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Abstract

This article reviews the available information on the founder grain crops (einkorn wheat, emmer wheat, barley, lentil, pea, chickpea, and flax) that started agriculture in Southwest Asia during the Pre-Pottery Neolithic period, some 11,000–10,000 years ago. It provides a critical assessment for recognizing domestication traits by focusing on two fields of study: biology and archaeobotany. The data in these fields have increased considerably during the past decade, and new research techniques have added much to our knowledge of progenitor plants and their domesticated derivatives. This article presents the current and accumulated knowledge regarding each plant and illustrates the new picture that emerged on the origin of agriculture.

Introduction

The occasion of the Wenner-Gren conference “The Origins of Agriculture: New Data, New Ideas” raised our awareness of the importance of critical, high-quality data. We suspected that the newly obtained data sets from well-dated archaeological contexts may generate new hypotheses concerning the first steps of the agricultural revolution in Southwest Asia.

For this purpose we concentrated on two domains: the newly obtained rich archaeobotanical assemblages and the molecular analysis of present-day accessions. Some of these new data are inconclusive for pinpointing the shift from wild to domesticated plants because of bad preservation of the archaeological samples and problematic field or lab procedures. Following the fruitful discussions during the conference, we critically reassessed the available archaeobotanical data in an attempt to “separate the grain from the chaff.” In the following pages we present the data sets, both old and new, that we consider to be reliable enough to indicate the first appearances of domesticated forms in the Fertile Crescent.

Eight plants are considered to be the domesticated founder crops in the Levant (i.e., the western “arc” of the “Fertile Crescent”; table 1). This assemblage includes three cereals (einkorn wheat Triticum monococcum, emmer wheat Triticum turgidum subsp. dicoccum, and barley Hordeum vulgare), four pulses (lentil Lens culinaris, pea Pisum sativum, chickpea Cicer arietinum, and bitter vetch Vicia ervilia), and a single oil and fiber crop (flax Linum usitatissimum).

Table 1. 

Eight founder grain (three grass [cereal], four legume [pulse], and one oil and fiber) crops and their wild progenitors that started Neolithic agriculture in the Levant

Domesticated cropWild progenitor
Common nameScientific nameCommon nameScientific name
Einkorn wheatTriticum monococcum subsp. monococcumWild einkornT. monococcum subsp. baeoticum
Emmer wheatTriticum turgidum subsp. dicoccumWild emmerT. turgidum subsp. dicoccoides
BarleyHordeum vulgare subsp. distichumWild barleyH. vulgare subsp. spontaneum
LentilLens culinarisWild lentilLens orientalis
PeaPisum sativumWild peaPisum humile
ChickpeaCicer arietinumWild chickpeaCicer reticulatum
Bitter vetchVicia erviliaWild bitter vetchVicia ervilia
FlaxLinum usitatissimumWild flaxLinum bienne

From many aspects these eight plants belong to the same group—the grain crops. All of them are annuals, self-pollinated, diploid (except emmer wheat), native to the Fertile Crescent belt, and interfertile within each crop and between the crop and its wild progenitor. Both the agronomic compensation and the dietary complementation between plants in this group have been appreciated since the early days of agriculture up until now.

The aim of this article is to review the available information on these founder crops that started agriculture in the Levant during the Pre-Pottery Neolithic (PPN) period some 11,000–10,000 years ago (table 2). We shall review the current knowledge of these plants by focusing on two fields of study: (i) biology and (ii) archaeobotany. Biological data derived from the research on living plants can indicate what the wild progenitors of the domesticated plants may have been and what the selection pressures for domesticated types were. Archaeobotanical information identifies the plants used by hunting and gathering groups, which were the plants first domesticated, and when and where these processes took place.

Table 2. 

Cultural-historical sequence for the Southern Levantine Pre-Pottery Neolithic periods

Period and entity/phaseCalibrated 14C years BP
Late Epipaleolithic, Final Natufian12,500–11,700
PPNA11,600–10,500
PPNB: 
 Early10,500–10,100
 Middle10,100–9500
 Late9500–8750
 Final/PPNC8750–8400
Early Pottery Neolithic, Yarmukian8400–7600

Sources. Goring-Morris and Belfer-Cohen 2011; Kuijt and Goring-Morris 2002.

Note. PPNA = Pre-Pottery Neolithic A; PPNB = Pre-Pottery Neolithic B; PPNC = Pre-Pottery Neolithic C.

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Large amounts of critical new data, botanical and archaeological, have been gathered in the Fertile Crescent belt in the past 40 years (fig. 1). These findings established the Levant as the critical area for understanding early domestication of both plants and animals (and see additionally Zeder 2011). (Note that we use the terms “Southwest Asia” and “Near East arc” interchangeably in this article; we also use the term “Levant” to refer to the western horn of the Fertile Crescent in the Near East.)

Figure 1. 
Figure 1. 

Distribution of Epipaleolithic and Pre-Pottery Neolithic sites mentioned in the text.

In this article we will adopt a critical assessment intended to recognize domestication traits in Levantine archaeobotany (Zohary, Hopf, and Weiss 2011). In some way this assessment is partially intended to follow and update Mark Nesbitt’s (2002) article. The basics of this assessment are twofold.

1. The most reliable trait for the identification of domestication is ear shattering in the cereals, pod indehiscence in the pulses, and capsule indehiscence in flax. In wild plants, ears and pods/capsules disarticulate at maturation and shatter the seed-dispersal devices or the seeds; in contrast, ears and pods/capsules stay intact in the domestic plant. This trait is the best effective diagnostic indication for recognizing grain-crop domestication (i.e., the shift from wild plants to domesticated plants), which is controlled mostly by a single major gene locus or two such loci (table 3).

Table 3. 

Recessive mutations that changed wild-type trait into domestic-type trait in the Near East founder crops

CropWild-type traitDomestication traitNo. recessive mutationsSource
Einkorn wheatShattering earsNonshattering ears1Love and Craig 1924
Emmer wheatShattering earsNonshattering ears2Sharma and Waines 1980
BarleyShattering earsNonshattering ears2Takahashi 1955; Zohary 1960:41
LentilDehiscent podDehiscent pod1Ladizinsky 1979
PeaDehiscent podIndehiscent pod1Waines 1975
ChickpeaDehiscent podIndehiscent pod1Kazan et al. 1993
Bitter vetchDehiscent podIndehiscent podUnknown 
FlaxDehiscent capsuleIndehiscent capsule1Diederichsen and Hammer 1995; Gill and Yermanos 1967

Note. These mutations are responsible for the shift from wild-type seed dispersal to human-dependent crops. There is no conclusive information yet regarding the situation in bitter vetch.

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2. The second diagnostic morphological trait is seed size. In the wild progenitors the seeds are relatively small, while in domesticated plants they are frequently larger. This trait takes longer to develop, it is variable within the plant community, it is apparently a later development under domestication, and it is not as diagnostic as the first trait. This trait is controlled by various genes and other factors.

Additional markers for domesticated traits (e.g., Fuller 2007) are relevant mostly for differentiating between wild and domesticated traits in living plants. This is apparently much less so in the archaeobotanical assemblage.

