New Evidence of Lateglacial Cereal Cultivation at Abu Hureyra on The

Home , Einkorn wheat, Emmer, Gordon Hillman, Tell Abu Hureyra, Tell Aswad

The Holocene 11,4 (2001) pp. 383–393

New evidence of Lateglacial cereal cultivation at Abu Hureyra on the Euphrates Gordon Hillman,1 Robert Hedges,2 Andrew Moore,3 Susan Colledge1 and Paul Pettitt2 (1Institute of Archaeology, University College London, 31–34 Gordon Square, London WC1H 0PY, UK; 2Research Laboratory for Archaeology and History, University of Oxford, 6 Keble Road, Oxford OX1 3QJ, UK; 3Ofﬁce of the Dean, College of Liberal Arts, Rochester Institute of Technology, 92 Lomb Memorial Drive, Rochester, New York 14623–5604, USA)

Received 1 August 2000; revised manuscript accepted 25 January 2001

Abstract: Hitherto, the earliest archaeological finds of domestic cereals in southwestern Asia have involved wheats and barleys dating from the beginning of the Holocene, 11–12000 calendar years ago. New evidence from the site of Abu Hureyra suggests that systematic cultivation of cereals in fact started well before the end of the Pleistocene – by at least 13000 years ago, and that rye was among the first crops. The evidence also indicates that hunter-gatherers at Abu Hureyra first started cultivating crops in response to a steep decline in wild plants that had served as staple foods for at least the preceding four centuries. The decline in these wild staples is attributable to a sudden, dry, cold, climatic reversal equivalent to the ‘Younger Dryas’ period. At Abu Hureyra, therefore, it appears that the primary trigger for the occupants to start cultivating caloric staples was climate change. It is these beginnings of cultivation in the late Pleistocene that gave rise to the integrated grain-livestock Neolithic farming systems of the early Holocene.

Key words: Agricultural origins, cereal cultivation, hunter-gatherers, domestication, palaeoclimate, rye, Abu Hureyra, Euphrates, southwestern Asia, Lateglaical, early Holocene.

Introduction lithic by a thousand years, and, for the moment, makes it the earliest known example of cultivation worldwide. (Note that all After more than two million years of obtaining food from the wild dates cited hereafter are in uncalibrated radiocarbon years before by gathering, hunting and scavenging, the development of farming the present – 14C yr BP.) propelled humankind into an entirely new trajectory. Worldwide, That this new evidence of early cultivation should come from the consequences for human society, demography, nutrition and the site of Abu Hureyra is no coincidence. So far, the site has ecology have been profound and often disastrous, and have proved unique in the detail of its continuous record of an in-situ moulded almost every aspect of modern life. transition from foraging to farming. It is also the ﬁrst site to pro- Decades of archaeological research and some thousands of pub- vide persuasive evidence of the pressures which seemingly drove lications have been devoted to exploring when, where and why the already sedentary hunter-gatherers to start the energy-expens- the shift from foraging to farming occurred. It appears to have ive business of cultivating their caloric staples. The evidence happened independently and at different times in at least three further reveals that the initial cultigens included rye, a cereal not (and possibly six or more) parts of the world. Evidence for some previously numbered among the ‘founder crops’ of southwestern of the earliest farming activities has come from southwestern Asia Asia (Zohary and Hopf, 1993). Yet, despite the present uniqueness (the ‘Near East’). Indeed, new evidence presented here from the of Abu Hureyra, we anticipate that equally early evidence of anal- Near Eastern site of Abu Hureyra now pushes back the beginnings ogous events will eventually be recovered from other sites in the of the systematic cultivation of cereals to before the end of the Fertile Crescent. Pleistocene, to ෂ13000 years ago (= 11000 years before the present (BP) in uncalibrated radiocarbon years; Stuiver et al., 1998). This predates the hitherto accepted start of the local Neo-

