Asouti Fuller 2012

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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: [email protected]

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


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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.


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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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