During the past decade or so, molecular studies became central in research toward understanding the beginning of agriculture. Whether the mode of domestication was monophyletic or polyphyletic, these studies went in two lines (Brown et al. 2009). In the first half of this decade, molecular studies were regarded as supporting monophyletic origin of a single localized event or at least very rare events (e.g., Badr et al. 2000; Heun et al. 1997; Özkan et al. 2002). In the second half, however, such studies were interpreted as supporting polyphyletic domestication of multiple events in more than one location, such as of einkorn wheat (Kilian et al. 2007) and barley (e.g., Molina-Cano et al. 2005; Morrell and Clegg 2007). An attempt is made to evaluate the available knowledge, crop by crop, for the above-mentioned signs of the earliest definite domesticates as well as molecular and other biological data.

From the eight founder crops, this article does not deal with bitter vetch Vicia ervilia. According to its large quantities in Neolithic contexts, this pulse might have been taken into domestication in Anatolia or the Levant (i.e., in the general area in which it still grows wild today). However, because currently there are no reliable diagnostic traits to morphologically discriminate between its wild and domesticated forms in archaeological remains, the early Turkish finds could be either.

The source of most dates is Radiocarbon CONTEXT database (Böhner and Schyle 2002–2006). Table 4 lists the uncalibrated and calibrated radiocarbon dates. In an attempt to simplify understanding of the domestication process, all dates in the text are representative dates only and were rounded to the nearest 50 years (please refer to table 4 for range of dates). Periods mentioned in this article, such as Pre-Pottery Neolithic A (PPNA) or Early Pre-Pottery Neolithic B (EPPNB), do not necessarily imply cultural similarities but are used for the convenience of comparing among sites that belong to the same approximate time period.

Table 4. 

Representative radiocarbon dates

SiteLab no.14C dates BPCal BP (1σ)13CDated materialPeriod
Abu HureyraOxA-8818870 ± 10010,180–9790Bone/ovicaprine, burntPPNB
Abu HureyraBM-1724R8020 ± 1009093–8710CharcoalPPNB
Ali KoshB-1227218540 ± 909600–9440Organic material/carbonPPN
Ali KoshB-1082568000 ± 509000–8770Organic material/carbonPPN
ArpachiyahP-5858064 ± 789090–8770Charcoal/ashChalcolithic
ArpachiyahBM-15316930 ± 607830–7680CharcoalChalcolithic
BeidhaP-13809128 ± 10310,480–10,200CharcoalPPNB
BeidhaP-13798546 ± 1009630–9440Pistacia, nutsPPNB
Cafer HöyükLy-44369560 ± 19011,200–10,650CharcoalEPPNB
Cafer HöyükLy-44378950 ± 8010,220–9920CharcoalEPPNB
Can Hasan IAA-411707853 ± 368700–8580Charcoal/juniperEarly Chalcolithic
Can Hasan IP-7936254 ± 787270–7020CharcoalEarly Chalcolithic
Can Hasan IIIHU-118584 ± 659610–9490CharcoalAceramic Neolithic
Can Hasan IIIHU-97874 ± 708780–8580CharcoalAceramic Neolithic
CayönüGrN-62439320 ± 5510,590–10,420CharcoalEPPNB
CayönüGrN-62448980 ± 8010,240–9930CharcoalEPPNB
Choga MamiBM-4836846 ± 1827930–7560CharcoalNeolithic/Samarra
GilgalPta-45889920 ± 7011,470–11,230CharcoalPPNA
GilgalPta-45859710 ± 7011,230–11,080CharcoalPPNA
JerichoP-3828956 ± 10310,230–9910CharcoalPPNB
JerichoBM-13208539 ± 649550–9480CharcoalPPNB
Kissonerga-MylouthkiaOxA-74609315 ± 6010,650–10,410NDCypro-EPPNB
Kissonerga-MylouthkiaAA-331299110 ± 7010,380–10,200NDCypro-EPPNB
MureybitP-12249492 ± 12210,880–10,580CharcoalPPNA
MureybitLv-60710,590 ± 14012,800–12,390CharcoalPPNA
Netiv HagdudPta-45579780 ± 9011,320–11,090CharcoalPPNA
Netiv HagdudPta-45569660 ± 7011,200–11,060CharcoalPPNA
Ohalo IIRT-162420,840 ± 29025,500–24,600−20.60CharcoalMasraqan
Ohalo IIRT-129717,500 ± 20020,950–20,350−22.70CharcoalMasraqan
Parekklisha-ShillourokambosLy-2909310 ± 8010,660–10,400NDCypro-EPPNB
Parekklisha-ShillourokambosLy-9308670 ± 809740–9530NDCypro-EPPNB
RamadGrN-44268210 ± 509270–9030CharcoalPPNB
RamadGrN-48218090 ± 509130–8980CharcoalPPNB
Rizokarpaso-Cape-Andreas-KastrosMC-8057775 ± 1258720–8410NDCypro-PPNB
Rizokarpaso-Cape-Andreas-KastrosMC-8077450 ± 1208390–8160NDCypro-PPNB
Tepe SabzI-15017460 ± 1608420–8050CharcoalObeid
Tepe SabzI-15005410 ± 1606400–5990CharcoalObeid
YiftahelKN 35719100 ± 8010,390–10,190Horsebean seedsMPPNB
YiftahelRT 736A8570 ± 1309740–9430Horsebean seedsMPPNB

Note. PPN = Pre-Pottery Neolithic; PPNA = Pre-Pottery Neolithic A; PPNB = Pre-Pottery Neolithic B; EPPNB = Early Pre-Pottery Neolithic B; MPPNB = Middle Pre-Pottery Neolithic B; ND = unknown.

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Cereals

Einkorn Wheat Triticum monococcum

Biology

Einkorn is a small plant, rarely more than 70 cm high, with a relatively low yield, but it survives on poor soils where other wheat types usually fail. It is a relatively uniform diploid (2n=2x=14 chromosomes) wheat with characteristic hulled grains and delicate ears and spikelets. Most domesticated einkorn varieties produce one grain per spikelet, hence its name, but cultivars with two grains exist as well (Harlan 1981; Schiemann 1948).

The wild ancestry of domesticated einkorn wheat is well established. Domesticated Triticum monococcum is closely related to a group of wild and weedy wheat forms spread over Southwest Asia and adjacent territories and traditionally referred to as “wild einkorn” or Triticum boeoticum. The most distinguishing trait between wild einkorn and domesticated einkorn is the mode of seed dispersal. Wild forms have brittle ears, and the individual spikelets disarticulate at maturity to disperse the seed. In domesticated einkorn the mature ear remains intact and breaks into individual spikelets only on pressure (threshing). The shape of the grain is another diagnostic character indicating domestication (van Zeist 1976). In domesticated forms, the kernels tend to be wider compared with the wild forms.

Wild einkorn is widely distributed over Western Asia and penetrates also into the southern Balkans (Harlan and Zohary 1966; Zohary, Harlan, and Vardi 1969). Its distribution center lies in the Fertile Crescent (i.e., northern Syria, southern Turkey, northern Iraq, and adjacent Iran, as well as some parts of western Anatolia; fig. 2). In these regions, wild einkorn is massively distributed as a component of oak park-forests and steppelike formations.