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Existing evidence for the start of same species (then much more widespread than they are today); cultivation in southwestern Asia and modifier genes. Clearly, the germination of spontaneously shed, wild-type spikelets will also have slowed fixation Finds of domesticates (Willcox, 1999). The process of cereal domestication materially alters the mor- As regards the first of these factors that could have decelerated phology of the plants concerned, and, in the archaeological record, domestication, it is perhaps inevitable that crops were, indeed, the start of cultivation is traditionally dated from the first appear- harvested while partially immature, because crops that are still in ance of remains of crop plants which clearly reveal such changes their wild state ripen very unevenly, and unripe harvesting would in their morphology. Hitherto, the earliest remains of domesticates have offered the most efficient means of avoiding heavy grain- in southwestern Asia have all been wheats (Harris, 1998; Garrard, loss through spontaneous shattering. This reinforces the 1999): Tell Aswad in the Damascus Basin yielded charred probability that there was, indeed, a period of ‘pre-domestication remains of emmer wheat of apparently domestic type from levels cultivation’ when cultivated crops continued to retain their wild- dated to ෂ9800 14C yr BP (van Zeist and Bakker-Heeres, 1979); type, brittle-rachised ears. This period might have been quite brief, Iraq ed-Dubb in northwestern Jordan produced domestic emmer but it could also theoretically have lasted several centuries. The or einkorn wheat from deposits dated to ෂ10000 14CyrBP fixation of other classic ‘domestic’ features such as large, plump (Colledge, 1994); and Tell Mureybit on the Middle Euphrates pro- grains in place of the skinny wild-type grains should theoretically duced two specimens of domestic emmer and another domestic have taken somewhat longer. Certainly, at the early Neolithic site wheat from levels dated to 10200–10000 14C yr BP, together with of Ç ayo¨nu¨, there was evidence that plump domestic-type grains two other domestic cereals of less certain date (van Zeist and probably did not emerge in local crops of emmer and einkorn Bakker-Heeres, 1986). Slightly later remains of domestic cereals until well after the fixation of semi-tough domestic rachis (van have been identified from sites such as Jericho in the Jordan Val- Zeist and de Roller, 1994). ley (Hopf, 1983), and from Cafer Hoÿu¨k (de Moulins, 1997) and Yet, if cereal crops retained their wild-type morphology during Ç ayo¨nu¨ (van Zeist and de Roller, 1994) in southeastern Turkey. the early stages of cultivation, how can we detect the point at However, the archaeobotanists who worked on these charred which cultivation first started? The archaeological evidence can remains stress that some of the grains could theoretically have come in at least four forms, including the following: trickled down from overlying, later levels, although at two of the (i) abrupt increases in remains of seeds of arable weeds sites the requisite overlying Neolithic levels are lacking. To be (‘segetals’) which cannot be explained by other forms of certain that the remains are of the same age as the levels where disturbance (Hillman and Davies, 1990b; 1992; Col- they were found, key specimens therefore need to be individually ledge, 1998); radiocarbon dated using accelerator mass spectrometry (AMS). (ii) indications of the heavy use of particular food plants by settlements in areas where extensive stands of such plants Evidence of pre-domestication cultivation could not have grown in the wild; There is also evidence from southwest Asia of some cultivation (iii) shifts in specific patterns of wear on flint sickle-blades having begun prior to the first definite appearance of these dom- (e.g., Unger-Hamilton, 1989; but see also Anderson, esticates (for discussion of the definition of terms such as culti- 1992); vation, domestication and agriculture, see Harris, 1989; 1990). (iv) archaeological evidence of concentrations of large, seden- This is only to be expected, as domestic traits would not necessar- tary populations that would not have been supportable ily have developed immediately cereals were first taken into culti- from hunting and gathering alone. vation. It is worth considering briefly how these ‘domestic’ traits become manifest. It is on one or more of these bases that pre-domestication culti- The first crops were sown from seed gathered from wild stands vation has tentatively been suggested at several early sites (Figure and were thus morphologically of the wild type, with brittle ears 1). Thus Colledge’s re-processing of the data from van Zeist and that shattered at maturity thereby allowing the seed to be dissemi- his colleagues’ analyses of the plant remains from Mureybit on the nated. In the absence of conscious human intervention, the culti- Middle Euphrates ranging in date from the Late Epipalaeolithic vation of wild-type wheats, barleys and ryes would have started (Mesolithic) to early Neolithic (ෂ10600 to 8900 14C yr BP) sug- to select for mutant tough-eared phenotypes of domestic mor- gests that, despite the wild-type morphology of most of the cereal phology only when these sown wild-type crops began to be har- grains, the associated seed remains actually come from weeds of vested by sickle-reaping or uprooting, in place of the harvesting arable cultivation (Colledge, 1998). Willcox has concluded the by beating that was favoured by most of the hunter-gatherers of recent times who foraged wild grass-grain as food (Bohrer, 1991; Harlan, 1992; Wilke et al., 1972). Such selection would also generally have required that the crops be sown on a new plot every year. Once sickle-reaping or uprooting was adopted, however, domestication in respect of the fixation of non-shattering (tough- rachised) ears in the crop population could theoretically have been complete well within a century (Hillman and Davies, 1990a). However, although it is possible to propose good reasons why the first farmers should have switched to harvesting by sickle- reaping or uprooting as soon as they started cultivating, it is possible that in fact such a switch was delayed, and that several centuries accordingly elapsed before the process of domestication could begin (Hillman and Davies, 1990b). Furthermore, even after these potentially domestication-inducing methods of harvesting were applied, several factors could have slowed dramatically the process of fixation of domestic traits. These include the effects of harvesting crops that were incompletely ripe; wet weather during Figure 1 The location of Abu Hureyra and other sites mentioned in the harvesting; introgression from nearby stands of wild cereals of the text.

