2006) reported charred examples of such seedless fruits
from Gilgal (ca 9300 BC). However, Denham (2007) cautions
that such mutants, occurring naturally in wild stands, may
have been preferred by gatherers, and fig seeds appear widespread at other Pre-Pottery Neolithic A and early Pre-Pottery
Neolithic B sites, e.g. Jericho, Gesher, Mureybit III, Tell
Qaramel, Jerf el Ahmar, and D’jade (Willcox et al. 2008;
Table 1 in Fuller et al. 2012), and it is possible that cultivation of Smyrna-type figs had begun alongside early sedentary Pre-Pottery Neolithic A villages cultivating cereals and
pulses. However, wild figs were likely still present within
natural riverine gallery vegetation (cf. Bogaard et al. 2017),
and collection entirely from the wild at this date cannot yet
be ruled out.
Some fig cultivars are triploid, and others tetraploid,
suggesting a role of polyploidy within both wild populations and early cultivars that may have potentially conferred
improved traits to their fruits (Falistocco 2016). Given the
ease and rapidity with which fig shrubs can establish in or
around middens, they may even have become part of the
Fig. 3 Archaeobotanical finds of fig within West Asia, North Africa
and adjacent regions (mainly Ficus carica but potentially also Ficus
sycomorus especially in North Africa) against the wild distribution
for common fig (Ficus carica). Sites are shown in millennium blocks
from 9700 to 4000 BC above, and from 4000 to 1000 BC below (ESM
Table S4)
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Vegetation History and Archaeobotany
human-constructed ecological niches of early Pre-Pottery
settlements in the Mediterranean Levant.
Archaeological finds of figs that fall outside their projected wild zone (Fig. 3; cf. map 17 in Zohary et al. 2012)
are first seen in the fourth millennium BC, including southern
Iran. Fourth millennium BC finds in the Nile Delta presumably represent the dispersal of cultivated F. carica, although
it remains unclear where in the Nile Valley native F. sycomorus was originally distributed, or how early it came under
cultivation (most likely through vegetative propagation).
However, such finds only begin in the third millennium BC
(Zohary et al. 2012, p 130) and its native pollinator wasp
(Ceratosolen arabicus), assumed to have been once present
in Egypt, has long since been extirpated and persists today
only in sub-Saharan Africa. F. sycomorus figs are sometimes
parthenocarpic, although more often they are notched or
scraped to induce fruit set (Harlan 1986). Both F. carica and
F. sycomorus are clearly differentiated in Egyptian art (and
later literature) from at least the Old Kingdom, c. 2500 BC
(Brewer et al. 1994), and cuneiform sources record the cultivation of assumingly common figs from a similar date in
Mesopotamia (Postgate 1987).
Grape (Vitis vinifera)
Similar to common fig, early finds of olive and grape appear
restricted to the Mediterranean Levant, where they form part
of the wild flora (Fig. 4). For grape, the earliest finds mainly
Fig. 4 Archaeobotanical finds of grape (Vitis vinifera) within West Asia, North Africa and adjacent regions against the wild distribution for
grape (Vitis sylvestris). Sites are shown in millennium blocks from 9000 to 4000 BC above, and from 4000 to 1000 BC below (ESM Table S5)
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Vegetation History and Archaeobotany
come from the Jordan Valley and the northern Fertile Crescent, i.e. the Upper Tigris Basin, dating to the ninth millennium BC, although two pips of Vitis are recorded from Ohalo
II (ca 21000 BC) on the Sea of Galilee (Kislev et al. 1992).
By the eighth millennium BC, grapes are more widespread in
intervening areas of the Levant and Cyprus. Whether wild
grapes were part of the native Cyprus flora, or came with the
pre-domestication cultivation package that included crops,
livestock, and commensals, is unknown. Claims have been
made for grape domestication and the origins of wine in the
early sixth millennium BC from the Caucasus (McGovern
2003; McGovern et al. 2017). However, grape macroremains
from the earliest periods have yet to be recovered, and evidence for early wine making is based on tartaric acid from
pottery residues. However, tartaric acid is commonly found
in other fruits (cf. Barnard et al. 2011), e.g. wild crab apples,
hawthorn, or Sorbus, the last notably being cited by Virgil
(Georgics Lib. III, 379–380) as consumed by Scythians as a
fermented alcoholic imitation of wine.