Figure 2. 
Figure 2. 

Distribution of wild einkorn wheat Triticum monococcum subsp. baeoticum (=Triticum baeoticum). The area in which wild einkorn is massively spread is shaded. Dots represent additional sites outside the main area harboring mainly weedy forms (Zohary, Hopf, and Weiss 2011).

Heun and colleagues (Heun, Haldorsen, and Vollan 2008; Heun et al. 1997) analyzed domesticated and wild einkorn from the Fertile Crescent and beyond for amplified fragment length polymorphism (AFLP) DNA analysis. According to their research, wild einkorn wheat was domesticated in the Karacadağ range in southeast Turkey. It also supports the assumption of its monophyletic origin. These findings were recently supported by Kilian et al. (2007), who found, however, that several domesticate lines arose at the beginning of einkorn cultivation (i.e., that einkorn went through a number of independent domestication events). These lines are all part of the einkorn gene pool that grows wild in southeastern Turkey.

Archaeobotany

The earliest definite domesticated einkorn wheat appears in two EPPNB sites, namely ca. 10,600–9900 cal BP Çayönü (van Zeist and de Roller 1991–1992, 2003) and Cafer Höyük (de Moulins 1997), southern Turkey (fig. 1). In these sites, situated within the present range of distribution of wild einkorn (fig. 2), numerous charred spikelet forks were found showing rough breakage scars.

From these localities, einkorn spreads farther south to Middle Pre-Pottery Neolithic B (MPPNB), ca. 10,200–9550 cal BP, Tell Aswad near Damascus (van Zeist and Bakker-Heeres 1982 [1985]) and Jericho (Hopf 1983), where its remains, however, are relatively few. Generally, this wheat appears to be less frequent than the emmer wheat and barley.

Emmer and Durum-Type Wheats Triticum turgidum

Biology

Triticum turgidum is a varied aggregate of domesticated and wild tetraploid (2n=4x=28 chromosomes) wheats. According to their response to threshing, the domestic turgidum wheats fall into two groups of cultivars that are recognizable in archaeological remains.

1. Hulled nonshattering emmer wheat, Triticum turgidum subsp. dicoccum (traditionally called Triticum dicoccum), in which the products of threshing are the individual spikelets. The grains remain invested by the glumes and pales. In domestic emmer, as in einkorn, threshing results in breaking the rachis of the ear in its weakest points below each spikelet. Emmer represents the primitive situation in domesticated turgidum wheats.

2. The more advanced free-threshing emmer, which evolved under domestication from hulled emmer. This is a group of intraspecific taxa; its common representative today is durum wheat, T. turgidum conv. durum. The glumes in all these domesticated tetraploid types are relatively thin. Threshing breaks the glumes at their bases and frees the naked grains. The rachis is usually uniformly tough. Threshing breaks it into irregular fragments.

Hulled emmer (T. turgidum subsp. dicoccum) was the principal wheat of Old World agriculture in the Neolithic and early Bronze Age.

Genetic and morphological evidence clearly indicates (for review, see Feldman, Lupton, and Miller 1995; Zohary 1969) that the domesticated tetraploid turgidum wheats (both hulled dicoccum forms and free-threshing durum-type varieties) are closely related to wild wheat native to Southwest Asia and traditionally called Triticum dicoccoides (wild emmer wheat). This is annual, predominantly self-pollinated wheat with characteristic large and brittle ears and big elongated grains that show a striking similarity to some domesticated emmer and durum cultivars.

In the tetraploid turgidum wheats, the most conspicuous diagnostic difference between wild and tame is the seed-dispersal mechanism (Zohary 1969; Zohary and Brick 1961). Wild dicoccoides wheats have brittle ears that shatter on maturity into individual spikelets. Each spikelet operates as an arrowlike device disseminating the seed by inserting them into the ground. The “wild-type” rachis disarticulation and the spikelet morphology reflect specialization in seed dissemination that ensures the survival of the wild forms under wild conditions. In the human-made system of reaping, threshing, and sowing, this major adaptation broke down, and selection resulted in the evolution of human-dependant nonbrittle types. Significantly, the shift from a brittle spike (in wild dicoccoides wheats) to a nonbrittle spike (“hulled type” in domesticated dicoccum wheats) is governed by a single recessive mutation in a major gene (table 3). In addition, wild and domesticated forms differ from one another in kernel morphology (van Zeist 1976). In domesticated dicoccum and durum forms, the grain tends to be wider and thicker as well as rounder in cross section compared with the wild dicoccoides counterpart.

Wild emmer, T. turgidum subsp. dicoccoides, is more restricted in its distribution and more confined in its ecology than wild einkorn. Its range covers Israel, Jordan, southwestern Syria, Lebanon, southeastern Turkey, northern Iraq, and western Iran (fig. 3).

Figure 3. 
Figure 3. 

Distribution of the wild tetraploid emmer wheat Triticum turgidum subsp. dicoccoides (=Triticum dicoccoides) and Timopheev’s wheat Triticum timopheevi subsp. araraticum (=Triticum araraticum; Zohary, Hopf, and Weiss 2011).

At first, molecular studies placed a probable single origin of emmer wheat in Turkey. Özkan and colleagues (Özkan et al. 2002; Salamini et al. 2002) examined 204 AFLP loci of 99 wild emmer lines from all over the Fertile Crescent belt and several dozen different cultivars. They concluded that the most likely place of origin for emmer wheat is the Karacadağ Mountains in southeastern Turkey. There is, however, still a debate regarding the validity of these results and the number of possible domestication events as attested by recent molecular studies (e.g., Brown et al. 2009; Feldman and Kislev 2007). As Feldman and Kislev (2007) pointed out recently, the data of Mori et al. (2003), Özkan et al. (2002, 2005), and Luo et al. (2007) may indicate a diphyletic or polyphyletic origin of domesticated emmer, possibly in the Karacadağ area, as well as in the southern Levant.

Archaeobotany

Hulled emmer wheat. The earliest fully convincing sign to date of domesticated emmer comes from the numerous spikelet forks retrieved from EPPNB, ca. 10,600–9900 cal BP, Çayönü (van Zeist and de Roller 1991–1992, 2003). Here, hundreds of spikelet forks were discovered, all showing rough breakage scars characteristic of domestic emmer. Numerous spikelet forks, with similar telltale rough scars, are also available from contemporary contexts at Cafer Höyük (de Moulins 1997). These finds indicate that at EPPNB, emmer domestication must have been well under way in the Fertile Crescent. We wish to have more data before being fully convinced about the domestication status of the EPPNB Cypriot find from Kissonerga-Mylouthkia and Shillourokambos (Murray 2003; Willcox 2000).

