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R E V I E W
From foraging to farming in the southern Levant:the development of Epipalaeolithic and Pre-pottery Neolithic
plant management strategies
Eleni Asouti Dorian Q. Fuller
Received: 14 May 2011 / Accepted: 29 October 2011 / Published online: 13 November 2011
Springer-Verlag 2011
Abstract This paper reviews the archaeobotanical record
of the transition from foraging to farming in the southernLevant. The concise presentation of the published botanical
evidence follows a critical assessment of: (a) the nature of
Epipalaeolithic plant management strategies, (b) the place
of the southern Levant in the polycentric development of
Near Eastern plant cultivation and domestication, and
(c) region-specific pathways for the emergence of domes-
ticated crop packages. Some inferences are drawn and
suggestions are made concerning the potential contribution
of archaeobotanical research to questions of broader
archaeological significance about socio-economic change
in the southern Levant during the Pre-pottery Neolithic.
Keywords Origins of agriculture Southern Levant
Natufian Neolithic Cultivation Plant domestication
Introduction
This paper aims to provide a critical overview of the
archaeobotanical record of the transition from foraging to
farming in the southern Levant (geographically encom-
passing southern Syria, Lebanon, Palestine, Israel and
Jordan). It compares datasets from different parts of the
region with the aim of generating hypotheses that may
enrich our current understanding of the long-term devel-opment of food production in this area. The chronological
boundaries of this review extend from the Epipalaeolithic
to the end of the Pre-pottery Neolithic B (*210007000
cal. B.C.). Such a broad time-span is justified by the
diversity of scholarly opinion debating short- versus long-
gestation models and single or multiple centres of origin
for the emergence of the domestication syndrome in cereals
and pulses (Abbo et al. 2010; Allaby et al. 2010; Colledge
et al. 2004; Fuller 2007; Fuller et al. 2011; Honne and
Heun 2009; Nesbitt 2002; Tanno and Willcox 2011; Weiss
et al. 2004; Willcox 2005). For this reason it is necessary to
consider datasets that bridge the Epipalaeolithic-PPN
divide and are, theoretically at least, capable of providing a
sound factual basis from which to evaluate the long-term
development of plant management strategies in this region.
Thematically, the paper addresses a number of key issues
that have preoccupied scholarly debate in recent years.
What was the nature and socio-ecological context of the
south Levantine plant management practices and how did
they change through time? Is it possible to trace a contin-
uous trajectory of co-evolving practices and political
economies, or were there important disjunctions en route to
food production? How were the south Levantine crop
packages constituted, and what was the contribution of
local developments versus regional influences in this pro-
cess? Given the limited journal space available, it is not
possible to provide here a detailed description of the
archaeological evidence or the history of archaeobotanical
research. The regional culture-historical entities, subsis-
tence economies and corresponding chronological periods
are summarised in Table 1 (see also Fig. 1 for a map with
the location of the sites mentioned in the text). Readers are
referred to other publications for more detailed discussions
Communicated by G. Willcox.
E. Asouti (&)
School of Archaeology, Classics and Egyptology, University
of Liverpool, Brownlow Street, Liverpool L69 3GS, UK
e-mail: E.Asouti@liverpool.ac.uk
D. Q. Fuller
Institute of Archaeology, University College London,
31-34 Gordon Square, London WC1H 0PY, UK
123
Veget Hist Archaeobot (2012) 21:149162
DOI 10.1007/s00334-011-0332-0
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(Bar-Yosef and Meadow 1995; Colledge 2001; Delage
2004; Henry 1989; Kuijt 2000; Kuijt and Goring-Morris
2002; Bienert et al. 2004; McCorriston 2009; Simmons
2007).
Epipalaeolithic plant management strategies:
a prelude to cultivation?
Evidence, in the form of well-preserved plant macrofossils,
for pre-Neolithic plant management strategies in the
southern Levant is extremely limited. At the time of writ-
ing, the only site with well-preserved and sufficiently
documented archaeobotanical remains is early Epipalaeo-
lithic Ohalo II, located on the southwestern shore of the
Sea of Galilee in the Dead Sea rift valley. Ohalo II con-
tained abundant waterlogged remains of wild-type barley
and emmer, oats, goat-grass, almond, hawthorn, pistachio,olive and grapes (Kislev et al. 1992). In the northern
Levant, Late Natufian Abu Hureyra 1 (Moore et al. 2000;
below) is the only other Epipalaeolithic site that has pro-
duced well-preserved archaeobotanical remains. Ohalo II is
exceptional not only for its excellent preservation but also
for the recovery of specialised food processing features: an
indoors large flat basalt stone, supported by small pebbles,
was associated with large quantities of charred barley and
grass seeds as well as several species of plants with
ethnographically reported medicinal uses (Weiss et al.
2008), and an outdoors stone-paved hearth that was cov-
ered with ash and charcoal that might have been used as a
baking installation (Piperno et al. 2004). There were also
indicators of multi-seasonal hunting, fishing and plant
gathering, a human burial, flint, bone and ground stoneconcentrations, as well as evidence of elaborate symbolic
behaviours (Kislev et al. 1992; Nadel 2003; Nadel et al.
2006).