Downloaded from hol.sagepub.com at Geographisches Institut on June 28, 2015 Gordon Hillman et al.: Early cereal cultivation at Abu Hureyra on the Euphrates, southwestern Asia 385 same for early Neolithic Jerf al-Ahmar (ෂ9750 14C yr BP) ent oak species allowed complementarity of mast cycles and (Willcox, 1998; Willcox and Fornite, 1999), while at early Neo- thence greater continuity of combined annual yields, the acorns lithic Netiv Hagdud in the Jordan Valley (ෂ9400 to 10000 14Cyr commonly served as a caloric staple; Mason, 1995b.) BP) the composition and archaeological context of plant remains Even the seed foods identified in the charred remains included has similarly prompted both Kislev and the excavator, Bar-Yosef, several that also seem to have served as staples, for example the to raise the possibility of pre-domestication cultivation there, too seeds of club-rush (Scirpus maritimus/tuberosus), Euphrates knot- (Kislev, 1997). grass (Polygonum corrigioloides), wild wheats and ryes (Triticum and Secale spp.) feather-grasses (Stipa spp.) and wild shrub-chenopods such as species of Anabasis, Hammada and members of New evidence from the site of Abu the tribe Camphorosmae. Significantly, each of these types of seed Hureyra food have also served as caloric staples among arid-zone hunter- gatherers in recent times. (Ethnohistorical examples are given in Moore et al., 2000: Chapter 12.) The remains of animal bones The results from a 25-year study of plant remains from the Epipa- also reveal an adundance of hunted animals, particularly gazelle laeolithic (Mesolithic) levels at Abu Hureyra not only accord with (Legge, 1996; Legge and Rowley-Conwy, 1987; Moore et al., the evidence from these sites but also have now tentatively pushed 2000: Chapter 13). The seasons of present-day local availability back the start of cultivation to substantially before the previously of each of the Ͼ100 seed foods and of the various animal species accepted start of the Neolithic. identified in the remains indicate that the hunter-gatherer settlement at Abu Hureyra was occupied year-round (Hillman et al., The settlement of Abu Hureyra 1997; Moore et al., 2000: Chapter 14). The prehistoric settlement of Abu Hureyra (35° 52Ј N, 38° 24Ј E) developed on a low promontory on the south side of the Euphrates The slide into aridity Valley, 130 km east of the present-day city of Aleppo. The first However, within 300–400 years of the foundation of the settle- settlement there, the Epipalaeolithic village of Abu Hureyra 1, ment, changes in the patterns of utilization of these food plants was inhabited from ෂ11500 to 10000 14C yr BP. The initial settle- recovered from Abu Hureyra indicate that the area was experienc- ment consisted of a cluster of semi-subterranean pit dwellings. ing a rapid slide into aridity (Moore and Hillman, 1992). This Once these had filled with occupation debris, they were succeeded seems to have paralleled a period of dry, cold conditions which by above-ground huts built of perishable materials. It has been has been observed in other parts of southwestern Asia in archaeol- estimated that, during the later centuries of occupation, the village ogical remains of wood charcoal (Baruch and Goring-Morris, of Abu Hureyra 1 was inhabited by 100–200 people (Moore et al., 1997), in pollen cores from lakes and mires (Baruch and Bottema, 2000: Chapter 5). In the following Intermediate Period (ෂ10000 1991; Bottema, 1995), from pollen in deep-sea cores (Rossignol- to 9400 14C yr BP), the settlement appears to have consisted of Strick, 1995; 1999), from varve sediments from Lake Van lightly constructed buildings similar to those of Abu Hureyra 1. (Landmann et al., 1996; Reimbold et al., 1996), from combi- This is attested by a modest deposit of occupation debris at the nations of mineral, diatom, pollen and molluscan remains from bottom of one of the excavation trenches. Continuity of occu- lakes in the Konya Basin (Kuzucuog˘lu and Roberts, 1997; Kuzuc- pation from Abu Hureyra 1 through the Intermediate Period is uog˘lu et al., 1999; Reed et al., 1999; Roberts et al., 1999; 2001), confirmed by a series of radiocarbon dates (Moore et al., 2000: from sediments from Lake Lisan (Yechieli et al., 1993), from Chapter 14). Following this intermediate episode of occupation, changes in the abundance and size of hunted animals (Davies, the settlement expanded rapidly during Abu Hureyra 2 times 1982), and from various other archaeological assemblages from (ෂ9400 to 7000 14C yr BP ). This village was composed of numer- the Levant (Bar-Yosef and Belfer-Cohen, 1992). This period of ous, rectilinear, multi-roomed family dwellings built of mud brick. aridity broadly coincided with the Younger Dryas climatic episode It is estimated that, at its apogee around 8000 14C yr BP, the (conventionally ෂ11000 to 10000 14C yr BP; Roberts, 1998) population of this village reached a maximum of 4000 to 6000 when, after several millennia of steadily increasing warmth and people, a direct consequence of the development of a farming way moisture, climate in several parts of the world abruptly reverted of life by its inhabitants. Large-scale flotation recovery was to colder, drier conditions (e.g., Bond et al., 1993; Harris, 2001; applied to virtually every deposit of each occupation level. In Kudrass et al., 1991; Roberts, 1998; Roberts et al., 1993). Recent almost every case it recovered vast numbers of plant remains. adjustment of the dates of the key pollen core from Hula in the Most of these were small, dense-tissued structures such as seeds Jordan Valley to take account of the hard water effect (Cappers, or wood fragments in which decay had been prevented through 2001) underlines the close chronological accord between the their having become charred on domestic fires. Younger Dryas episode seen in the Hula core and the start of seemingly parallel events at Abu Hureyra, even though Abu Hure- The first few centuries of occupation: a time of yra occupied a more marginal environment and could be expected plenty for hunter-gatherers to have experienced the effects of advancing desiccation some- The abundant plant remains from Abu Hureyra 1 reveal that the what earlier. wild plant foods gathered by the hunter-gatherers who settled the These changes associated with the Younger Dryas inevitably site ෂ11500 14C yr BP included Ͼ100 species of edible seeds had profound effects on the spatial and seasonal patterns of avail- and fruits gathered from: (i) the marshes and gallery-forest of the ability of plant and animal food resources, the more so because Euphrates Valley; (ii) two distinct subzones of the local terebinth- the changes followed a period of marked climatic amelioration. almond (Pistacia-Amygdalus) woodland-steppe; (iii) slightly more For the area around the northern Fertile Crescent, it has been poss- distant areas of oak-dominated park-woodland (Moore et al., ible to model the shifts in vegetation distribution as a set of maps 2000: Chapter 12). These seed foods and fruits were doubtless (Moore et al., 2000: Figure 3.18; see also Roberts, 1998: Figure supplemented by a range of edible roots and leafy foods which 5.2). These maps suggest that the changes in vegetation cover did not survive in identifiable form among the charred remains, were even more radical than hitherto assumed, and they underline and possibly also by acorns from the mixed oak park-woodland. the danger of using, in unmodified form, the present-day distri- (Virtually all recent hunter-gatherers with access to acorns appear bution of the wild ancestors of the first crops to indicate where to have used them as food; Mason, 1992; 1995a; but see also they were initially taken into cultivation, as in Lev-Yadun et al. McCorriston, 1994; and when overlapping distributions of differ- (2000) and, indeed, in other recently published proposals for the

Downloaded from hol.sagepub.com at Geographisches Institut on June 28, 2015 386 The Holocene 11 (2001) probable location(s) of the earliest agriculture. Caution in this regard has likewise been urged by Jones et al. (1998) in response to the arguments advanced by Heun et al. (1997). (See also Jones and Brown, 2000.) At Epipalaeolithic Abu Hureyra, the profound changes in vegetation distribution inevitably resulted in dramatic changes in the utilization of many of their wild foods. These changes in food use seem to have followed a distinct sequence. The ﬁrst to be affected was the use of fruits and seeds of drought-sensitive plants of oak- dominated park-woodland: after 300–400 years of the people of Abu Hureyra gathering these foods, they suddenly stopped using them (Figure 2a), probably because they no longer grew within easy reach. (Prior to the Younger Dryas, oak-dominated park- woodland is likely to have extended to within some 15 km of the settlement, i.e., within the foraging range for key seasonal foods of many recent hunter-gatherers; Moore et al., 2000: Figure 12.1.) These park-woodland foods included the edible seeds of an aspho- del (Asphodelus microcarpus type), and probably also acorns which could well have served as another caloric staple. Next, there was a seemingly marked decline in the use of wild lentils and other large-seeded legumes, and these then disappear completely (Figure 2b). (Their subsequent resurgence is discussed later.) Hitherto, these would have thrived in the dry fringes of nearby oak park-woodland and in the moistest zone of woodland- steppe adjacent to Abu Hureyra. The decline in their use probably represents the loss of local populations of many (maybe all) the large-seeded legumes and other food plants typical of these particular formations, including not only the lentils, but also the wild peas, vetches and some of the wild cereals. Certainly, Figure 2c shows the use of wild wheats and ryes seemingly peaking shortly after the wild lentils and then going into a similar overall decline, albeit with oscillations. Thereafter, wild wheats and ryes appear to have been used less heavily. The feather-grasses (Stipa spp.) are more drought-resistant than either the large-seeded legumes or even the wild ryes and wheats, and use of their grain correspondingly peaks later still, prior to then declining abruptly, as with the others (Figure 2d). The most drought-resistant of all the local staples were the shrubby chenopods, and the use of their seeds is correspondingly the last to peak, but even they eventually go into sharp decline (Figure 2e). By that point, local vegetation outside the Euphrates Valley adjacent to Abu Hureyra appears to have changed from a very moist form of woodland-steppe to a much drier equivalent, with arid, treeless steppe probably encroaching from the southeast. Nevertheless, the continued use of terebinth (Pistacia atlantica and/or P. khinjuk) nutlets – albeit in greatly reduced and ever- declining numbers (Moore et al. 2000: 341) and the continued presence of some terebinth charcoal (Roitel and Willcox, 2000: 545) both attest to the survival of at least a few terebinth trees somewhere within the site catchment. However, the overall pic- ture is one of sharply increased aridity. (The frequency distributions of all 150+ seed species found in Epipalaeolithic levels are presented in Moore et al., 2000: Figure 12.7; and all 16 of the wood-charcoal taxa in Roitel and Willcox, 2000: Figure A6.1.)