The early archaeological finds of grape from the Levant
and northern Fertile Crescent more strongly support initial
cultivation and domestication in this region, where a diverse
array of annual crops was already long established. Late
Neolithic finds of grape (c. 4300 BC) from northern Greece
are likely related to their collection from the wild (Valamoti et al. 2007; Valamoti 2015), while wine production
is supported by finds of burnt pressed grapes along with
ceramic residues from jars at Dikili Tash (Garnier and Valamoti 2016). Thus, we can regard exploitation of grapes for
early wine production as potentially widespread throughout
the area of wild grape distribution. Likewise, evidence for
wine making is also evident from around 4000 BC at Areni
Cave, Armenia (Barnard et al. 2011), also within the range
of wild grape, although in this case it may relate to cultivated
grapes.
It is also from the oak woodlands of western Asia, where
wild grapes grow, that Saccharomyces cerevisiae originated.
Saccharomyces cerevisiae is the key yeast responsible for
turning grapes to wine, and phylogenetics suggest that bread
yeasts derived from these early wines (Legras et al. 2007),
while beer yeasts had other distinct origins in part (Gonçalves et al. 2016). Thus, the evolution of leavened doughs,
such as those that presumably filled Uruk-era ceramic bread
moulds, e.g., the bevel-rimmed bowl (Goulder 2010), represent a technological exaptation of wine fermentation.
The transfer of yeasts from grape wine into cereal products including beer and leavened bread was appreciated by
Miller and Wetterstrom (2000) and Sherratt (1999). Indeed,
these chemical transformations of food stuffs allowed the
full nutritional advantages of agriculture to be more fully
realized, as fermentation made available more digestible
nutrients, as well as “new forms of storage and cooking possibilities” (Miller and Wetterstrom 2000, p 1,126).
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Evidence for the start of systematic cultivation remains
elusive. Morphological change in terms of grape pip lengthening would certainly be indicative but may only evolve later
in the domestication process, and is yet to be systematically
studied for West Asian Vitis (see Bacilieri et al. 2017). Nevertheless, what is clear is the period when grapes expanded
beyond their natural wild distribution, which can be taken as
indicative of cultivation, if not domestication (Fig. 4). Grape
pips, as well as grape pollen, are recorded beyond their wild
range in eastern Iran by the fifth millennium BC, a finding
further supported by grape pollen in Lake Zeribar, western
Iran from c. 4300 BC (Miller 2008). By the end of the fourth
millennium BC, grapes were present in the Egyptian Nile
Valley, and probably the Indus Valley, where they became
an important part of urban societies by the third millennium
BC. The trade in wine can be seen to be well established in
Western Asia within written sources by the third millennium BC (Postgate 1987; Tengberg 2012). Leavened breads,
a probable by-product of Mesopotamian urbanisation, also
likely spread along expanding trade networks, including into
Egypt, as part of wider Mesopotamian cultural influence (see
Wengrow 2006).
Olive (Olea europaea)
Early olive finds are frequent throughout the Levant and
Cyprus, generally within their expected wild range, which,
while similar to that of grape, is more restricted to the fringes
of the Mediterranean (Fig. 5). Olive stones from Levantine
sites show increased length indicative of cultivation from
at least 4500 BC and continuing into the early Bronze Age,
after 3500 BC (Dighton et al. 2017; Fuller 2018). Earlier
evidence for pressing olives comes from northern Jordan ca
5200 BC (Dighton et al. 2017) and coastal Israel, although
these may be of wild olive (Tengberg 2012), suggesting cultivation was perhaps established earlier, through a mixture
of seed propagation (to account for seed size increase) and
vegetative propagation. Ultimately, vegetative propagation
came to dominate and is key to maintaining varieties. Earlier finds dating from the eighth millennium BC at Halulua
on the Upper Euphrates (Willcox 1998), lie only 100 km
beyond the present wild range, and might indicate a wider
wild distribution than present (Fig. 5). However, finds from
fourth millennium Hassek Höyük, southern Turkey (Gregor
1992), some 150–200 km beyond present wild distributions,
are more plausibly associated with its dispersal as a cultivar.