Had such attributes been found in association with Early
Natufian complex hunter-gatherer habitations, they
would have been hailed as signifiers of new socio-eco-
nomic behaviours linked to sedentism. The assumption that
Early Natufian symbolic behaviours, sedentism, territori-
ality and increasing population densities represented a
departure by an order of magnitude from early and middle
Epipalaeolithic phenomena, hence presaging by some
3,000 years Neolithic food producing societies, is deeplyingrained in the literature (cf. Bar-Yosef1998; Bar-Yosef
and Belfer-Cohen 1989; Henry 1989; Hayden 2004; Sim-
mons 2007). Fragmentary as it is, the southern Levantine
record contests such views. Recent fieldwork at Uyun al-
Hammam in Wadi Ziqlab, northern Jordan (Maher et al.
2011a) and Kharaneh IV in the Azraq basin of the Eastern
Desert (Richter et al. 2011) has suggested that organised
cemeteries, logistic management of resources, long-range
exchange networks and elaborate symbolic behaviours
Table 1 Summary of southwest Asian late Epipalaeolithicearly Neolithic culturalhistorical entities, their broad chronological framework and
associated key developments in regional subsistence economies
Chrono-cultural horizons Cal. years B.C. Near Eastern culture-historical entities and key features of regional
subsistence economies
Late Epipalaeolithic *1200010000 Natufian (Levant, south Anatolian coast); Epipalaeolithic of the
northeastern Fertile Crescent (Taurus-Zagros arc): hunting-gathering
Pre-pottery Neolithic A (PPNA) *105008700 PPNA habitations at Gobekli Tepe, Cayonu (SE Anatolia): hunting-
gathering (pre-domestication cultivation?)
Khiamian (northern, southern? Levant); early PPN of the northeastern
Fertile Crescent: hunting-gathering
Mureybetian (northern Levant); Sultanian (southern Levant): hunting-
gathering, pre-domestication cultivation
Early PPNB *87008200 Northern Levant, southeast Anatolia (persistence of the PPNA in the
southern Levant?); early PPN of the northeastern Fertile Crescent;
early Cypro-PPNB; earliest known Neolithic settlement in central
Anatolia: pre-domestication or mixed cultivation, hunting-gathering,
herding, first appearance of domesticated crop packages
Middle PPNB *82007500 MPPNB cultures of the southern Levant; aceramic Neolithic cultures
of the northern Levant, southeast and central Anatolia, Cyprus, and
the Zagros: diverse habitation patterns and subsistence practices
observed region-wide
Late PPNB *75007000 Late aceramic Neolithic cultures; southern Levantine megasites:
establishment and expansion of mixed agro-pastoral economies
based on cereals, pulses and caprine herding region-wide;
completion of the plant domestication process; widespread adoption
of domesticated crop packages in mixed agropastoral economies
150 Veget Hist Archaeobot (2012) 21:149162
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were manifested in earlier periods as well. Althoughanalyses of botanical finds from these projects are still
pending, it is likely that they will produce important evi-
dence on the nature of little known Epipalaeolithic plant
management practices.
South Levantine Natufian sites with published macro-
botanical remains are few, including Early Natufian Wadi
Hammeh 27 that contained wild-type barley, pistachio
nuts, figs and legumes (Colledge 2001) and Hayonim Cave
that preserved two fragments of wild-type barley, almond
nuts and legumes (lupin seeds and possibly pea; Hopf and
Bar-Yosef 1987) and Wadi Jilat 6 containing mainly
chenopods and sedges (Colledge 2001). Poor preservation
of charred plant remains and the lack of systematic
retrieval from older excavations are believed to be the
principal causes for the low visibility of plant remains at
Natufian sites (Bar-Yosef 1998). Plant consumption prac-
tices might be another explanation, with fewer fires used
for cooking plant foods such as vegetables, while tubers
and root crops could have been roasted by direct heating
and thus preserved only as unidentifiable residues
(Colledge 1991). Such observations have not prevented
scholars from proposing that plant-based subsistencefocused on intensive cereal gathering and even cultivation.
In the absence of tangible botanical evidence, material
culture such as sickle blades and ground stone (cf. Edwards
2007; Wright 1991) has been used to claim routine cereal
management and consumption. Coupled with the evidence
for substantial architecture and multi-seasonal habitation in
the Mediterranean Woodland Zone (MWZ) of the Levan-
tine littoral, such indicators have contributed to the per-
ception of the Natufian as a precursor and a threshold to
later Neolithic village life and food production (see
discussions in Henry 1989; Bar-Yosef 1998; Cauvin 2000;
Valla 2000; Byrd 2005; Simmons 2007).