To date, the earliest conspicuous grain-size increase is reported from EPPNB Tell Aswad, near Damascus, Syria (van Zeist and Bakker-Heeres 1982 [1985]). Here dicoccum-like plump kernels start to appear in the lowest habitation level, Ia (ca. 10,500–10,200 cal BP; Willcox 2005), and also in MPPNB phases II (ca. 10,200–9550 cal BP). In these phases, however, no rachis segments have been retrieved. Significantly, no wild dicoccoides-like narrow kernels were retrieved from Tell Aswad. As van Zeist and Bakker-Heeres emphasize, the continuous presence of morphologically discernible dicoccum kernels, the total absence (from the very start) of dicoccoides-like material, and the extreme dryness (less than 200 mm annual rainfall) suggest that emmer wheat was introduced into the Damascus Basin as a domesticated cereal not later than the second half of the eleventh millennium BP. From ca. 10,100–8700 cal BP onward, charred grains that morphologically conform with dicoccum appear also at Tell Abu Hureyra, northeast Syria, again with no rachis segments (Hillman 1975, 2000; Hillman, Colledge, and Harris 1989), and in contemporary PPNB Can Hasan III and Çatalhöyük East, Konya plain, Turkey (Fairbairn, Near, and Martinoli 2005; Fairbairn et al. 2002, 2007; Helbaek 1964, 1969; Hillman 1972, 1978), Ali Kosh, Deh Luran Plain, Khuzistan (Helbaek 1969), and Jericho, Israel (Hopf 1983). From the very beginnings of agriculture in Southwest Asia, emmer is one of the principal cereals, and it prevails quantitatively over domesticated barley and domesticated einkorn.

Free-threshing emmer wheats. Free-threshing wheat forms, identifiable by their rachis fragments, make their appearance in Southwest Asia soon after the firm establishment of emmer wheat cultivation (for review, see Maier 1996). They are already present among the plant remains of Late Pre-Pottery Neolithic B (LPPNB), ca. 9600–8600 cal BP, Can Hasan III, Turkey (Hillman 1972), and of MPPNB, ca. 10,200–9550 cal BP, Tell Aswad (van Zeist and Bakker-Heeres 1982 [1985]) and LPPNB, ca. 9450–8600 cal BP, Tell Bouqras (van Zeist and Waterbolk-van Rooijen 1985), Syria. They also occur in the MPPNB/LPPNB, ca. 9300–9000 cal BP, Çatalhöyük, Turkey (Fairbairn, Near, and Martinoli 2005; Fairbairn et al. 2002; Helbaek 1964), and Tell Ramad, Syria (van Zeist and Bakker-Heeres 1982 [1985]).

Barley Hordeum vulgare

Biology

Domesticated barley Hordeum vulgare subsp. vulgare is one of the main cereals of the belt of Mediterranean agriculture and a founder crop of Old World Neolithic food production. All over this area barley is a common companion of wheat, but in comparison with the latter, it is regarded as an inferior staple. Yet barley withstands drier and warmer environments, poorer soils, and some salinity. Because of these qualities, it has been the principal grain produced in numerous areas and an important element of the human diet. Barley is also the main cereal used for beer fermentation in the Old World. The crop was and still is a most important feed supplement for domestic animals.

Barley is a diploid (2n=2x=14 chromosomes) and predominantly self-pollinated crop. All cultivars have nonbrittle ears, a sharp contrast with wild barleys, in which ears are always brittle. Nonbrittleness in domesticated barley is governed by a recessive mutation in either one of two tightly linked “brittle” genes (table 3).

Barley ears have a unique structure. They contain triplets of spikelets arranged alternately on the rachis. According to the morphology of the spikelets, domestic barley can be divided into two principal types.

1. Two-rowed forms, traditionally called Hordeum distichum, in which only the median spikelet in each triplet is fertile and usually armed with a prominent awn. The two lateral spikelets are reduced and are grainless and awnless. Each ear thus contains only two rows of fertile spikelets.

2. Six-rowed forms, traditionally called Hordeum hexastichum, in which the three spikelets in each triplet bear seed and usually all are awned. Ears in these varieties therefore have six rows of fertile spikelets.

Wild barley is spread over the East Mediterranean Basin and the West Asiatic countries (fig. 4), penetrating as far as Turkmenia, Afghanistan, Ladakh, and Tibet. Wild barley occupies both primary and segetal human-made habitats. Its distribution center lies in the Fertile Crescent. In this area, wild barley is continuously and massively distributed. It constitutes an important annual component of open herbaceous formations, and it is particularly common in the summer-dry deciduous oak park-forest belt east, north, and west of the Syrian Desert and the Euphrates Basin and on the slopes facing the Jordan Rift Valley. From here, it spills over the drier steppes and semideserts.

Figure 4. 
Figure 4. 

Distribution of wild barley Hordeum vulgare subsp. spontaneum (=Hordeum spontaneum). The area in which wild barley is massively spread is shaded. Dots represent additional sites mainly of weedy forms. Wild barley extends eastward beyond the boundaries of this map as far as Tibet (Zohary, Hopf, and Weiss 2011).

The origins of barley are still not fully understood. Early crossing experiments and chloroplast DNA (cpDNA) typing have suggested that barley is of one, two, or at most a very few major domestication events (Zohary 1999). Later, Badr et al. (2000) examined 400 AFLP loci in wild and domesticated lines and found that barley is probably of a monophyletic origin in the Israel-Jordan area. However, in recent studies that included sequencing of seven genetic loci, Morell and Clegg (2007; as well as other molecular studies by Molina-Cano et al. [2005], [Saisho and Puruggana 2007], and Wang, Yu, and Ding [2009]) suggested two origins: one within the Fertile Crescent and a second farther east, possibly at the eastern edge of the Iranian Plateau. Apparently, European and North African barley is largely connected to the Fertile Crescent, while much of Asian barley is connected to the eastern center.

Archaeobotany

Unmistakable remains of nonbrittle barley (i.e., forms that could survive only under domestication) came from phase II (ca. 10,200–9550 cal BP) in Tell Aswad (van Zeist and Bakker-Heeres 1982 [1985]) and from ca. 9450 to 9300 cal BP Jarmo, Iraq. In the latter site (Braidwood 1960; Helbaek 1959a, 1960, 1966), Helbaek was the first to show two-rowed barley remains still closely resembling wild spontaneum but also displaying a nonbrittle rachis. Similar finds were reported and verified by Hopf (1983) in PPN Jericho. Indicative clues come from ca. 9600–8750 cal BP Ali Kosh (Helbaek 1969), where the brittle spontaneum-like material characterized the lower layers, and in the upper strata it was replaced by nonbrittle broad-seeded barley forms. Hulled barley has been found in Cyprus from the EPPNB, ca. 10,650–9550 cal BP (Murray 2003; Willcox 2000).

Domesticated barley continued to be a principal grain crop in Southwest Asia throughout the Neolithic period. Its remains have been recovered side by side with wheats in most Neolithic sites. Shortly afterward, we are faced with more advanced forms (i.e., six-rowed hulled as well as naked cultivars of barley).

In conclusion, the archaeological finds indicate that barley is a founder crop of the Levantine Neolithic agriculture and a close companion of emmer and einkorn wheats. The archaeological remains make it also possible to trace the main developments of barley under domestication: first the fixation of nonbrittle mutations and subsequently the emergence of six-rowed hulled and naked types.