In the third millennium BC, additional finds come from east
of the wild range, including one in eastern Iraq, probably
representative of trade. By the third millennium BC, olive
trees were being managed and pressed for oil at Ebla, Syria,
as seen through textual evidence (Tengberg 2012), and
archaeological evidence in parts of Greece and the Aegean
Vegetation History and Archaeobotany
Fig. 5 Archaeobotanical finds
of Olive (Olea europaea) within
West Asia, North Africa and
adjacent regions against its wild
distribution. Sites are shown
in millennium blocks from
8000 to 4000 BC above, and from
4000 to 1000 BC below (ESM
Table S6)
(Margaritis 2013). Olives appear to be established later than
grapes in Egypt, possibly only in the New Kingdom when
shaduf irrigation facilitated the expansion and diversification of garden and orchard cultivation (Eyre 1994). Earlier
finds may indicate importation of olives to Egypt during the
Middle Kingdom, from ca 2000 BC (Caracuta et al. 2018).
Date palm (Phoenix dactylifera)
Date palm differs from those crops discussed above in originating probably from the Arabian Peninsula, with the wild
distribution potentially stretching across the Gulf of Oman
into Pakistan (Fig. 6), but seemingly less likely into North
Africa (Flowers et al. 2019; cf. Gros-Balthazard et al. 2016).
This is in agreement with Late Pre-Pottery Neolithic evidence for date palm from northern Israel, which indicates
the presence of the closely related P. theophrasti, (Kislev
et al. 2004), which explains the eighth millennium records
for Phoenix from Ain Ghazal and Atlit-Yam in Jordan and
Israel, respectively (see Rivera et al. 2014; Flowers et al.
2019). Taken together, the implication is that the wild distribution of P. theophrasti originally spread along the eastern
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Vegetation History and Archaeobotany
Fig. 6 Archaeobotanical finds of Phoenix dactylifera within West
Asia, North Africa and adjacent regions against the inferred wild
range for wild Phoenix dactylifera and P. theophrasti. Sites are
shown in millennium blocks with finds of Phoenix dactylifera
and P. theophrasti from 7500 to 5000 BC above, and all finds from
6000 to 1000 BC below (ESM Table S7)
edge of the Mediterranean, making it unlikely that the (early
Holocene) distribution of wild P. dactylifera spread this far
north or indeed west into Africa (contra map 17 in Zohary
et al. 2012).
The earliest finds of true date, P. dactylifera, come from
eastern Arabian sites in the sixth millennium BC, supporting
Arabia as the likely origin of wild date and early cultivars,
especially given the recent recognition of relict wild populations in Oman (Gros-Balthazard et al. 2017; Flowers et al.
2019). Early finds in southeast Iran and southern Pakistan
potentially indicate former more easterly wild populations
or perhaps early dispersal across the Persian Gulf/Gulf of
Oman (Fig. 6). Finds from Eridu in Southern Mesopotamia,
Teileilat Ghassul in the Jordan valley, and El Omari in the
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Vegetation History and Archaeobotany
Lower Nile, indicate the age of domesticated palm moving
north and west (Fig. 6; Zohary et al. 2012, p 134), suggesting cultivation probably commenced as early as ca 4500 BC
(Fuller 2018). Artistic evidence, including early pictographic
script, from southern Mesopotamia indicates that date palm
cultivation was well-established by Late Uruk times (late
fourth millennium BC) (Miller et al. 2016), and in the third
millennium BC date palms are commonly referred to as a
major constituent of fruit orchards and frequently associated
with burials (Tengberg 2012). Measurements on date stones
demonstrate that a general directional size change towards
elongated date stones was still occurring after the Late Uruk
period, from the third through to the first millennium BC
(Fig. 3 in Fuller 2018). Despite a few Predynastic and early
Dynastic reports from Egypt, date palm cultivation in Egypt
and Nubia only appears well established from the Middle
Kingdom onwards, becoming increasingly important from
the New Kingdom period when the shaduf improved irrigation of gardens and orchards (Eyre 1994).