Figure 2 The decline in abundance of charred seed remains of five major groups of food plants from Abu Hureyra during a period coinciding with the Younger Dryas. (a) Seeds of food plants of oak-dominated park-woodland. (b) Seeds of large-seeded legumes (mostly lentils). (c) Grains of wild-type ryes and wheats. (d) Grains of feather-grasss. (e) Seeds of shrubby chenopods. The levels date from ෂ11500 14C yr BP at the bottom of phase 1 to ෂ10000 14C yr BP at the top of phase 3. In (b) to (e), the frequency of each species is expressed as a percentage of the total number of charred remains of seeds of dryland species recovered from the same levels. Because the five histograms of (a) represent much smaller numbers, they are not expressed as percentages, but rather as the numbers of seeds identified per 200 litres of excavated deposit (see Moore et al., 2000).

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Throughout this dramatic decline in resources from the park- had previously been the principal source of food. After all, wild woodland and woodland-steppe zones, the heavy use of two key cereals were just one of several apparent staples represented in foods from plants of the valley bottom continued without any sys- the charred remains, and there were probably also other, soft- tematic reduction. These foods were the nutlets of club-rush tissued staple foods such as wild roots and tubers (and perhaps (Scirpus maritimus/tuberosus) and of Euphrates knotgrass acorns, as well) that have left no identifiable trace. Rather, wild (Polygonum corrigioloides). Their continued use suggests that cereals were probably a critical seasonal resource during early to overbank flooding of nearby stretches of the Euphrates Valley middle summer, in the month or so preceding the ripening of grain occurred sufficiently often to allow both plants to continue to of the prolific, large-grained feather-grasses. grow in substantial quantities. This conclusion accords with the The choice of wild cereals as the first cultigens, rather than one recent suggestion by Reimbold et al. (1996) that the higher of the other wild caloric staples, reflects the fact that, of the vari- sediment-deposition rates they observed in Lake Van in East Ana- ous staples, they were the only ones amenable to annual culti- tolia during the Younger Dryas were due to factors such as sparse vation on a large scale in densely sown stands, they produce a vegetation and soil wash-out (linked with dry conditions) that heavy seed yield (unlike perennial equivalents), and were able to would have led to higher discharges into the local rivers, including produce an easily stored product in a single season. Cereals also the headwaters of the River Euphrates. yield a secondary product: straw, that is useful as tinder, thatching Given the decline (or complete disappearance) of most of the and bedding, and that would one day also prove useful as fodder other key plant food resources, one might ask why the inhabitants for domestic herbivores. did not simply leave Abu Hureyra for better locations. In fact, The areas chosen for cultivation would have been those with this option almost certainly did not exist. The Abu Hureyra area enhanced soil moisture, such as breaks in slope and shallow wadi- already provided the richest resources for both wild plants and bottoms not subject to spring spates. Yet while the soil moisture animals in this part of the Euphrates catchment, and other areas in such locations would have been sufficient to support wild cere- within reach were apparently experiencing precisely the same als, even during the Younger Dryas (in the same way that, in slide into increased aridity. Potentially more favourable areas in present-day arid steppe, such locations can often support crops the western Levant would already have been occupied by other such as cotton that normally require irrigation), competition from groups, who would not have welcomed immigrants, especially dense scrub that forms the natural vegetation in these locations with their own food supplies in decline. The people of Abu Hure- would have prevented wild cereals from establishing themselves yra were therefore forced to find a local solution. spontaneously. In consequence, wild-type cereals could have grown in these habitats only once cultivators had cleared the The start of cultivation scrub, tilled the soil, and sowed seed-corn which they themselves Their solution was, it seems, to cultivate certain of their wild cere- had gathered for this purpose from the ever more distant wild als, namely an annual wild rye (either Secale cereale L. subsp. stands. (Sowing autumn-germinating crops such as these primitive vavilovii (Grossh.) Zhuk. or S. iranicum Kobyl.) and probably also ryes and wheats in the Euphrates Valley bottom would have been a wild wheat. In the succession of losses of staple wild foods from rendered pointless by the risk of over-bank flooding through win- the Abu Hureyra menu, the first indications of cultivation come ter and into the spring. That crops were not, indeed, sown on the soon after the decline in wild cereals. The evidence for the start valley floor at this time is confirmed by the absence of valley- of cultivation at Abu Hureyra is as follows. bottom species in the ascendent weed flora.) (a) Despite the worsening aridity, from shortly before ෂ11000 In the centuries following the first appearance of domestic-type 14C yr BP seeds of drought-intolerant weeds characteristic of rye grains, with the Younger Dryas still maintaining its grip, we local, rainfed, arable cultivation suddenly increased from the rela- continue to encounter charred remains of rye grains of the same tively low levels they had maintained for four centuries hitherto. domestic morphology, together with remains of the lentils. By (b) Wheats and ryes grains of a skinny morphology resembling the Intermediate Period (ෂ10000 14C yr BP), however, when the the wild type continued to be used (albeit now in much smaller Younger Dryas had just ended, the rye and lentils had been joined quantities), despite the dry conditions of the Younger Dryas hav- by other domesticated cereals such as domestic einkorn. By the ing almost certainly eliminated all wild stands from the area. time the Neolithic was fully developed at the settlement (by (c) Soon afterwards, we encounter the first rye grains of the ෂ8500 14C yr BP), these three crops had been joined by domestic domestic type, Secale cereale L. subsp. cereale Zhuk., and rye emmer, bread wheat, macaroni wheat, barley, chickpeas and field grains of this type continue to occur thereafter. (Although the pre- beans (de Moulins, 1997; 2000). All these latter crops had preceding four centuries-worth of deposits yielded wild-type rye sumably been taken into cultivation and domesticated at one or grains in considerable abundance, there had been no trace of any more (as yet unidentified) locations elsewhere. domesticates. The features distinguishing the domestic grains from their wild progenitors are indicated in Figure 4.) (d) Two or three centuries later, lentils and other large-seeded Detailed evidence for the start of legumes suddenly reappeared and started to increase in abundance cultivation (Figure 2c), this despite the fact that (i) they had, by then, been locally absent for some centuries, and (ii) the arid conditions Having now considered the events at Abu Hureyra in overview, which probably caused their disappearance still prevailed and it is appropriate to examine the three main lines of evidence for would have prevented any wild stands re-establishing themselves the start of cultivation in more detail, and then to consider their in the area. significance. In combination, the four lines of evidence seem to suggest that some of the occupants of Abu Hureyra had probably started to The rise of arid-zone weed floras cultivate – first cereals such as rye and wheat, and second some While the use of most of the major wild caloric staples at Abu large-seeded legumes including lentils. The fact that the apparent Hureyra was declining (or, in the case of the two valley-bottom start of cultivation shortly before ෂ11000 14C yrs BP coincided staples, remaining unchanged), seed remains of three other with a marked overall reduction in the use of wild wheats and important groups of plants started to increase abruptly from the ryes further suggests that it was probably major losses of wild start of phase 2, shortly before ෂ11000 14C yr BP These plants stands of these cereals which initially triggered their cultivation. (Figure 3) were the small-seeded legumes (c. 20 species of Such a conclusion does not necessarily imply that the wild cereals Trigonella, Medicago and Trifolium), small-grained grasses