Pulses

Lentil Lens culinaris

Biology

Lentil ranks among the oldest and the most-appreciated grain legumes of the Old World (Smartt 1990; Zohary 1995). In Mediterranean agriculture it is a characteristic companion of wheat and barley. Compared with the cereals, yields are relatively low, but lentil stands out as one of the most nutritious and tasty pulses. The protein content is about 25%, and lentil constitutes an important meat substitute in peasant communities.

The domesticated crop Lens culinaris (=syn. L. culinaris subsp. culinaris; Lens esculenta) manifests a wide range of morphological variation in both its vegetative parts and its reproductive parts. Like many other annual grain crops, lentil is predominantly self-pollinated, diploid (2n=2x=14), and interfertile, or largely so.

Conventionally, lentil cultivars are grouped in two intergrading clusters of seed sizes: (a) small-seeded lentils (subsp. microsperma), with small pods and small 3–6-mm seeds, and (b) large-seeded lentils (subsp. macrosperma), with larger pods and with seeds attaining 6–9 mm in diameter. Macrosperma forms are to be regarded as more advanced; they start to appear rather late in archaeological sequences, only in the third millennium BP. The occurrence of pods, or their fragments, is extremely rare.

As we described in the introduction to this article, the first sign of domestication is the retention of the seed in the pod (pod’s indehiscence) and the second is the gradual increase in seed size. The pod’s indehiscence is governed by a single mutation (table 3), the nondehiscent condition being recessive to the dehiscent one.

The wild progenitor of the domesticated plant, subsp. culinaris, shows close morphological, cytogenetic, and molecular affinities to wild subsp. orientalis, native to the East Mediterranean Basin and Southwest and Central Asia (fig. 5; Ferguson et al. 1998, 2000; Ladizinsky 1993; van Oss, Aron, and Ladizinsky 1997; Zohary and Hopf 1973). In fact, subsp. orientalis looks like a miniaturized subsp. culinaris but bears pods that burst open immediately after maturation.

Figure 5. 
Figure 5. 

Distribution of wild lentil Lens culinaris subsp. orientalis (=Lens orientalis; Zohary, Hopf, and Weiss 2011). Wild lentil extends eastward beyond the boundaries of this map into north Afghanistan and adjacent Central Asia.

Lens orientalis grows primarily on shallow stony soils and in gravelly hillsides in open or steppelike habitats. It also enters disturbed localities such as stony patches or stone heaps bordering orchards and cereal cultivation. In most parts of its distribution, L. orientalis is rather inconspicuous or even rare. It usually forms small scattered colonies. However, on stony slopes of Mount Hermon, the Anti-Lebanon, the oak park-forest belt of southern Turkey, and the western escarpments of the Zagros range, L. orientalis is occasionally locally common at 1,200–1,600 m altitude (Ladizinsky and Abbo 1993). Frequently it grows side by side with bitter vetch (Vicia ervilia).

As argued by Zohary (1999), the rich chromosomal polymorphism found in the wild progenitor compared with the chromosomal uniformity in the cultivars suggests that this pulse crop was taken into domestication only once or very few times.

Ladizinsky (1999) examined cpDNA restriction patterns. Using those results and additional information from crossability and chromosomal architecture led him to conclude that Lens originated from the territories around southern Turkey and north Syria. The same areas were also mentioned in later recombinant DNA studies conducted by Sonnante and coworkers (Sonnante, Galasso, and Pignone 2003; Sonnante, Hammer, and Pignone 2009).

Archaeobotany

Lentils seem to be closely associated with the start of wheat and barley domestication in the Levant. Very possibly this legume was introduced into domestication in this region together with emmer, einkorn, and barley; that is, it should be regarded as a founder crop of Old World Neolithic agriculture (Zohary and Hopf 1973).

At the end of the tenth and in the ninth millennia BP, charred seeds of lentil appear in most of the PPNB early farming villages in Southwest Asia. The seeds are still similar in size to those of wild forms (2.5–3.0 mm in diameter) and usually do not occur in quantities. Yet they are always associated with domesticated wheat and barley. Among the richest sites are ca. 10,200–9550 cal BP Tell Aswad (van Zeist and Bakker-Heeres 1982 [1985]), ca. 10,200–8700 cal BP Tell Abu Hureyra (Hillman 1975, 2000, 2001; Hillman, Colledge, and Harris 1989), ca. 10,250–9500 cal BP Jericho (Hopf 1983), ca. 10,600–9900 cal BP Çayönü (van Zeist 1972; van Zeist and de Roller 1991–1992, 2003), and ca. 9600–8800 cal BP Ali Kosh, Iran (Helbaek 1969).

A large hoard of carbonized lentils was recovered from the MPPNB, ca. 10,400–9450 cal BP, Yiftah’el, north Israel. The size of the hoard (ca. 1,400,000 seeds) and its contamination by the fruits of the weed Galium tricornutum, a characteristic weed in lentil cultivation, indicate that there and then lentil was already domesticated (Garfinkel, Kislev, and Zohary 1988). Large amounts of lentil seeds were discovered also in somewhat later phases of the Neolithic settlements in Southwest Asia: in ca. 9450–9300 cal BP Jarmo, Iraq (Braidwood 1960; Helbaek 1959a, 1960, 1966); in ca. 9250–9000 cal BP Tell Ramad, Syria (van Zeist and Bakker-Heeres 1982 [1985]); in ca. 8200–7800 cal BP ceramic Hacilar (Helbaek 1970); and in ca. 8350–7750 cal BP Tepe Sabz, Deh Luran Valley, Iran (Helbaek 1970). The Tepe Sabz lentils had already attained 4.2 mm in diameter. This is an obvious development under domestication. Lentils are repeatedly encountered in the early European and Southwest Asian Pottery Neolithic sites situated far outside the distribution area of L. orientalis, suggesting that these lentils were already domesticated.

In summary, archaeological remains do not provide us yet with direct diagnostic traits (such as indehiscent pod remains) for a determination of the start of lentil domestication. Moreover, it is doubtful whether comparative morphology will provide us with such clues in the future. Yet once Neolithic agriculture is soundly established, cultivation of lentil is part of it. The available archaeological information on early remains of lentil comes from the Levant, the very territory over which wild L. orientalis is distributed.

Pea Pisum sativum

Biology

The pea ranks among the oldest grain legumes of the Old World. From its early beginnings, this crop has been a close companion of wheat and barley domestication (Zohary and Hopf 1973). Pea Pisum sativum is well adapted to both warm Mediterranean-type and cool temperate conditions. In peasant communities in Southwest Asia, the Mediterranean Basin, temperate Europe, Ethiopia, and northwestern India, it constitutes an important source of protein for human consumption. The protein content of the seed is about 22%. Today, pea ranks among the world’s most important pulses (Davies 1995; Smartt 1990).