Other authors have proposed different models suggest-
ing that tree fruits and nuts were used alongside grasses and
legumes in the MWZ, with grasses being more prominent
in the steppe and woodland-steppe habitats east of the river
Jordan (cf. Colledge 2001; Olzewski 1993, 2010; Savard
et al. 2006; contra McCorriston 1994). Plant remains from
Natufian sites outside the MWZ are also few, including
charred grain fragments (Byrd and Colledge 1991; Coll-
edge 2001) and phytoliths (Olzewski 2010). With the
Fig. 1 Map showing the location of the sites mentioned in the text (included are the sites listed in Tables 2 and 3
Veget Hist Archaeobot (2012) 21:149162 151
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exception of a few large sites that might have functioned as
base camps, most habitations in the steppe were small,
indicating high degrees of mobility; material culture finds
included ground stone, bedrock mortars and cup holes, but
no sickle blades (Olzewski 2010). It is therefore likely that
Natufian grass management practices did not always or
necessarily go hand in hand with reduced mobility, storage
and large permanent habitations. Preliminary analyses ofphytoliths from Early and Late Natufian sites situated in the
MWZ (Hilazon Tachtit, Ain Mallaha) have suggested that
woody plants were more prominent in the Early Natufian,
while in the Late Natufian non-cereal grasses were domi-
nant (Rosen 2010). Against the expectations of models
proposing that Late Natufian subsistence increasingly
focused on the intensive management and cultivation of
cereals (Hillman et al. 2001; Moore and Hillman 1992;
Sherratt 1997) most phytoliths have derived from non-
cereal grass taxa. Rosen (2010) proposes that the harvest-
ing of small-seeded grasses formed a stable adaptive
strategy to the abrupt climatic deterioration of the YoungerDryas (YD*109009600 cal. B.C.; Alley et al. 1993;
Severinghaus et al. 1998). Its key attribute was the diver-
sification of gathered plant resources, rather than the cul-
tivation of cereals on alluvial surfaces and lake margins as
suggested by the classic models. A recent detailed re-
evaluation of the archaeobotanical evidence from Abu
Hureyra 1 by Colledge and Conolly (2010) has questioned
previous propositions by Hillman et al. (2001) that the
inhabitants of the site began cultivating cereals and large-
seeded legumes as a response to their reduced availability
during the YD. Wild-type cereals contributed\15% of dry
land taxa in all Epipalaeolithic phases (13), while other
potential food plants appear in higher frequencies in phases
23 (YD) including small-seeded grasses, legumes,
chenopods, feather grasses and valley-bottom plants such
as Polygonum and Scirpus (Colledge and Conolly 2010).
They interpret this evidence as indicating the diversifica-
tion of wild plant gathering during the YD. In turn, such an
explanation, rather than the intensification of cereal man-
agement practices, may be responsible for the increased
frequency of ground stone artefacts in Late Natufian sites
(Colledge and Conolly 2010, p. 136; see also Dubreuil and
Plisson 2010).
The early PPN (10thmid-9th millennia):
pre-domestication cultivation
The beginning of the PPN in the southern Levant is
considered to be coeval with the end of the YD
at *9600 cal. B.C. Recent assessments of the south Lev-
antine radiocarbon dates (Maher et al. 2011b and refer-
ences therein) have suggested that the onset of the PPNA
may be somewhat earlier at certain sites such as Wadi
Faynan 16/WF 16 starting into the YD. South Levantine
early PPN sites with relatively well-preserved and pub-
lished archaeobotanical assemblages include Netiv Hag-
dud (Kislev 1997), Gilgal I (Weiss et al. 2006), Zahrat
adh-Dhra (ZAD) 2 (Edwards et al. 2004), Iraq ed-Dubb
(Colledge 2001) and el-Hemmeh (White and Macarewicz
2011). Jericho (Hopf 1983) was excavated in the 1950swhen flotation sampling was not practised, and has thus
produced few materials. The study of plant remains from
Dhra is still at a preliminary stage (Colledge et al. 2008).
Previous sampling at WF16 has indicated reliance on
gathered resources with very limited contributions from
wild-type cereals (Jenkins and Rosen 2007; Kennedy
2007). Recent fieldwork has revealed large horizontal
exposures of buildings and external areas that were sys-
tematically sampled for the retrieval of plant remains
(Finlayson et al. 2009). Therefore forthcoming archaeo-
botanical analyses may change this picture.
None of these sites has provided incontestable evidencefor domesticated-type cereals (Nesbitt 2002; see also
Table 2). On the contrary, there is increasingly accumu-
lating evidence for the pre-domestication cultivation of
barley and, less conclusively, emmer and pulses. Parthe-
nocarpic figs might also have been planted by vegetative
propagation (Kislev et al. 2006; cf. Denham 2007). In
addition, there is the inference of dead-end early culti-
vars such as oats at Gilgal I (Weiss et al. 2006) and Vicia
peregrina (rambling vetch) at Netiv Hagdud (Melamed
et al. 2008). The case for pre-domestication cereal culti-
vation at Gilgal I is based on the finds of large stores of
wild-type oats and barley (Weiss et al. 2006), while at
Netiv Hagdud the abundant finds of V. peregrina are
considered as indicators of cultivation and possibly storage
(Melamed et al. 2008). Lens (lentil), too, was found in large
concentrations, while the existence of an arable weed flora
has been inferred from the presence of non-cereal taxa. At
both sites, the absence of basal collars from the barley ears
has suggested to Kislev et al. (2004) that harvesting might
have been done by collecting disarticulated spikelets from
the ground rather than with a sickle. A mixture of ground-
picking and sickle harvesting strategies is proposed for
Hemmeh (White and Macarewicz 2011). At ZAD 2
(Edwards et al. 2004) barley rachises were mainly of the
wild type, although the proportion of non-shattering types
was higher than what would be expected in wild popula-
tions: at least 8% if all indeterminate are regarded as wild
but rising to 29% when only determined rachises are
considered. This suggests that some selection under culti-
vation was underway. Average grain size was similar to
measurements obtained from Jerf el Ahmar (early levels)
and Tell Qaramel, although a smaller proportion of larger
grains might also indicate that selection for increasing
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grain size had also begun (for a list of all measurements,
see Fuller et al. 2011). This observation and the presence of
possible arable weed taxa suggest pre-domestication cul-
tivation of barley; on the other hand the wild-type emmer
wheat found at ZAD 2 was probably gathered (Edwards
et al. 2004).