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remains of small-seeded legumes and grasses at Abu Hureyra from shortly before ෂ11000 14C yr BP merely represented rather desperate, intensified gathering of these low-quality foods from their last surviving stands to compensate for the loss of the pre- ferred caloric staples (Hillman et al., 1989). However, the parallel increase in the remains of stony-seeded gromwells cannot be explained on this basis. The fact that all three groups represent the classic weeds of rainfed cereal cultivation of gypsiferous soils in the steppe of central Syria suggests a more convincing explanation for their synchronous and abrupt increase, namely that they were invading new areas of cultivation near Abu Hureyra, and that their presence on the site is attributable to casual arrival with the harvested crops. If they had not been growing as weeds of crops sown in areas of enhanced soil moisture, the increasing aridity of the native woodland-steppe around Abu Hureyra should actually have caused these particular plants to decline in abundance rather than to increase. After all, they were originally all natives of moist woodland-steppe where they can still be found today (albeit with the tree components of this formation often now absent due to recent felling). Although we now regard them as typical weeds of local ‘arid-zone’, rainfed cultivation, they are insufficiently drought-tolerant for their weedy, colonizing tendencies to have allowed them to replace any of the other plants of woodland- steppe that were disappearing due to the drought conditions of the Younger Dryas. In this regard, their ecology contrasts with that of weeds such as some species of poppy (Papaver spp.) which, in pristine native floras (i.e., in areas without any cultivation or other forms of heavy disturbance), achieve their greatest abundance in full-blown, treeless steppe, and which Neil Roberts (personal communication, 1999) notes to have increased during the Younger Dryas in many pollen diagrams without any indi- cation of an anthropogenic explanation. The sudden increase of these three particular groups of weeds at Epipalaeolithic Abu Hureyra is thus explicable only by the removal of competing vegetation from habitats with enhanced soil moisture combined with disturbance sufficient to fragment the hard crust of the local gypsiferous soils. Domestic animals could not have been the source of this disturbance, as sheep and goats, the first apparent domesticates at Abu Hureyra, did not arrive until ෂ9400 14C yr BP, and domestic cattle and pigs later still (Legge, 1996; Moore et al., 2000: Chapter 13). Similarly, the disturbance cannot be attributed to a shift to year-round occupation of the site because we have seen that some, at least, of the Abu Hureyra population had already been sedentary for the preceding 400 years. Scrub clearance and tillage associated with cultivation in areas of enhanced water availability (from which competing vegetation had been cleared) seems to offer the only satisfactory explanation. Once harvested, the crops with their admixture of weeds would have been threshed and winnowed. The unwanted weed seeds Figure 3 Changes in the relative abundance of seed remains of three groups of classic weeds of arid-zone cultivation during the Epipalaeolithic were probably then sieved or hand-sorted from the cereal grain (Mesolithic) Period at Abu Hureyra: (a) charred seed remains of small- and discarded on domestic fires, where a proportion got preserved seeded legumes; (b) charred remains of small-grained grasses; (c) silicified by charring. Certainly, discarded weed seeds separated by sieve- seeds remains of dryland gromwells. As in Figure 2, the levels date from cleaning, together with skinny ‘tail grains’ from the host crop, ෂ11500 14C yr BP at the bottom of phase 1 to ෂ10000 14CyrBPatthe dominate the charred remains on many later archaeological sites top of phase 3. In (a) and (b), the seed frequencies represent percentages (Hillman, 1984; Jones, 1987). (‘Tail grain’ comprises the under- of all dryland plants preserved by charring (i.e., with the very abundant sized grain that passes through the grain-cleaning sieves together silicified seeds of the gromwells excluded); in (c), they represent percent- with the small weed seeds, and gets discarded with this waste ages of all seed remains – both charred and silicified. fraction.)