Pea is a diploid (2n=2x=14 chromosomes) and predominantly self-pollinated crop. As a consequence of the self-pollination system, variation in pea is molded in numerous true breeding lines. Domesticated pea shows a wide range of morphological variation. As we already described, the first sign of domestication is the retention of the seed in the pod (pod’s indehiscence), and the second is the gradual increase in seed size, from 3–4 to 6–8 mm in diameter. The pod’s indehiscence is governed by a single mutation (table 3), the nondehiscent condition being recessive to the dehiscent one. A third character here is the reduction of the relatively thick texture and rough surface of the seed coat, resulting in the breakdown of the germination inhibition of wild peas.

The crop complex of P. sativum contains the variable collection of pea cultivars and its closely related wild races. The wild forms of P. sativum fall into two main morphological types (fig. 6): (i) Pisum sativum subsp. elatius, a tall “maquis type” that thrives as a sporadic climber in maquis formations in the relatively mesic parts of the Mediterranean region and as a weed, and (ii) Pisum sativum var. pumilio (known formally as Pisum humile), a shorter, more xeric “steppe type” geographically restricted to southwest Asia. It occurs in the deciduous oak park-forest belt and in open steppelike herbaceous vegetation formations characteristic of the Fertile Crescent (i.e., in the same zone that harbors the wild progenitors of wheat, barley, lentil, and flax). From such primary habitats humile peas spill over to secondary habitats and occasionally infest cereal cultivation.

Figure 6. 
Figure 6. 

Distribution of the two main wild races of pea Pisum sativum: (i) “steppe-type” humile forms and (ii) “maquis-type” elatius forms (Zohary, Hopf, and Weiss 2011). In the West Mediterranean Basin, wild elatius peas extend beyond the boundaries of the map and reach as far as Spain.

The available evidence from the living plants implicates the wild humile peas as the progenitor stock for pea domestication. Humile peas show closer morphological similarities than elatius peas to the domesticated aggregate and grow in steppelike habitats (i.e., under open conditions similar to the cultivated field). Within humile peas, the Turkish and the south Levant forms having chromosomes identical to those present in the cultivars should be regarded as the primary ancestral stock. This is also supported by cpDNA comparisons. Yet it is very likely that the two subspecies contributed some genes to the cultivated ensemble through occasional secondary hybridization (Ben Ze’ev and Zohary 1973; Palmer, Jorgensen, and Thompson 1985).

Nasiri and coworkers (Nasiri, Haghnazari, and Saba 2009) have examined SSR (microsatellite) markers of wild peas and cultivars. They have found that these markers effectively differentiate between the cultivars and the wild accessions. In addition, Pisum sativum subsp. fulvum was found to be the closest relative of the cultivars, but it should be noted that subsp. humile was not tested in this study.

Archaeobotany

Remains of peas are present in many of the PPNB farming villages that developed in the Fertile Crescent arc from 10,500 cal BP years onward. Some of the earliest finds were retrieved from ca. 10,200–9550 cal BP Tell Aswad in south Syria (van Zeist and Bakker-Heeres 1982 [1985]), ca. 10,600–9900 cal BP Çayönü in southeast Turkey (van Zeist and de Roller 1991–1992, 2003), ca. 10,250–9500 cal BP PPNB Jericho (Hopf 1983), and ‘Ain Ghazal, Jordan (Rollefson et al. 1985). Much richer remains were available from somewhat later Neolithic phases in the Levant. Large quantities of carbonized seed accompany the domesticated wheats and barley in MPPNB/LPPNB Çatalhöyük (Fairbairn, Near, and Martinoli 2005; Fairbairn et al. 2002; Helbaek 1964) and Final PPNB/PPNC Erbaba (van Zeist and Buitenhuis 1983).

In contrast to those of the wheats and barley, the earliest archaeological remains of pea do not provide us with simple traits for a foolproof recognition of domestication (Zohary and Hopf 1973). In peas under domestication there is a general trend toward an increase in the size of the seed and the length of the hilum, but such changes occurred gradually in the course of domestication. In early finds there is a considerable overlapping in the dimensions of wild and domesticated forms. Perhaps the most reliable indication of domestication in peas is provided by the surface of the seed coat. Wild peas are characterized by a rough or granular surface, while domesticated varieties have smooth seed coats. However, seed coats survive only very rarely, and if they do not, it is impossible to ascertain whether the material retrieved represents wild or domesticated forms. The lower levels (ca. 10,600–9900 cal BP) of Çayönü (van Zeist and de Roller 1991–1992) retained some fragments of rough-surface wild seed coats. Wild-type seed coats occur even much later in Pottery Neolithic Hacilar (Helbaek 1970). Significantly, the remains from MPPNB/LPPNB Çatalhöyük (Helbaek 1964) and Bouqras (van Zeist and Waterbolk-van Rooijen 1985) and those from LPPNB Çayönü and Can Hasan I (Renfrew 1968) include a single seed with smooth seed coat characteristic of domesticated varieties. This smoothness suggests that domestication of peas in the Levant is as old, or almost as old, as the domestication of wheat and barley.

Although it is currently less definite, the archaeological evidence establishes pea as one of the founder crops of the Levantine Neolithic agriculture. Since this early start, pea seems to be a consistent and common element of food production and a common companion of wheats and barley. The evidence from the living plants complements the archaeological finds. The wild humile forms, with chromosomes and cpDNA identical to those prevailing in the cultivated crop, should be regarded as the closest wild relatives from which this pulse crop evolved.

Chickpea Cicer arietinum

Biology

Chickpea is a valued grain legume of the traditional agriculture in the Mediterranean Basin and Western Asia as well as India and Ethiopia. It is a member of the grain ensemble found in Levantine Neolithic and Bronze Age remains. Chickpea is adapted to a subtropical or Mediterranean-type climate; it grows almost exclusively in the postrainy season on moisture stored in the soil. Like lentil and pea, chickpea (with a seed protein content of some 20%) constitutes an important meat substitute in peasant communities.

Domesticated chickpea Cicer arietinum, like all other eight founder crops, is a predominantly self-pollinated annual crop with pods containing one to two seeds. The cultivars show a wide range of variation in size, color, and shape of the seed and in the size and form of leaves and flowers. All domesticated varieties are diploid (2n=2x=16 chromosomes) and interfertile. Chickpea landraces are grouped into two interconnected clusters (Smartt 1990; Smithson, Thompson, and Summerfield 1985). Large-seeded varieties (known as “Kabuli” type) with relatively smooth, rounded, light-colored seed coats and pale cream flowers predominate in the Mediterranean countries and Southwest Asia. Varieties producing small wrinkled seeds (“Desi” type) with dark-colored seed coats and usually purple flowers prevail in the eastern and southern parts of the distribution area of the crop (i.e., in India, Afghanistan, and Ethiopia).

As in most other seed legumes, the conspicuous features of evolution under domestication in chickpea are the retention of seeds in the pods (pod’s indehiscence) and the gradual increase in seed size, from 3.5 to 6.0 mm and more. The pod’s indehiscence is governed by a single mutation (table 3). Another change under domestication is the development of a smooth seed coat and the reduction of its thickness. Seed remains are the only material recovered to date in archaeological excavations.