The presence of cereal domesticates has been claimed at
Iraq-ed Dubb and Jericho. The former did not contain
domesticated-type barley rachises; wild-type barley was
probably cultivated (Colledge 2001; Fuller 2007). Colledge
(2001) has reported a single einkorn-like wheat grain of the
wild-type size range, to which she has attributed cultivar
status, based on its occurrence outside the assumed natural
distribution for einkorn. However, the presence of einkorn
at Iraq-ed Dubb has been questioned by Willcox (2007)
and its cultivar status rejected by Nesbitt (2002). Wheat
spikelet forks were not identified to species level, which
leaves open the possibility that they derived from wild-type
Table 2 Summary of archaeobotanical data for wild- and domesti-
cated-type crops and their progenitors in the southern Levant
(including figs). x present (non-quantified, or sample size too small
to quantify), D domesticated-type, d partial development of domes-
tication syndrome (large seed size or presence of a sizeable minority
of non-shattering rachises), ? possibly present, A-s absence consid-
ered significant for region/period, and is genuine rather than being
attributable to sampling and identification biases. Semi quantitative
ranking for taxon representation: XXX very frequent/dominant, XX
frequent, X present, low frequency. Note that wild-type two-grained
einkorn grain is indistinguishable from rye based on grain morphol-
ogy alone; the status of PPN finds of Vicia faba (broad bean) as
domesticated remains unresolved; C-14 age ranges represent the
summed probability of the calendar age for each site at 1r;
probabilities were calculated and calibrations performed using the
OxCal 3.10 software (Bronk Ramsey 2005) and the most recently
revised calibration curve (IntCal09; Reimer et al. 2009)
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emmer rather than einkorn. Colledge (2001) has also
inferred the presence of an arable weed flora. Overall, pre-
domestication barley cultivation at Iraq-ed Dubb seems
likely; however, it remains very uncertain whether einkorn
was cultivated.
Carbonised plant macrofossil evidence for plant man-
agement activities at Jericho is, as noted already, limited.
Cereals were certainly used, as indicated by hand-pickedcharred remains and plant impressions (Hopf 1983).
However, their status as locally cultivated and/or gathered
foodstuffs is somewhat unclear. From Jerichos PPNA
phase (equivalent to the late PPNAEPPNB regional
horizon; see Tables 1 and 2) few specimens were retrieved:
three grains attributed to einkorn, 34 to emmer and 32 to
barley. Hopf (1983) has reported a mixture of larger
(domesticated-type) and thinner (wild-type) barley and
emmer grains. Lentil and (possibly) chickpea fragments
were also present. Pulses appear in higher frequencies
and with more convincing evidence for the presence of
domesticated crop types in Jerichos PPNB phase(equivalent to the MPPNB horizon). Chickpea was proba-
bly cultivated as it is found outside its natural distribution
and probably only slightly later than the earliest chickpeas
in the northern Levant, at Tell el-Kerkh (Tanno and Will-
cox 2006). Lentils also showed a shift to larger average
diameters by comparison to PPNA wild types. Therefore, it
is reasonable to consider Jerichos PPNA plant assemblage
as representative of pre-domestication cultivation with
partially developed domestication syndrome. Cultivation is
also indicated by the occurrence of chaff impressions in
mud bricks suggesting that dehusking waste from crop
processing was used for the production of building mate-
rials. At the same time, Jerichos PPNB plant assemblage
represents the transition to the cultivation of fully domes-
ticated crops.
Another transitional site is Tell Aswad in the Damascus
basin. Aswad (EPPNB) has produced a large assemblage of
barley rachises predominantly of the shattering wild type,
albeit with a sizeable minority of domesticated-type chaff
(cf. van Zeist and Bakker-Heeres 1985; Fuller 2007;
Stordeur et al. 2010). By its later phases, barley grains
show a significant increase in size compared to the wild-
type measured regional assemblages (Fuller et al. 2011).
Grain morphology indicates that emmer rather than einkorn
makes up the majority of the Triticum assemblage, but
wild- and domesticated-type spikelet bases have not been
positively identified and quantified (van Zeist and Bakker-
Heeres 1985). As regards pulses, lentils appear to be
enlarged and are thus considered to be of the domesticated
type (Fuller et al. 2011). Aswad has also produced an
arable weed flora, including about half of the diagnostic
taxa identified from Djade and Jerf el Ahmar (Willcox
et al. 2008; Willcox 2011).