namely peach, apricot, and jujube (or Chinese date). These
three species are likely native to China and are referenced in
the Late Bronze Age poems of the Xijing (Shih Ching) (see
Keng 1974; Anderson 1988). While their archaeobotanical
remains occur sporadically throughout the Neolithic and
Bronze Age, in neither Southern (Fig. 7) nor Northern China
(Fig. 8), as defined by the Qinling Huaihe Line, is there an
obvious increase that signifies systematic fruit cultivation.
Taken individually we can see that the record for each of
the three fruit species suggests increased use and probable
domestication starting in the later fourth or third millennium BC, following the prime cereal domestications, when
Neolithic settlement patterns shifted towards urbanism. It
is likely there was a shift from wild exploitation, alongside
cereal domestication, to primarily sourcing fruit from cultivation. Such a scenario can be related to the expansion of
agriculture across the Neolithic, which was accompanied by
deforestation as seen through a marked decline in arboreal
pollen at a regional level (see Fig. 5 in Stevens and Fuller
2017).
Results: East Asia—the domestication
and dispersal of peach, apricot and Chinese
jujube
Peach (Amygdalus persica, including A. ferganensis)
As in Western Asia, the earliest Chinese civilizations were
centred on cereals—millets (Setaria italica and Panicum
miliaceum) in the Yellow River Basin and rice (Oryza
sativa) in the Yangtze—that were dominant in each respective region by ca 4000 BC (Liu and Chen 2012; Stevens and
Fuller 2017). In terms of annual grain staples, early Chinese
agriculture is less diverse than that of West Asia, although
morphological changes suggest soybeans were domesticated
in Central China (the Yellow River basin) during the Middle
Neolithic, between 3500 and 1500 BC (Lee et al. 2011; Fuller
et al. 2014), while wheat was introduced to China around
2000 BC (Stevens et al. 2016). However, it should be noted
that foxtail millet became by far the most dominant crop
within the Yangshao period in northern China in the period
between domestication and urbanisation, while the increase
in millet crops in Southern China is largely due to the expansion of millet agriculture from the Yellow River into southern parts of China predominantly after 3500 BC, where it
was a minor accompaniment of rice agriculture (Stevens
and Fuller 2017). Urbanisation in China was a dispersed
phenomenon with some large regional centres appearing in
the third millennium BC in the rice-growing Yangtze (e.g.
Shijiahe, Liangzhu), but with urbanisation in the Central
Plains established by ca 2000 BC at sites with Setaria italica
as the dominant grain (e.g. Taosi and Erlitou) (Liu and Chen
2003; Liu 2007).
The fruits we discuss are those with a good archaeobotanical record, by virtue of their hard, woody endocarps:
Unambiguous wild populations of A. persica (syn. Prunus
persica), are not clearly documented, although many scholars have inferred Chinese origins (e.g. Li 1970; Keng 1974;
Huang et al. 2008; Weisskopf and Fuller 2014a). Many also
support Darwin’s conclusion that the wild ancestor of peach
is nowhere to be found (Yazbek and Al-Zein 2014). Nevertheless, Pliocene fossils (> 2.6 million years old) from
Yunnan (Southwest China) affirm an indigenous wild distribution in at least part of China (Su et al. 2015). One challenge is that related species, whose stones display similarly
ridged or deeply rugose endocarps, occur both in wild and
in semi-cultivated states, such as A. mira which is grown
and consumed in parts of southwestern China (Sichuan and
Yunnan). Other related wild peaches, including mountain
peach (A. davidiana) and Gansu peach (A. kansuensis), are
widely distributed across Northern China, but there are no
wild peach species present today within the eastern provinces south of Shandong. Recent taxonomy groups all wild
peaches into a strongly monophyletic subgeneric section
Persica, distinct from the western Eurasian section Amygdalus of almonds (Yazbek and Oh 2013; Yazbek and Al-Zein
2014). The distribution of section Persica provides a broad
geography from within which true wild peaches are likely
to have been brought into cultivation. This range is plotted
in Fig. 9, following Delplanke et al. (2016), and excluding small outlier populations on the Tibetan Plateau, and in
Mongolia and Kazakhstan.