The domesticated chickpea C. arietinum is a member of a leguminous genus comprising some 40 species centered in Central and Western Asia (Cole, Maxted, and van der Maesen 1998; van der Maesen 1972). The domesticated pulse shows close morphological affinities and an almost identical seed protein profile to two wild species of chickpea: Cicer echinospermum and Cicer reticulatum. These two wild chickpeas are diploid self-pollinated annuals known only from southeastern Turkey. The available morphological and cytogenetic evidence implicates C. reticulatum as the wild ancestor of the domesticated plant (Ladizinsky and Adler 1976), and it is therefore referred to as C. arietinum subsp. reticulatum. Because the distribution of the wild progenitor is restricted to southeast Turkey (fig. 7), the area of origin of the domesticated crop can be outlined there.

Figure 7. 
Figure 7. 

Distribution of wild chickpea Cicer arietinum subsp. reticulatum (=Cicer reticulatum; Zohary, Hopf, and Weiss 2011).

Archaeobotany

Like lentil and pea, chickpea seems to be closely associated with the start of food production in the Levant, but unlike them it is much rarer in Neolithic contexts. Earlier chickpea seeds were found in EPPNB, ca. 10,600–10,250 cal BP, Tell el-Kerkh, northwest Syria (Tanno and Willcox 2006). A few of these seeds are somewhat larger and plumper than the wild progenitor, and the authors argue (Tanno and Willcox 2006:200) that this might indicate that they are an “intermediate stage” between wild and domesticated chickpeas. Some charred chickpea seeds were recovered from EPPNB, ca. 10,600–9900 cal BP, Çayönü (van Zeist 1972; van Zeist and de Roller 1991–1992). A few more were found in the MPPNB level of Tell Abu Hureyra, northern Syria (Hillman 1975, 2000), and Aşıklı Höyük (van Zeist and de Roller 1995) in Turkey. The seeds from these sites correspond in size to those of C. reticulatum. Because these sites are situated within (or close to) the very restricted geographic distribution area of the wild progenitor, it is difficult to be sure whether they represent wild or domestic plants. The seeds retrieved from MPPNB Jericho (Hopf 1983) and ‘Ain Ghazal (Rollefson et al. 1985) and those from LPPNB Ramad near Damascus (van Zeist and Bakker-Heeres 1982 [1985]) very probably represent domesticated forms. The latter sites lie far away from the territory of the wild progenitor, and the specimens from Jericho seem to have smooth seed coats.

The evidence from the living plants and the plant remains discovered in archaeological excavations indicate that C. arietinum belongs to the early Neolithic grain-crop assemblage of the Levant. Archaeological data are still limited and less definite, but in this legume, the delimitation of the place of origin is relatively simple: the wild progenitor of the domesticated chickpea is endemic to the central part of Fertile Crescent. Here, very likely, this pulse was first brought into domestication.

Fiber and Oil Plants

Flax Linum usitatissimum

Biology

Flax Linum usitatissimum is an annual crop with characteristic slender strong stems and rounded capsules that in domesticated forms do not dehisce but retain the oval compressed shining seed. The crop is diploid (2n=2x=30 chromosomes) and predominantly self-pollinated. Consequently, variation has been molded in the form of numerous true breeding lines and aggregates of landraces. Two specializations are apparent: (i) oil varieties, which are relatively short (30–70 cm) and branched and usually bear large seeds, and (ii) fiber varieties, which are taller and sparsely branched and usually produce small seeds. Transitional forms, cultivated for both oil and fiber, occur as well.

Flax was a principal oil and fiber source in the Old World and probably the earliest domesticated plant used for textiles. Until recently, flax was extensively cultivated in vast areas of Eurasia (Durrant 1976). In antiquity, flax fibers were the principal vegetable fiber used for weaving textiles in Europe and Western Asia. The seed contains about 40% oil, and in peasant communities linseed was used as a source for edible oil and high-grade lighting oil.

The fibers for spinning are obtained from the tall stems, which are harvested before the maturation of the seed. Traditionally, they were first dried and then immersed (wetted) in water to allow the microbial decomposition (retting) of the pectin connecting the fibers with other cells and tissues of the stem. After retting, the stems were dried, and the fibers (averaging 4 cm in length) were separated by pounding (breaking) and combing.

Domesticated flax L. usitatissimum is most closely related to wild Linum bienne (syn. Linum angustifolium). These two flaxes have the same chromosome number (see above), intercross readily, and are fully interfertile (Gill and Yermanos 1967). Linum bienne—with its characteristic strong branches, blue flowers, and dehiscent capsules—is widely distributed over West Europe, the Mediterranean Basin, North Africa, Southwest Asia, Iran, and Caucasia (fig. 8). Some wild forms are annual, and others are biennial or perennial; all are predominantly self-pollinated. Linum bienne grows mainly in wet places such as moist grassy areas, springs, seepage areas on rocky slopes, moist clay soils, and marshy lands. On the basis of its close morphological and cytogenetic affinities to the domesticated crop, L. bienne is identified as the wild progenitor of L. usitatissimum (Diederichsen and Hammer 1995). The main changes under domestication are the shift to nonsplitting capsules and the increase of seed size (like in many grain crops) as well as the selection for higher oil yield or longer stems with a high amount of long fibers.

Figure 8. 
Figure 8. 

Distribution of wild flax Linum usitatissimum subsp. bienne (=Linum bienne; Zohary, Hopf, and Weiss 2011).

Phylogenetic evidence from modern accessions (Allaby et al. 2005; Fu and Allaby 2010) gives indications that (i) there was a single domestication event for flax and (ii) the oil-producing variety was domesticated, suggesting that domestication selected for the larger oil-rich seeds rather than its fibers.

Archaeobotany

Flax was apparently used by humans already before its domestication. Recently, twisted and dyed flax fibers were reported in Upper Paleolithic Dzudzuana Cave, Georgia (ca. 30,000 years old; Bergfjord et al. 2010; Kvavadze et al. 2009, 2010). The oldest wild linseed remains retrieved from archaeological sites in Southwest Asia come from ca. 10,900–9900 cal BP Tell Mureybit (van Zeist and Casparie 1968). Soon after, seeds of flax were found in many of the PPNB farming villages that appeared in the Fertile Crescent from 10,500 cal BP onward (fig. 1). Some of the earlier finds come from ca. 10,600–9900 cal BP Çayönü, Turkey (van Zeist 1972; van Zeist and de Roller 1991–1992, 2003); ca. 10,200–9550 cal BP Tell Aswad, near Damascus, Syria (van Zeist and Bakker-Heeres 1982 [1985]); Ali Kosh, Iran (Helbaek 1969); Jericho, Israel (Hopf 1983); and ‘Ain Ghazal, Jordan (Rollefson et al. 1985). The seeds are still small, similar in size to those of wild bienne forms, yet they are almost always associated with domesticated wheats and barley.