(Hordeum bulbosum and species of Eremopyrum and the H. muri- Continued use of cereals during the driest parts of num aggregate) and stony-seeded dryland gromwells (Arnebia and the Younger Dryas Buglossoides spp. of the borage family). Cereal cultivation is also suggested by the continued use of wheats Our ecological studies over the past 25 years indicate that these and ryes (albeit at now reduced levels) after the point at which plants would originally have been natives of the local, moist the evidence suggests that wild stands of these species could no woodland-steppe. We initially considered that the increase in the longer have grown in the area. Certainly, if the aridity was

Downloaded from hol.sagepub.com at Geographisches Institut on June 28, 2015 Gordon Hillman et al.: Early cereal cultivation at Abu Hureyra on the Euphrates, southwestern Asia 389 sufficient to drastically reduce the much more drought-resistant of them is illustrated in Figure 4 alongside a typical ancient grain feather-grasses and, thereafter, the even more drought-resistant of wild-type rye for comparison. (Further grains are illustrated in shrubby chenopods, then it will certainly have eliminated most of Moore et al., 2000: Figures 12.6 and 12.23.) the wild wheats and ryes. More specifically, our models of vegetation that could potentially exist in the present day in the absence of deforestation, cultivation and grazing by domestic ani- AMS 14C dating of the cereal grains mals indicate that, even under current conditions, no extensive stands of wild wheats or rye could grow within 60–70 km of Abu Because only a small number of grains of the domestic type sur- Hureyra (Moore et al., 2000: Figure 3.18). In the middle of the vived in the charred remains from the Epipalaeolithic occupation, Younger Dryas, conditions in southwestern Asia appear to have it has been essential to establish that they had not worked their been significantly drier than those of the present day (Baruch and way down from overlying Neolithic levels. Key specimens were Bottema, 1991; Baruch, 1994), so the nearest stands of wild cere- therefore dated individually by AMS 14C dating. als would have been even farther from Abu Hureyra. The con- Of the five domestic-type rye grains AMS dated so far, three tinued presence and use of wheat and rye at this time therefore have given Epipalaeolithic dates matching other dates from the suggests that they were actually under cultivation, whether as levels in which they were found, namely 11140 Ϯ 100 14CyrBP brittle-rachised wild types, or as tough-rachised domesticates (OxA-8718); 10930 Ϯ 120 14C yr BP (OxA-6685); and 10610 whose grains were mostly still skinny and resembled those of the Ϯ 100 14C yr BP (OxA-8719). The fourth grain was dated to Abu wild types. Hureyra’s Intermediate Period: 9860 Ϯ 220 14C yr BP (OxA- 6996). The fifth proved to be Neolithic in age: 8275 Ϯ 65 14Cyr The presence of domestic cereals Identifiable remains of domesticates make their first appearance at Abu Hureyra 1 in phase 2, ෂ11000 14C yr BP, a century or so after the reduction in use of wild-type cereals, and the abrupt rise in weed floras. Domesticates can be distinguished from wild types using grain size and shape, and the presence of a rough abscission scar at the rachis node on remains of subunits of the ear termed ‘spikelets’. (A rough abscission scar implies that disarticulation of the ear into its component spikelets did not occur spontaneously, as it does in wild ears, but had been induced by threshing; e.g., Kislev, 1992; Zohary, 1992; Hillman and Davies, 1990a; 1992.) It could be argued that, because such features are to some degree phenotypically plastic, they are unreliable indicators of domestication. For example, the expression of rachis brittleness during threshing is affected by the state of maturity and wetness, and the completeness of grain fill (and thence grain breadth and thickness, but less so grain length) by soil moisture, soil fertility (peripherally), and, more exceptionally, by reducing the ratio of photosynthate sink to source, for example by destroying part of each ear (Heun, personal communication, 1998). However, factors of this sort seem unlikely to account for the sudden appearance of exceptionally fat, long domestic-type grains in the Abu Hureyra remains from 11000 BP onwards, after a complete absence of such grains among the large numbers of wild-type grains recovered from the preceding 400 years’ worth of archaeological deposits. Similarly, the sudden appearance of the domestic-type grains cannot be attributed to the fact that, like most grain remains from archaeological sites, those from Abu Hureyra were preserved by charring. Charring certainly affects grain shape, but while it generally increases the breadth and thickness of cereal grains, it simultaneously reduces their length (Hopf, 1955). It cannot therefore Figure 4 Examples of charred remains of rye grains from Epipalaeolithic account for the domestic-type grains being massively longer (as (Mesolithic) levels at Abu Hureyra drawn to the same scale: (a) a typical well as broader and thicker) than any of the large numbers of grain of wild rye, Secale cf. cereale L. subsp. vavilovii (Grossh.) Zhuk., wild-type grains found in earlier levels. The sudden appearance from phase 1 which pre-dates the earliest evidence of cultivation; (b) a of such grains from 11000 BP onwards is much more likely to be typical example of one of the nine grains of domestic-type rye, S. cereale the result of selection under cultivation, although in populations as L. cf. subsp. cereale Zhuk., found in phases 2 and 3 of the small (and as isolated from wild stands) as those probably sown Epipalaeolithic. This specimen comes from slightly higher levels than the as the first crops at Abu Hureyra, the effects of genetic drift cer- wild specimen (a), and is AMS 14C dated to phase 2 of the Epipalaeolithic 14 tainly cannot be discounted (Martin Jones, personal communi- occupation, with a date of 10930 Ϯ 120 C yr BP (OxA-6685). The cation, 2000). That selection under cultivation was probably a piv- domestic-type grains differ from the wild types primarily in being significantly larger not only in breadth and thickness but also in length. As such, otal factor is further suggested by the fact that, from 11000 BP the domestic-type grains were instantly recognizable among the mixture onwards, domestic-type grains of similar morphology continue to of smaller grains with which the earliest specimens were found, and were occur right through into the fully developed Neolithic period when clearly distinguishable from the mass of wild-type grains which were the agriculture was indisputably well established. only forms present in all the preceding levels. (For drawings of more of We have so far found nine of these massive, domestic-type rye the charred specimens of wild and domestic-type grains recovered from grains in phases 2 and 3 of Epipalaeolithic Abu Hureyra 1. One Abu Hureyra, see Moore et al., 2000: Figures 12.6 and 12.23.)

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BP (OxA-5843). These last two grains were clearly intrusive from and its eventual irreversibility, this systematic cultivation of layers overlying the Epipalaeolithic deposits. However, all five caloric staples would have been altogether different from any earl- grains shared the same typically domestic morphology. ier forms of ecological intervention such as the small-scale sowing The four other grains of domestic-type rye from these early of non-staples, the occasional planting of food-producing trees levels have so far been withheld from 14C dating because it is such as oaks, the tending of stands of wild food-plants by weeding possible that they have been chemically contaminated by out less favoured species, or landscape management using fire or adhesives (silver dag) used earlier in the project during scanning drainage to enhance food yields from wild species. Palaeo- electron microscopy (Colledge, 1988). Two of the four were also environmental/archaeological evidence suggests that a range of contaminated by solvents used in infrared spectroscopy (McLaren such manipulations had already been applied by hunter-gatherer et al., 1990; Hillman et al., 1993). However, alternative forms of populations in various parts of the world for many millennia (e.g., chemical pre-treatment might allow them to be dated in due Harris, 1989), but that they had not led inexorably to the large- course. scale cultivation of caloric staples and fully fledged agriculture, Each of the five grains of domestic rye that have been dated no more than such practices had among the many hunter-gatherers so far (as with the 27 wild-type grains that have also been dated) who adopted similar strategies in recent times. It was only when were chemically treated to remove potential organic contaminants people were forced, as at Abu Hureyra, to start the systematic of a different radiocarbon content, then oxidized to CO2, and their cultivation of caloric staples that the evolution of agrarian and, 14C/12C content measured by accelerator mass spectrometry using eventually, urban society was set in motion.