Genetic work suggests two scenarios: a single domestication followed by multiple feral peach cultivars, or multiple
independent domestications giving rise to multiple observed
13
Vegetation History and Archaeobotany
Fig. 7 Percentage of North Chinese sites divided into 1 millennium time blocks from 8000 to 0 BC with remains of Chinese jujube, peach and
apricot above and broomcorn millet, foxtail millet, rice, soybean and wheat below (ESM Table S2)
traits (Akagi et al. 2016). This same study demonstrated
hybridization between single accessions of wild A. davidiana and A. mira, and cultivated A. persica. This observation, together with the lack of an obvious genetic progenitor
from Northern China, despite the presence of two wild peach
species there today, and given that the wild peach ancestor
is likely to have been geographically isolated from closely
related species, would support domestication from a now
extinct progenitor that came under cultivation within at least
13
the Lower Yangtze, where the earliest archaeological evidence has been recovered, but possibly also the Huai River
and/or Middle Yangtze. Under this scenario early records of
peach from Northern China might represent their collection
from the wild rather than early cultivation.
The earliest archaeobotanical records of peach from
the lower Yangtze, Yangtze delta, Middle Yangtze, Huai
Valley and Northern China date from 6000 to 3500 BC
(Fig. 9). Stones of A. persica from around 6000 BC to ca
Vegetation History and Archaeobotany
Fig. 8 Percentage of Southern Chinese sites divided into 1 millennium time blocks from 7000 to 0 BC with remains of Chinese jujube, peach and
apricot above and broomcorn millet, foxtail millet, rice, and soybean below (ESM Table S2)
2300–2000 BC were studied from sites within the Lower
Yangtze and Yangtze delta (Zheng et al. 2014). This study
noted that stones became larger and less spherical through
time, and while little change is seen between 6000–5000 BC,
from 3500 to 2000 BC there is a considerable increase in
size (Fig. 2 in Fuller 2018; Fig. 6 in Zheng et al. 2014). This
suggests the initiation of cultivation, at least in the Lower
Yangtze, began around 4000 BC, coinciding with the end
of rice domestication. Genetic work also indicates a strong
bottleneck in the cultivar around 3000 − 2000 BC, when
compared to the wild mountain peach A. davidiana (Faust
and Timon 1995). That peaches were under cultivation by
3000 − 2000 BC is supported not just by the increase in stone
size, but also the first occurrence of peach in the Shandong
region. The subsequent appearance of peach in Late Neolithic Kashmir and in Late Jomon Kyushu, Japan, during the
second millennium BC, suggests translocation of the cultivated tree (Weisskopf and Fuller 2014a; Zheng et al. 2014;
13
Vegetation History and Archaeobotany
Fig. 9 Archaeobotanical finds of Amygdalus persica and A. cf. persica from China and India and the inferred probable range of section Persica
(ESM Table S8)
Stevens et al. 2016). In summary, morphological change was
under way in the third millennium BC in Chinese peaches
with translocation to distant regions by the second millennium BC taking place during the period of urbanization.
Apricot (Armeniaca vulgaris)
As with peach, the domestication history of apricot raises a
number of issues. While the wild ancestor is unknown, wild
or feral populations are widely distributed not only throughout China, but also in Central Asia, the Caucasus Mountains,
and along the Himalayan foothills. As long ago as 1882, de
Candolle (1885, pp 215–218) argued for a Chinese origin,
based on the recovery of small wild-type fruits from Henan,
together with historical records showing a slow dispersal
into Europe and West Asia, a clear great antiquity of use in
China, and linguistic evidence including its long-established
single character in Chinese, xing (杏), comprising the compounds for tree/wood and mouth. Genetic evidence (microsatellite markers) indicated three clusters of diversity focused
on the Caucasus, eastern Central Asia, and Central China
(Maghuly et al. 2005), although this study lacked systematic
sampling of Chinese cultivars or wild populations. However,
it seems likely domestication originated in China, with later
introgression with wild populations in Xinjiang and the Caucasus (Weisskopf and Fuller 2014b), diffusing at a later date
via central Asia into Eastern Europe (Maghuly et al. 2005).