Middle to late PPNB (late 9th7th millennia):
the emergence of crop packages
Although the assumption that the MPPNB of the southern
Levant witnessed the adoption of, and reliance on, domes-
ticated crop packages is well-established (cf. Kuijt and
Goring-Morris 2002; Simmons 2007), a closer examination
of the available botanical data reveals a more complexpicture. The impression created is one of diversity in the
choice of cultivars managed by different communities,
while wild-type crops also persist (Table 2). Beidha con-
tains such an idiosyncratic crop assemblage: while the
presence of naked barley implies that some barley was
certainly domesticated, also indicated by larger grain size,
chaff morphology suggests that cultivation focused on wild-
type barley (Helbk 1966, p. 355). Emmer includes a
mixture of wild and domesticated forms, deduced from
grain measurements (Feldman and Kislev 2007), while
Colledge (2001) has identified a single two-grained einkorn
grain that falls within the domesticated-type size range. Ofthe pulses, lentils demonstrate a change to larger average
diameters compared to earlier types, while chickpea is
outside its natural distribution. At Nahal Hemar, emmer
contained a mixture of wild and domesticated types (Feld-
man and Kislev 2007), while barley has been classed as
domesticated-type based on rachis morphology (Nesbitt
2002). Lentils were found in large quantities and showed a
shift to larger average diameters thus indicating their status
as a domesticated crop (Weiss et al. 2006). At Wadi Jilat 7,
barley rachises were predominantly of the wild type. Wheat
remains included both wild- and domesticated-type einkorn
grains alongside some domesticated-type spikelet forks that
have not been assigned to a particular species (Colledge
2001). Domesticated-type emmer grains are also reported
alongside arable weeds; the Jilat 7 botanical assemblage can
thus be considered as representative of the partial devel-
opment of the domestication syndrome (Colledge 2001).
Of the few MPPNB south Levantine sites with published
archaeobotanical assemblages, only Jericho contains
incontestable evidence of a mature domesticated crop
package and moreover one in which trends that began in
the PPNA appear to become fully articulated during the
PPNB: barley, emmer and (perhaps significantly in view of
its contested presence in the PPNA phase) 1-grained ein-
korn, flax, lentil and pea. At other MPPNB sites such as
Beidha we can observe the continuation of a mixture of
different PPNA traditions, with the cultivation of local
wild-type crops (barley, emmer) alongside pulses such as
lentil and pea. Yiftahel (Area C) is exceptional in this
respect, as it has provided evidence for the storage of
lentils and broad beans but none for cereal cultivation or
storage (Garfinkel 1987). The sole site other than Jericho
that contained domesticated-type barley is Nahal Hemar
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(Kislev 1988). While Jericho was a large (*10 acres)
settlement, Nahal Hemar was a cave site that contained
important indicators of ritual activities, being interpreted as
a sacred place and repository of ritual paraphernalia (Goren
et al. 1993; Bar-Yosef and Meadow 1995, p. 82).
This situation appears to have changed in significant
ways during the LPPNB. The sites of Ain Ghazal, Basta
and Wadi Fidan 1 are the sole examples of LPPNB southernLevantine megasites (large [8 ha densely populated
habitation sites found primarily along a north-south axis in
western Jordan) from which there are published archaeo-
botanical assemblages (for a general discussion of the south
Levantine megasites cf. Simmons 2007, pp. 175197.
Note that the stratigraphic resolution of the Ain Ghazal
botanical assemblages reported in the literature is very low.
Samples were retrieved for analysis during the first season
of excavation at the site (Rollefson et al. 1985). We there-
fore assume that the majority of charred macroremains
probably derived from LPPNB layers/contexts). These sites
have provided some evidence of mature domesticated croppackages (Table 2). Domesticated-type emmer and bar-
ley are found alongside lentils, peas and chickpea. Derived
(secondary) cereal domesticates (free-threshing wheat,
naked barley) that could reproduce only under full human
control were also present at Ain Ghazal (Rollefson et al.
1985). At Wadi Fidan 1 the assemblage is more cereal-
oriented with the presence of einkorn, emmer, barley and
free-threshing wheat. Developed arable weed floras have
also been reported (Colledge 2001). By contrast, in steppe
sites such as Wadi Jilat 13, barley appears to be predomi-
nantly of the wild type, while einkorn and emmer are
mainly of the domesticated type; in Azraq 31 cereals
(domesticated-type barley and free-threshing wheat) were
the sole recovered crops (Colledge 2001).
Discussion
The socio-ecological dynamics of plant domestication
The presence of domesticated-type crops is inferred from
the occurrence of tough rachises in the archaeobotanical
record. However, tough rachises near the base of the ear are
natural components in small proportions of wild cereals
(Kislev 1999) while metrical analyses have also demon-
strated that the transition to large grain size was a very
gradual process (Willcox 2004; Fuller et al. 2011). Thus our
approach to characterising south Levantine PPN botanical
assemblages has been a cautious one: full domesticated
status was assigned where non-shattering rachis remains
represented the majority of the assemblage, or they were
found at late sites alongside significantly enlarged grains.
Partial domestication status was assigned where non-
shattering rachises were present, but in a minority, or when
grains showed on average larger sizes compared to grains
retrieved from early sites. Such a measured approach pro-
vides a more realistic assessment of PPN crop management
strategies compared to simplistic distinctions drawn
between wild and domesticated crops that are some-
times uncritically considered as equivalent to agricultural
and gathering practices respectively.Constructed on these principles, Table 2 summarises the
current status of the south Levantine PPN archaeobotanical
assemblages arrayed by chronological order (expressed in
dates cal. B.C.), while Table 3 shows the region-wide spread
of domesticated crop packages. As it becomes evident from
the data presented in the previous sections, there is currently
no evidence pre-dating the MPPNB horizon for the culti-
vation of domesticated-type crop packages in the southern
Levant. This absence of evidence has been used by some
authors (Abbo et al. 2010; Honne and Heun 2009) for
proposing that, contrary to the polycentric origins argument
(cf. Willcox 2005), domesticated crops appeared during theEPPNB in the north, and that crop species were subse-
quently diffused as packages to other regions from their
agricultural homeland in southeast Anatolia.