CO2 as the ion source substrate. Charred seeds may easily dissolve in alkali, so this step in the acid-base-acid treatment to remove Why cultivate rye? contaminant humics was omitted, and replaced with prolonged Once they had been driven to start cultivating, why did they ultrasonication in hot water. (Other work on this site has shown bother with rye when wheat was available? In fact, compared with that humic components in seeds and burnt bone have a similar wild and primitive domestic wheats, rye offers several advantages. radiocarbon content to that of the original material; Battern et al., Even in the wild state, rye grain is much more easily threshed 1986.) Sample OxA-6685 was first treated with a from the spikelets, and the chaff more easily winnowed from the methanol/acetone mixture to extract possible suspected paraffin grain; the wild rye grain is less glassy than those of the wild and detergent contamination. wheats and so more easily prepared as food, whether by roasting, grinding to flour, or boiling and mushing; and the ease with which it can be mushed would have made it attractive as a weaning food. Discussion Also, once eaten, the starch of rye foods is hydrolyzed to sugars more slowly, and it correspondingly produces a lower insulin Systematic cultivation of potential staples on a scale that makes response than does the starch of equivalent wheat products pre- a significant contribution to caloric need involves enormous effort. pared in the same way and consumed in the same combinations. Thus, although cultivation can yield more calories per unit area Rye-based foods are therefore more sustaining (Leinonen et al., of land than the gathering of equivalent staples from the wild, it 1999; Miller et al., 1996). These advantages might also account simultaneously involves much higher energy inputs per unit of for the apparently prominent role of rye at a slightly later site, caloric return, and a corresponding loss of leisure time. It probably Tell Mureybit, upstream from Abu Hureyra, as reported by also disrupted the peoples’ ethical/spiritual perceptions of their Willcox and Fornite (1999). relationship with their environment, disrupted their concepts of Furthermore, once the rye was domesticated, additional advan- ownership of resources and rights of access, and required a radical tages would have become apparent: combined with its loose chaff, revision of division of labour within (and possibly between) the fully tough rachis of even the most primitive domestic ryes families. would have allowed grains to be freed by a single threshing, and The systematic cultivation of caloric staples such as cereals is the straw and chaff to be eliminated by a single round of win- therefore likely to have been adopted only in response to stress, nowing. Such a cereal is termed ‘free-threshing’. Present chrono- and at Abu Hureyra we have seen that the trigger was probably logies suggest that equivalent free-threshing forms of wheat the sharply reduced availability of local stands of wild cereals due evolved only after a further 2500 years of farming. (The planned to Younger Dryas aridity, following probable earlier losses of AMS dating of single grains of bread wheat from Intermediate other staples such as acorns, the seeds and roots of asphodels, and Period Abu Hureyra and from levels of equivalent age, ෂ10000 further food-plants of oak park-woodland. Their decision to start 14C yr BP, at Mureybit will probably show them to have intruded cultivating the wild-type cereals was then further reinforced (and from later levels.) These advantages of rye are likely to have been made more irreversible) by the ensuing sharp reductions in other obvious to all. Its Achilles’ heel, however, may eventually have wild staples such as grain from the feather-grasses and, thereafter, been its vulnerability to any early onset of summer drought during seed from the shrubby chenopods. its relatively slow maturation (Blumler, 1996), its susceptibility As hunter-gatherers, their susceptibility to the climatically to infection by the ergot fungus when sown on damp soil, and induced changes in their wild food resources would also have the vulnerability of its grain to predation by rodents, even while been exacerbated by the fact that they occupied the site year- it is still ripening in the ear. round and appear to have done so ever since the site was first In addition, once rye was domesticated and grown more widely, settled ෂ11500 14C yr BP. (Hillman et al., 1997; Moore et al., its outbreeding behaviour (if established at that stage) would have 2000). Compared with more transient occupation, year-round sed- facilitated the unwanted re-introduction of agronomically entism is likely to have entailed more intensive utilization of the disadvantageous wild-type characteristics from any wild stands of local wild resources and correspondingly narrower safety margins. outbreeding rye that had re-colonized the region following the Furthermore, if (as is generally assumed) sedentism prompted fas- return of moister conditions at the start of the Holocene. We have ter population growth, then these safety margins would already found that, even with a nominally cleistogamous in-breeder such have been getting progressively narrower for the preceding four as barley, introgression from wild stands into cultivated crops centuries. Thus, when most of the intensively exploited wild food growing in Turkey regularly occurs over distances in excess of 1 resources were eventually decimated by advancing aridity, sea- km and produces crops with a conspicuous component of brittle- sonal shortages were inevitable. rachised ears, even in six-rowed forms. With outbreeders such as It should be noted that, in its profound effects on human society most modern domestic ryes (or the maize involved in the GMO