Fragments of a capsule from ca. 10,250–9500 cal BP MPPNB Jericho (Hopf 1983) is probably the earliest indication we have today of domesticated flax. Another indication of early flax domestication comes from linseed remains recovered from LPPNB, ca. 9250–9000 cal BP, levels of Tell Ramad, Syria. The calculated size of the seed from this site, corrected for charring shrinkage, ranges from 3.2 to 4.1 mm in length. This is already within the size class of the L. usitatissimum seed, the lower limit of which lies at 3.0 mm. It is therefore an attractive indication for flax domestication under rain-dependent conditions before ca. 8600 cal BP (van Zeist and Bakker-Heeres 1975). Such domesticated linseeds were found in Pottery Neolithic Nahal Zehora, Mount Carmel, Israel, ca. 8150–7850 cal BP (Kislev and Hartmann, forthcoming). In the Mesopotamian Basin, linseed sizes in ca. 8400–6000 cal BP Tell Sabz, Iran (Helbaek 1969), and in Halafian Arpachiya (Helbaek 1959b) are even bigger (4.7–4.8 mm long). As argued by Helbaek, these large seeds indicate advanced domestication and demonstrate that flax was part of the irrigated grain agriculture system that evolved in this region. In Choga Mami, Iraq, Helbaek (1972) reports a rare find in the earlier, ca. 7950–7550 cal BP, stratum, which became frequent during the following one, dated to the second half of the eighth millennium BP. Linseed also appears in several later Southwest Asian Neolithic and Bronze Age sites.

Remains of flax textiles also make an early appearance. The best examples come from the drier parts of Southwest Asia, where because of low humidity, woven material survived without carbonization. Pieces of exquisitely woven linen were discovered among PPNB (beginning of the ninth millennium BP) remains in Nahal Hemar cave near the south tip of the Dead Sea, Israel (Schick 1988). A single piece of linen was found in Neolithic (eighth millennium BP) Fayum, Egypt (Caton-Thompson and Gardner 1934).

The available archaeological evidence clearly suggests that flax belongs to the first group of grain crops that started agriculture in the Levant during the MPPNB. The gradual increase in seed size and the use of linen indicate that flax domestication was very probably already practiced in the PPNB Levant as attested first by MPPNB capsules and then by LPPNB seeds and linen textile. The evidence from the living plants shows that wild bienne forms are widespread in the arc and fully supports a Fertile Crescent domestication.

Conclusions

This article deals with current biological and archaeobotanical information from Southwest Asia (fig. 1) in an attempt to identify the earliest finds of domesticated grain crops. If we adopt the criteria suggested in the beginning of this article, we can now separate the appearance of the eight founder crops into several distinguishable groups and dates.

From the group of eight Southwest Asian founder crops, two crops seem to appear earlier as cultivated wild plants. These “pioneer crops” are (i) wild barley, found in PPNA Gilgal, and (ii) wild lentil, found in PPNA Jerf el-Ahmar and in PPNA Netiv Hagdud (Weiss, Kislev, and Hartmann 2006). Some lines of evidence suggest that cultivation of wild einkorn, wild emmer, wild barley, wild rye, and wild lentil might have been practiced at PPNA Jerf el Ahmar and Dja’de, northern Syria (Willcox, Fornite, and Herveux 2008).

From the group of founder crops, the earliest definite domesticated plants are einkorn wheat and emmer wheat from two EPPNB sites in Turkey, Cafer Höyük (de Moulins 1997) and Çayönü (van Zeist 1972; van Zeist and de Roller 1991–1992, 2003). Unfortunately, study of the third possible site in this cluster, Nevali Çori (Pasternak 1998), was published only as a preliminary publication and therefore remains uncertain.

The earliest definite domestic forms of barley and lentil first appear during the MPPNB. In this period, domesticated barley is present in Aswad (van Zeist and Bakker-Heeres 1982 [1985]) and later on (still in the MPPNB) in Jarmo (Helbaek 1969). Domesticated lentil appears in MPPNB Yiftah’el (Garfinkel, Kislev, and Zohary 1988).

The earliest domesticated flax was retrieved from MPPNB Jericho (Hopf 1983) and later in LPPNB Tell Ramad (van Zeist and Bakker-Heeres 1982 [1985]) and Nahal Hemar, Israel (Schick 1988).

The evidence regarding the remaining three founder crops—chickpea, pea, and bitter vetch—is as yet inconclusive. The geographical boundaries of the first group of domesticated plants therefore focus on a rather restricted part of the Fertile Crescent (i.e., from southeastern Turkey in the north to Israel in the south).

We see now (and see Zeder 2011 for details and references) the first wave of animal domestication (all herbivores: goat, sheep, cow, and pig) more or less at the same time as early plant domestication. It is possible, therefore, that the remains of food processing for human food—stalks and inflorescences—were used for animal feed of the emerging Neolithic culture.

At present, no agreement prevails among researchers and between research disciplines regarding the mode of origin of the “first wave” of the Southwest Asian grain crops (einkorn wheat, emmer wheat, barley, lentil, pea, chickpea, bitter vetch, and flax). Was it of monophyletic origin (derived from the same ancestral taxon representing a single event of domestication) or of polyphyletic derivation (resulting from several ancestral taxa representing several independent events)?

The accumulated morphological, anatomical, and cytogenetic information seems to support monophyletic speciation as the plausible mode of origin (Zohary 1996, 1999). Important in domestication is the interaction between annaulity, reproductive biology, and self-pollination and selection; self-pollination is significant because it causes the plant community to be built of almost only pure inbred lines. The combination of selfing and selection of inbred lines further supports this notion because it causes the selection toward domestic types to be efficient and fast.

As far as the molecular research goes, however, there are currently two views based on analytical as well as methodological perspectives. Some (e.g., Badr et al. 2000; Heun et al. 1997; Özkan et al. 2002) support a monophyletic origin, while others (e.g., Kilian et al. 2007; Molina-Cano et al. 2005; Morrell and Clegg 2007) support a polyphyletic one. It seems that this major research question will get more attention in the near future. No doubt the research avenue of molecular analysis of domesticated crops and their origin is a fast-growing field that has not yet reached its climax.

Although it is beyond the scope of this article, it is important to mention the relatively new data available from Cyprus (e.g., Peltenburg et al. 2000, 2001; Vigne 2011; Vigne et al. 2000; Willcox 2000). It is apparent that already during the EPPNB, human pioneers crossed the sea and settled the island, bringing with them the Neolithic culture. These finds hint that the beginning of agriculture on the mainland, as attested in this article, was transferred almost immediately (on an archaeological timescale) outside the core area. Cyprus, as an isolated phenomenon, teaches us the pace of the biological and cultural shifts from wild to domesticated plants and from hunter-gatherers to farmers.

This article is partially based on materials from D. Zohary, M. Hopf, and E. Weiss, Domestication of Plants in the Old World, 4th edition, forthcoming from Oxford University Press.

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Notes

Ehud Weiss is Senior Lecturer at the Institute of Archaeology, Martin (Szusz) Department of Land of Israel Studies and Archaeology, Bar-Ilan University (Ramat-Gan, 52900 Israel []) and Visiting Scientist, Kimmel Center for Archaeological Sciences, Weizmann Institute of Science (Rehovot, 76100 Israel). Daniel Zohary is Professor in the Department of Evolution, Systematics, and Ecology, Hebrew University Jerusalem (91904 Israel).