The earliest records for apricot in China are widely dispersed, covering northeastern China, the Huai River, and the
lower Yangtze delta region (Fig. 10). Regarding wild species
in China it might be noted that the Flora of China (Lu and
Bartholomew 2003) lists a number of wild species of apricot
13
including Armeniaca holosericea in north-western China, A.
hongpingensis in Hubei and Hunan, and a number of other
species distributed across Southern China. Of some interest
is that the modern distributions of A. mandshurica, A. sibirica and wild A. vulgaris var. ansu in Northern China overlap
with each other, which, given that all can readily hybridise, suggests that some may be feral populations. Today
only wild A. vulgaris var. ansu is reported from Jiangsu,
with no records from modern Zhejiang, which suggests, as
with peach, that at least one area of likely domestication
lies within the lower Yangtze or Yangtze delta, given the
region’s continuous archaeobotanical record. In northern
China, early records could potentially represent other wild
species of Armeniaca, although there are no genetic studies
as yet to indicate the relationship or indeed differentiation
between wild species within China.
Early specimens of apricot are reported at around
4000–3000 BC from Neolithic settlements in Ukraine (see
Zohary et al. 2012), but these are difficult to reconcile with
the general pattern of evidence for apricot. It is possible
that they represent a separate Caucasian origin, but caution is warranted given that reports of Chinese millets from
Ukraine at this time are equally problematic (MotuzaiteMatuzeviciute et al. 2013; Stevens et al. 2016). We therefore
conclude that cultivated apricot originates in China, and that
its occurrence in Kashmir at the end of the local Neolithic
during the second millennium BC marks the first dispersal
of cultivated apricots outside of China (Stevens et al. 2016).
Vegetation History and Archaeobotany
Fig. 10 Archaeobotanical finds of Armeniaca from China, India and Nepal, from c. 6000–0 BC (ESM Table S9)
Chinese jujube (Ziziphus jujuba var. jujuba)
There exists some confusion regarding the taxonomy of
this species, and presently the two cultivars are often both
assigned to Z. jujuba Mill. However, archaeological and
genetic evidence indicates that they should be separated
into two distinct species and hence domestication events: Z.
jujuba var. jujuba and Z. mauritiana Lam. (Indian jujube).
Genetic studies show clear separation between the cultivated
Indian and Chinese jujube, together with its wild progenitor, sour jujube (Z. jujuba Mill. var. spinosa (Bunge) Hu ex
H. F. Chow, syn. Z. acidojujuba, Z. vulgaris var. spinosa)
(Ma et al. 2011). Further, recent genetic studies in China
suggest that the clustering of two wild varieties of Z. jujuba
var. spinosa with separate clades of cultivars might relate to
two distinct domestication events (Xu et al. 2016), or two
regional patterns of introgression.
Wild jujubes are characteristic of secondary growth in
the temperate deciduous broadleaf forests of Central and
Northern China, from Liaoning in the east towards Gansu
and Qinghai in the west (Wang 1961, pp 74–92). The wild
progenitor Z. jujuba var. spinosa is widely distributed in
Northern China, as well as the mountain foothills of Anhui
and Jiangsu, with other congeneric species confined to
southwest China (Fig. 11 shows its potential wild distribution). A number of early records exist for the Upper Huai
River/Middle Yellow River, with a single early record in
the Middle Huai. The continuous archaeobotanical record
within this region suggests it was most likely first cultivated
here alongside millet. Unlike peach and apricot, jujube is
largely absent from archaeobotanical records for the lower
Yangtze. Findings outside this zone, including in Yunnan,
would suggest that domesticated, cultivated varieties were
present by at least the second millennium BC.