From an archaeological viewpoint there are two lines of
argument with which such hypotheses can be addressed
(for detailed discussions of the evolutionary pace and
mechanisms, and the genetics arguments surrounding crop
domestication cf. Fuller et al. 2011; Allaby et al. 2010
respectively). The first has to do with the radiocarbon dates
and the second with the comparative analysis of the ar-
chaeobotanical data from each site. Table 3 presents in
summary form published archaeobotanical assemblages
and radiocarbon age ranges from PPN sites across south-
west Asia dating to the 9th7th millennia cal. B.C. (includ-
ing sites in Cyprus, Syria, central and southeast Anatolia,
the Zagros and the southern Levant). Three trends are
immediately evident which underline not only the validity
but, perhaps more importantly, the ecological and (hitherto
little investigated) socio-ecological complexity of the
polycentric origins argument:
(a) To date, the site of Kissonerga-Mylouthkia (KM) in
Cyprus contains the earliest evidence for the existence
of a fully domesticated crop package associated
with a weed flora anywhere in southwest Asia; all PPN
age determinations were obtained on grain, with the
earliest on a barley grain (Peltenburg et al.2003, p. 84).
The sum of calculated probabilities of the age deter-
minations at 1r places the beginning of KM earlier
than the mainland sites of Kerkh, Neval Cori, Cayonu
and Jericho. The causes for this early appearance of the
domestication syndrome in Cyprus may be attributed
to founder effects (cf. Lucas et al. 2011). While recent
Veget Hist Archaeobot (2012) 21:149162 155
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evidence for PPNA habitations on the island (McCart-
ney et al. 2010) theoretically leaves open the possibil-
ity of indigenous crop ancestry, the range of taxa
retrieved from KM casts doubt on the local origin of its
domesticated crop package. It seems far more likely
that it represents a characteristic example of short
gestation due to the dislocation of wild-type cultivars
from their natural habitats. Interestingly, the somewhat
later PPN site of Shillourokambos (although possibly
affected by issues of preservation) contained fewer
taxa including wild-type barley and domesticated-type
emmer. The Cypriot record thus indicates that the
ecological complexities of early PPN plant manage-
ment practices cannot be captured by unidirectional
Table 3 A comparison of archaeobotanical data for crop species and
fig presence in select assemblages across the region, with southern
Levant sites in bold (for an explanation of the symbols and the
derivation of C-14 age ranges see Table 2); Cayonu has a long
occupation through the PPN. The sum of 23 dates (excluding those
with large standard deviations and the basal level date, which is too
early) produces a strong modal peak at*8300 cal. B.C. which is
taken as the date for the early PPN finds of einkorn, emmer and pea at
the site; at Askl the majority of dates have derived from later Level
2, which was also the focus of the archaeobotanical analyses
published to date (van Zeist and de Roller 1995); at Catalhoyuk the
summed 1r probability of 65 dates (excluding those with large
standard deviations) provided an extended modal range of 71006400
B.C. The earliest (aceramic) levels of the mound have produced few
reliable dates by comparison to the ceramic Neolithic phases
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crop dispersal models that are based on putative crop
genealogies.
(b) It has become almost common place in the literature to
describe the Neval Cori and Cayonu botanical assem-
blages (Table 3) as the earliest examples of domesti-
cated crops by authors citing secondary sources (Abbo
et al. 2010; Simmons 2007, pp. 140141, both citing
Nesbitt 2002). In reality, however, domesticated-typecereal chaff represents a sizeable minority of the chaff
recovered from Neval Cori (Tanno and Willcox 2006;
contra Pasternak 1998; Haldorsen et al. 2011). The
same is the case for Cayonu where, in the absence of
better stratigraphic resolutionfor the botanical data, the
single strong modal peak observed at*8300 cal.
B.C. for its PPN phases is the sole date that can be
realistically assigned to the earliest occurrence of
domesticated-type einkorn, emmer and pea (Fuller
et al. 2011). In turn, this date corresponds well with the
end of PPNA Jericho (*8350 cal. B.C.)and is alsovery
close to the beginning of Jerichos PPNB phase(?8200 cal. B.C. if not older; for a detailed commentary
on the Jericho radiocarbon dates see Benz nd) In other
words, the chronological gap between Jericho and
Cayonu as regards the earliest appearance of domes-
ticated crops is more apparent than real. The same
argument appliesto Cafer Hoyuk (the earliest southeast
Anatolian site with evidence for a developed crop
package) which starts at*8300 cal. B.C. (Table 3).
(c) As discussed already, Jerichos PPNB botanical
assemblage appears to continue and amplify trends
that were already underway during the PPNA. Com-
pared to other MPPNB south Levantine sites, it is alsonoteworthy that while Jericho contains a well-devel-
oped crop package this does not seem to be the case
at Beidha, Yiftahel and Nahal Hemar. These sites
appear to continue local traditions of barley/emmer/
lentil management and cultivation, with the sporadic
presence of other pulses and flax, exclusive in the case
of Yiftahel. Domesticated crop packages do not
become widespread until the LPPNB, and then
predominantly in the context of the megasites. This
is a phenomenon that can be observed in other
southwest Asian regions as well as including sites
that do not always overlap chronologically, for
example the substantial communities of Askl, Abu
Hureyra 2A-C and Catalhoyuk. At the same time,
however, it does not register at other sites such as Ganj
Dareh and Jarmo in the Zagros area, despite their
chronological lateness that makes them good can-
didates for the introduction of developed crop pack-
ages from a hypothetical centre in the west. It
follows that crop packages cannot be considered as
the product of dispersal from a single centre
because their occurrence was chronologically diffuse
and, moreover, spatially discontinuous. Especially in
the southern Levant, non-local cereal crops were few:
one-grained einkorn (that does not appear to have held
a particularly prominent position in south Levantine
PPN cultivation systems) and two-grained einkorn
which was probably absent throughout the south (with
a few reported finds that might actually representsmall-grained emmer).