Downloaded from hol.sagepub.com at Geographisches Institut on June 28, 2015 Gordon Hillman et al.: Early cereal cultivation at Abu Hureyra on the Euphrates, southwestern Asia 391 debate), introgression of this intensity is likely to occur over much trickle of additional families succumbing to the pressure to culti- greater distances. vate as drought conditions progressively eliminated the last stands The debut of grains of domestic-type rye so soon after the start of wild cereals still surviving within foraging range in the hilly of cultivation (as indicated by the rise of segetal weeds) suggests hinterland. Combined with the series of disappearances of wild that there was only a very brief period of pre-domestication culti- foods, the explosive expansion of cultivation, together with its vation, and this poses certain problems. Most of the modern forms effect in further restricting mobility, would have sharply increased of domestic rye are predominantly outbreeding, and, if the wild dependence on the narrow range of cultivable foods (notably the ancestor was also outbreeding (as are at least some modern popu- cereals) and led to a corresponding collapse of overall dietary lations of Secale cereale L. subsp. vavilovii (Grossh.) Zhuk.) then, diversity. Certainly, even for the seed foods, the small number of without conscious selection, domestication would theoretically plants (perhaps 7–8) which eventually proved amenable to culti- have taken several millennia (Hillman and Davies, 1990b). The vation contrasted starkly with the 100+ seed foods hitherto gath- rapid emergence of domestic rye at Abu Hureyra therefore sug- ered by their hunter-gatherer forebears. gests that: In this narrowing of dietary diversity, it was cereal consumption (a) some decades of unusually hot summers had induced self- that played the key role. Overall, the dietary role of cereals seems fertilization (Gertz and Wricke, 1991) and therefore allowed likely to have expanded dramatically from that of seasonal staple more rapid fixation of any recessive alleles advantaged by to dominant staple year-round. Such heavy consumption of the harvesting regime; or cereals has psychoactive and other physiological effects that (b) the populations of wild rye that provided seed-grain for the would have further reinforced the selfsame cereal dependency. first crops were largely inbreeding in any case, rather like When cereals are consumed in quantity, exorphins can apparently the wild Secale iranicum Kobyl. (Kuckuck and Kranz, 1957; activate reward centres in the brain and produce a sense of well- Jaaska, 1998; Kobyljanskij, 1989); and/or being that tends to produce cravings for continued regular con- (c) conscious selection was applied sooner than we had hith- sumption (Wadley and Martin, 1993). Perversely, equivalent crav- erto supposed. ings for cereals are reported in cases of physiological intolerance of the gliadins of the Triticeae (the grass tribe that includes wheat, On into the Holocene: the evolution of complex rye and barley), for example, in some patients suffering with cel- agro-pastoral systems, and a growing obsession with iac disease (Blostoff and Gamlin, 1989). While we would argue cereal consumption against the suggestion of Wadley and Martin that the effects of At Abu Hureyra, the transition from the exclusive use of wild exorphins drove the very inception of cereal cultivation, we accept foods to eventual dependence on cultigens as the principal sources that they would have reinforced its irreversibility. of starch took over 2500 years to complete. More specifically, plant remains from ensuing Neolithic levels reveal that small quantities of three of the erstwhile wild caloric staples (the seeds Summary of club-rush, Euphrates knotgrass and feather-grass) continued to ෂ 14 be used until 8500 C yr BP, albeit on an ever-diminishing scale So far, Abu Hureyra provides the clearest, continuous record (de Moulins, 1997; 2000). anywhere in the world of an in-situ transition from foraging to The terminal stages of the waning use of these erstwhile wild farming, although eventually other such sites will doubtless be staples (together with the piecemeal abandonment of most of the found. The evidence suggests that, on the Middle Euphrates (and remaining secondary wild starch foods), probably reflects not just almost certainly elsewhere in southwestern Asia), these transform- the now reduced need for starch sources from the wild but also ations were set in train before the end of the Pleistocene, in a constraints on foraging resulting from: (a) the heavy time and climatic context different from that supposed hitherto, and among energy inputs involved in crop husbandry, which not only directly hunter-gatherers who were already largely sedentary. Here, at limited the time available for foraging but probably also curtailed least, the primary trigger appears to have been the critically the longer hunting forays and, with them, access to those wild reduced availability of key wild plant staples during the arid con- foods hitherto gathered from more remote areas of the erstwhile ditions of the Younger Dryas climatic episode. This early incep- economic catchment; (b) the erosive effect (on the local survival tion of cultivation then set the scene for the development and of these wild food-plants) of tillage, the maintenance of mono- rapid spread of integrated agro-pastoral economies in the early cultures and, eventually, the emergence of pastoralism and ever Holocene. While the advent of agriculture is often hailed as a more intense grazing pressures on local vegetation; (c) the evapor- triumph in ‘laying the foundations of civilization’, the social, ation of knowledge of the identity, ecology and methods of gath- demographic, nutritional and ecological consequences of the ensu- ering and processing of wild food species that were no longer ing chain-reaction have mostly proved disastrous, and continue to regularly encountered; and (d) changes in ecological and spiritual mould almost every aspect of modern life. perceptions concerning the place of humans in (and their identification with) their natural environment. By ෂ8500 14C yr BP use of the wild starch foods had seemingly ceased altogether, cultivation had become part of a complex Acknowledgements agricultural socio-economic system incorporating the rearing of livestock, and burgeoning farming populations were expanding We wish to acknowledge, first, the role of David Harris as the then in all directions, irreversibly eliminating former hunter-gatherer Head of Department of Human Environment of UCL’s Institute of lifeways and the ecosystems of which they had formed a part. Archaeology in facilitating (and participating in) a critical phase Nevertheless, although the ‘tail end’ of the use of erstwhile of the laboratory and field studies; second, the influence of valu- wild staples such as the feather-grasses, club-rush and knotgrass able discussions over many years with Daniel Zohary, Frances took more than two millennia finally to disappear, the steep rise McLaren, David Harris, Dominique de Moulins, George Willcox, in weed floras described above suggests that, as soon as cereal Tony Legge, Peter Rowley-Conwy, Andrew Garrard, Uri Baruch, cultivation was initiated, it expanded very rapidly and that this Neil Roberts, Ann Butler and Terry Miller. We are also grateful expansion continued apace for some centuries. This rapid expan- to Derek Roe, Neil Roberts, George Willcox and Martin Jones sion probably represents the first families of cultivators growing for kindly editing the script and proposing many improvements. ever larger areas of cereals year-on-year, as well as a steady Work on the plant remains from Abu Hureyra and related plant-

Downloaded from hol.sagepub.com at Geographisches Institut on June 28, 2015 392 The Holocene 11 (2001) ecological fieldwork was supported by a research grant from the —— 1990: Settling down and breaking ground: rethinking the Neolithic British Government’s Science and Engineering Research Council Revolution. Stichting Nederlands Museum voor Anthropologie en Praehis- and by major portions of four research grants from the Natural torie, Twaalfde Kroon-Voordracht, Amsterdam, The Netherlands. Environment Research Council, and the University of London’s —— 1998: The origins of agriculture in Southwest Asia. The Review of Gwynne-Vaughan Trust and Central Research Fund. Archaeology 19, 5–11. —— 2001: Climatic change and the beginnings of agriculture: the case for the Younger Dryas. In Lister, A., and Rothschild, L., editors, Evolution on Planet Earth, New York: Academic Press. 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