Differentiation between the more globular stones of
wild sour jujube and the narrower pointy stones of modern cultivars (see Liu et al. 2008, pp 137–138) allow the
two to be distinguished and suggest a potential means for
tracking Chinese jujube cultivation. While no such studies
have been conducted for China, the identification of jujube
stones as wild or cultivated stones from a number of sites
does allow some preliminary observations to be made. At
Xijincheng, Henan, cultivated stones were only identified
from later Tang/Song dynasty deposits, with wild-type
stones seen in Longshan deposits (Chen et al. 2010), while
definitive cultivated stones resembling modern specimens
were present in the Early Western Han period (Zhao and
Wang 2016). Given the two to three millennia of morphological domestication seen for other arboreal domesticates,
these data suggest that cultivation potentially begun in the
Late Longshan period. Slightly elongated and acute stones
from Yangshao Yuqiao, Henan (ca 3000 BC ), may suggest that selection resulting from cultivation had started at
this date, although the later Longshan stones from nearby
Xiawu still included round wild types some 1,000 years
later (Fuller and Zhang 2007). As with other trees today,
Chinese jujube is reproduced through grafting, but as discussed above the observed morphological changes within
the stone shape implies that until at least the Han period its
cultivation was through planting the stone. Planting from
13
Vegetation History and Archaeobotany
Fig. 11 Archaeobotanical finds of Ziziphus jujuba var. jujuba (Chinese jujube) from China and the inferred probable range of wild Z. jujuba var.
spinosa (ESM Table S10)
seed would have also maintained high levels of variation
and potential reversion to more wild-like forms via crosspollination, and thus high variation in shape among early
archaeological finds is to be expected.
Discussion: a comparative summary of fruit
tree cultivation before urbanization
The evidence presented above demonstrates how the domestication of the major fruit trees, with the possible exception
of figs, likely falls within the time period that follows the
domestication of annual grain crops and establishment of
agriculture, but before the emergence of large urban centres
of population within both regions. In his definition of the
Urban Revolution, Childe (1936, 1950) laid emphasis on
craft specialization, metallurgy, the trade of craft items over
long distances, and the accumulation of surplus and wealth
in early cities, where ruling classes lived and monumental public buildings were created. The reconsideration of
fruit-tree domestication outlined above, in line with Childe’s
theory of urbanization, indicate a shift from the cultivation
and domestication of annual staples, which can be regarded
as driven by subsistence needs centred on maintaining food
security, to domestications that relate to Childe’s other
aspects of subsistence, such as trade, craft production, and
defining social status.
It might be noted that several authors have highlighted
agricultural intensification as key to the process of urbanization (e.g. Trigger 2003; Yoffee 2005). However, isotope
13
data from cereal grains spanning the Neolithic to Bronze
Age indicate a general decline in manuring intensity in West
Asia, suggesting that more extensive rather than intensive
agricultural systems supported urbanism (Araus et al. 2014;
Styring et al. 2017). Although the creation of irrigation systems during the fourth millennium BC in at least Southern
Mesopotamia (Wilkinson and Jotheri 2019) is likely to have
played a role in such changes, agricultural intensification
is better demonstrated for more recent periods of human
history where larger populations have been progressively
supported by the cultivation of less land per person (Ellis
et al. 2018), and it is against this that these earliest trends
towards extensification must be reconciled. Part of this reconciliation can be found in the emergence of longer supply
chains of trade that contributed to feeding early cities, but
to this we may add the development of a new mosaic of land
use, encompassing a wider variety of crops, that operated on
different temporal cycles.
The period between the appearance of domesticated cereals that supported large Neolithic mega-sites like Çatalhöyük
in the seventh millennium BC (Bogaard et al. 2017) and the
rise of urbanism in the West Asia was around 4,000 years.