The longue duree of Epipalaeolithic and PPN plant
management strategies in the southern Levant
Established narratives represent the cultivation of domes-
ticated crops as one of the cornerstones of the Neolithic
Revolution which gave rise to new forms of social
organisation and set the foundations for the rise of social
complexity in later periods. Yet, by focusing almost
exclusively on the domestication process, the vexingquestion of the historical development of PPN plant man-
agement practices has been largely overlooked. What
insights could they provide for understanding the long-term
development of south Levantine PPN subsistence econo-
mies? That developed crop packages seem to have been
the exception rather than the rule before the LPPNB also
raises another important question: was plant domestication
the enabler or the outcome of population aggregation
manifested in the LPPNB megasites?
The available evidence permits outlining some pre-
liminary hypotheses, which although speculative at this
stage, should be further explored by future, problem-ori-
ented research. Epipalaeolithic plant management practices
do not emerge as pre-adaptations en route to food pro-
duction. They should be more realistically perceived as a
diverse array of foraging adaptations to locally fluctuating
plant resources. Cereals were not the focus of plant gath-
ering and are also unlikely to have been cultivated.
Potential causes are the high labour costs involved in field
preparation and crop processing which, combined with
delayed economic returns and the climate deterioration of
the YD during the Late Natufian, rendered intensive cereal
management a subsistence option that was both costly and
risky. This statement does not imply that cereals were not
used at all, only that they are unlikely to have been long-
term staple foods as is sometimes implied in the literature;
in all probability they represented a lesser component of
Epipalaeolithic diets that could have been adjusted when
available resources became scarce, as happened during the
YD. Overall the available evidence suggests that south
Levantine Epipalaeolithic plant management strategies
were probably flexible, being adapted to changing micro-
ecologies and local resource management histories.
Veget Hist Archaeobot (2012) 21:149162 157
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However, much more research is required for improving
our current understanding of plant gathering practices and
the nature of plant-people relationships during this period.
With the onset of the Holocene a long period of sub-
stantial climate amelioration began, which was character-
ised by higher temperatures and longer rainfall periods in
the annual cycle (Rosen 2007). Climate improvement led to
the proliferation and expansion of plant communities, whilehigher CO2 levels than in the Pleistocene increased edible
biomass available from wild seed plants (Cunniff et al.
2010). Nevertheless, net availability did not automatically
translate to intensification of plant management activities. It
was not until the later part of the early PPN that new
practices appeared at several sites, such as the collective
storage of clean crops (gathered and/or cultivated).
With the significant exception of Jericho, evidence for
MPPNB developed crop packages is limited. Despite
this scarcity of evidence, the MPPNB emerges largely as
continuing local early PPN traditions of plant management.
A tangible break with earlier practices can be discerned atthe LPPNB megasites with the widespread occurrence of
mature domesticated crop packages and the near disap-
pearance from their botanical assemblages of wild-type
crops (however, the quantity and quality of the available
archaeobotanical datasets from the southern Levant is
generally low with very few botanical assemblages
retrieved and studied at the required standard. For this
reason the interpretations offered here can only be viewed
as preliminary statements. Sampling and analysis must be
undertaken at more sites, including different site types, for
establishing a reliable picture of the nature and pace of
change in late PPN subsistence economies). This trans-
formation is broadly coeval with the transference of storage
and cooking facilities inside domestic spaces, amplifying
trends that were already under way during the MPPNB
(Wright 2000; Kuijt 2008a), not just in the southern
Levant, but also in sites in the north and in central Anatolia
such as Halula and Askl, and may also be correlated with
a change towards obtaining more meat from herded ani-
mals rather than hunted fauna (Vigne 2011, p. 177). The
contextual associations and spatial distribution of better
studied botanical assemblages from similarly large habi-
tation sites such as Catalhoyuk (*13 ha) in central Ana-
tolia have also indicated that crop storage and plant
consumption were private, household-centred affairs
(Bogaard et al. 2009). It is possible to hypothesise therefore
that population aggregation acted as an incentive for the
intensification of plant cultivation aimed at meeting the
nutritional needs of individual households, which took
the form of small-scale, intensive, household-operated
horticulture (sensu Bogaard 2005). In the case of Cat-
alhoyuk, aggregation manifested as the nucleation of
indigenous cultivator-forager communities (Baird 2006,
2010) that was accompanied by the adoption of several
crop and animal domesticates (Asouti 2006a). In the
southern Levant, a greater commitment to farming entailed
the intensified production of crops with a long history of
local management (emmer, barley, lentil) alongside pre-
viously introduced cultivars (einkorn, chickpea, broad
bean). Coeval with the intensification of cultivation was the
widespread adoption of goat and sheep pastoralism (Con-olly et al. 2011). The integration of crop packages with
animal herding brought about the establishment of highly
productive, mixed agro-pastoral systems (Harris 2002).