In China, we might estimate this period to have been shorter,
with some 1,500 years separating the end of domestication
and the rise of urbanism (from ca 4000 to 2500 BC). One of
the agricultural developments that emerged in this period,
as emphasized by Sherratt (1999), was the introduction of
commodity crops. Value was added in the transformation
of these crops into dried fruits, oil, wine etc., increasing
their longevity while reducing their bulk and facilitating
Vegetation History and Archaeobotany
their transportation as prestige goods. The first cities then
emerged as part of this process, as centres with specialized
social and economic functions, drawing in raw agricultural
produce from their surrounding hinterland, transforming it
into added value commodities, and redistributing the products to an increasingly large non-farming population (Trigger 2003; Sherratt 2011). As argued by Renfrew (1972) in
his assessment of the development of Aegean social complexity, the new exploitation of crops including olive and
grape created “a new flexibility in subsistence strategies,
and […] the possibility of production specialization in single commodities” (Renfrew 1972, p 280; also; Margaritis
2013). In other words, agricultural commodity specialization
occurred at the forefront of trends to craft specialization.
The emergence of perennial domesticates during these
periods then provided a new basis for manufactured food
commodities for exchange to be routed through cities,
accompanying the general increase in trade of specialised
craft items. This secondary agricultural revolution can be
seen as a parallel development comparable with the specialization of animal secondary products such as wool, milk, and
animal traction (Sherratt 1999).
In West Asia these developments first arose on the peripheries of urban Mesopotamia, but the exchange of such items
soon came under the control of urban elites (Wengrow 2008),
increasing their value by association with emergent highranking social classes, and their institutions and temples.
The production of textiles was similarly a process that drew
on raw produce from the periphery that was transformed in
value by urban labour. As explored by McCorrsiton (1997)
for Mesopotamia, this involved high levels of wool imported
into cities alongside the more labour-intensive processing
of flax. Flax for linen, wool sheep, and perennial crops all
contributed to a new form of land use encompassing longerterm (perennial) production, what we might term investment
agriculture, to differentiate it from the annual cropping, or
sustainability agriculture, that focused on annual production
of staple foods. This investment agriculture often relied on
longer-term input of labour into land, relying on accepted
norms of long-term land tenure, something that urban societies and states tended to enforce. In contrast to annual cereal
and pulse crops, fruit and vines required longer-term investment with delayed returns. This created a tension between
increasing the production of staple calories to support denser
populations and allocating land for commodity crops, a
tension well-documented from historical periods, e.g. the
Islamic expansion of cotton production in Iran at the expense
of further grain production (Bulliet 2009). However, the
expansion of crop diversity through long-lived perennials,
such as tree-fruits and textile crops, has been less thoroughly
documented for the later Neolithic, Chalcolithic and Early
Bronze Age.
The cultivation of perennial crops can be seen as establishing a new philosophy of land use, requiring longer-term
investment, which included the transformation of crop products into value-added commodities for further exchange.
Thus the cultivation of perennials can be argued to have been
a prerequisite for some of the social changes that underpinned urbanization. These include developments in land
ownership and rights of tenure associated with emergent
elite bodies, resulting from increased social stratification.
Indeed, some of the earliest written law codes in ancient
Egypt, Mesopotamia, Israel (Ellickson and Thorland 1995)
and China (Zhang 2014) enshrined ownership of agriculturally productive land.
In the case of China, archaeobotanical research has
clearly documented the establishment of cereal agriculture
in the Neolithic by ca 4500–4000 BC in both the Yangtze
and Yellow River basins, but less attention has been paid to
the diversification in arboreal domesticates. The evidence
presented in this paper suggests that tree fruit cultivation
had begun by ca 3000 BC and domestication was underway
during the third millennium BC when urbanism emerged by
the end of that millennium. The extent to which tree-fruit
commodities, or other cash crops such as those for textiles
(hemp, ramie, silk raised on mulberry orchards), were also
tied to urban elite controls and redistribution requires further
research, but in general we hypothesize similar processes
of agricultural diversification towards perennial investment
systems in China as in Mesopotamia.
Acknowledgements This paper was supported by a European Research
Council grant “Comparative Pathways to Agriculture” (ComPAg, no.
323842). It was prepared to honour the distinguished archaeobotanical
career of Naomi Miller.
Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativeco
mmons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate
credit to the original author(s) and the source, provide a link to the
Creative Commons license, and indicate if changes were made.
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