The timing of the appearance of domesticated crop
packages with population aggregation in large sedentary
communities suggests that it was this greater commitment
to farming that accelerated the pace of indigenous crop
domestication and the dispersal of non-local taxa. What
were the causes of south Levantine LPPNB population
aggregation in the first place is a vexing question that, to
date, has not found satisfactory answers, while the causes
of the decline and, in some cases, the abandonment of themegasites are also poorly understood (cf. Bienert et al.
2004; Simmons 2007). Environmental degradation seems
an improbable candidate (Goring-Morris and Belfer-Cohen
2010). Sites such as Ain Ghazal, appear to have been
sustainable with regard to their environmental footprint
(Campbell 2010 contra Rollefson and Kohler-Rollefson
1992). More plausible explanations are likely to emerge
from integrated contextual analyses of the southern Lev-
antine PPNB political economies encompassing plant and
animal management strategies, storage, land ownership and
territory definition, and the structure and organisation of
south Levantine PPNB households and communities.
Conclusions
Despite over five decades of fieldwork and data collection in
the southern Levant, the current state of the botanical evi-
dence does not permit us to provide region-specific answers
to key questions concerning the pace and mechanisms of the
development of the domestication syndrome that have
become central in archaeobotanical research in recent years.
This is due to the limited availability of sufficiently large,
quantified datasets from multiple PPN sites documenting the
evolution of key domestication markers such as large seed
size andshatteringhabit (cf.Fuller et al.2011). However, it is
possible to draw some preliminary conclusions, in the form
of hypotheses to be tested by future research, regarding the
development of plant management practices through time.
Overall, the available evidence suggests that cereal and pulse
crop progenitors were not intensively managed or cultivated
in Epipalaeolithic southern Levant. As the botanical finds
from Ohalo II indicate, cereals were harvested possibly in a
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pattern of opportunistic exploitation of foraging territories
that nonetheless did not take the form of sustained, long-term
dependence on cereals as staples. Climatic instability was
probably an underlying factor (cf. Richerson et al. 2001)
although the absence of evidence for intensive cereal
exploitation from the Early Natufian as well would seem to
suggest that, as a general rule, plant-based subsistence was
diversified rather than specialised. This observation alsoagrees with the finds of zooarchaeological investigations
indicating the persistence of south Levantine traditions of
broad-spectrum hunting (cf. Stiner and Munro 2002). Fur-
thermore, while the intensification of plant gathering activ-
ities in the YD is a possibility, it appears to have involved a
diverse array of taxa other than crop species progenitors.
The predominant pattern of PPNA plant management
practices has been increasingly described in recent years as
pre-domestication cultivation? gathering. However, to
date, there have been few attempts to define more precisely
the contribution of each component to subsistence econo-
mies. Recent work has suggested that cereals probablyplayed an important role in communal contexts (Kuijt and
Finlayson 2009). However, more and better studied
botanical datasets are needed for reconstructing with
greater precision the nature of gathering and cultivation
activities, and their linkages with different PPNA habita-
tion types (including multi-seasonal and short-lived/tran-
sient sites).
Despite their limitations, the available MPPNB datasets
suggest that whereas at sites such as Jericho domestication
trends were amplified, other communities such as Beidha
and Yiftahel followed divergent paths. The regional dis-
tribution of wild- and domesticated-type crops is not uni-
form. It is possible (empirical demonstration pending) that
the differences observed between sites may relate to hith-
erto unknown roles that plant production and consumption
played in the negotiation and reproduction of community
identities (cf. Asouti 2006b, pp. 112113). Such issues
have been the object of in-depth analyses focusing on
MPPNB burial practices and mortuary customs (Kuijt
2008b and references therein) but have not figured at all in
discussions of subsistence socioeconomics. For archaeo-
botanical studies in particular, higher-order interpretations
are hindered by the lack of reporting of the contextual
associations of plant remains. Such site by site reporting is
necessary for exploring specific practices associated with
plant consumption, storage and discard, and how they
might have related to other domains of social life. This
observation is valid for the PPNA and the LPPNB as well,
but its pertinence is particularly felt in the case of the
MPPNB, whereby archaeobotanical analysis has occurred
on a very limited scale and for very few assemblages
compared to the number of excavated and published sites.
Limited as it is, the available evidence suggests that
LPPNB south Levantine plant production was based on the
cultivation of crops with a long history of local manage-
ment extending back thousands of years. Levantine crop
packages themselves appear to have emerged as the
direct outcome of population aggregation, manifested in
the south Levantine megasites, that brought about the
intensification of subsistence production due to increasedcommunity size. Subsistence production primarily took the
form of intensive, small-scale, mixed agro-pastoral sys-
tems. Present data limitations do not permit examining
with greater spatial and temporal resolution how this pro-
cess unfolded before and during the lifetime of the south
Levantine megasites and what, if any, was its contri-
bution to their decline, or how it varied between different
site types and ecological zones. Much more fieldwork and
analysis are required before archaeobotanical research can
attain a contextualised understanding of the historical
development of early Neolithic plant cultivation in the
southern Levant.
Acknowledgments We wish to thank George Willcox for the
invitation to contribute a paper to this special issue of Vegetation
History and Archaeobotany. We also acknowledge the contribution of
the three anonymous reviewers whose constructive comments and
criticism improved the scope and content of our paper.
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