Vegetation History and Archaeobotany (2019) 28:449–463 https://doi.org/10.1007/s00334-018-0702-y ORIGINAL ARTICLE Re-analysis of archaeobotanical remains from pre- and early agricultural sites provides no evidence for a narrowing of the wild plant food spectrum during the origins of agriculture in southwest Asia Michael Wallace1 · Glynis Jones1 · Michael Charles2 · Emily Forster2 · Eleanor Stillman3 · Vincent Bonhomme3,4 · Alexandra Livarda5 · Colin P. Osborne6 · Mark Rees6 · Georg Frenck6,7 · Catherine Preece6,8 Received: 26 February 2018 / Accepted: 22 October 2018 / Published online: 17 November 2018 © The Author(s) 2018 Abstract Archaeobotanical evidence from southwest Asia is often interpreted as showing that the spectrum of wild plant foods narrowed during the origins of agriculture, but it has long been acknowledged that the recognition of wild plants as foods is problematic. Here, we systematically combine compositional and contextual evidence to recognise the wild plants for which there is strong evidence of their deliberate collection as food at pre-agricultural and early agricultural sites across southwest Asia. Through sample-by-sample analysis of archaeobotanical remains, a robust link is established between the archaeological evidence and its interpretation in terms of food use, which permits a re-evaluation of the evidence for the exploitation of a broad spectrum of wild plant foods at pre-agricultural sites, and the extent to which this changed during the development of early agriculture. Our results show that relatively few of the wild taxa found at pre- and early agricultural sites can be confidently recognised as contributing to the human diet, and we found no evidence for a narrowing of the plant food spectrum during the adoption of agriculture. This has implications for how we understand the processes leading to the domestication of crops, and points towards a mutualistic relationship between people and plants as a driving force during the development of agriculture. Keywords Archaeobotany · Neolithic · Pre-Pottery Neolithic · Wild plant foods · Wild plants · Broad spectrum Communicated by A. Fairbairn. Electronic supplementary material The online version of this article (https​://doi.org/10.1007/s0033​4-018-0702-y) contains supplementary material, which is available to authorized users. 5 * Michael Wallace Department of Classics and Archaeology, University m.p.wallace@sheffield.ac.uk of Nottingham, University Park, Nottingham NG7 2RD, UK 6 1 Department of Animal and Plant Sciences, University Department of Archaeology, University of Sheffield, of Sheffield, Alfred Denny Building, Western Bank, Minalloy House, 10‑16 Regent Street, Sheffield S1 3NJ, UK Sheffield S10 2TN, UK 2 Department of Archaeology, University of Oxford, 36 7 Department of Ecology, University of Innsbruck, Beaumont Street, Oxford OX1 2PG, UK Technikerstraße 25, 6020 Innsbruck, Austria 3 School of Mathematics and Statistics, University of Sheffield, 8 CREAF, Campus de Bellaterra (UAB), Edifici C, Hicks Building, Hounsfield Road, Sheffield S3 7RH, UK 08193 Cerdanyola del Vallès, Spain 4 Institut des Sciences de l’Evolution‑Montpellier (ISEM‑UMR 5554), Equipe Dynamique de la Biodiversité, Anthropo‑écologie, Université de Montpellier, 34095 Montpellier Cedex 2, France 13 Vol.:(0123456789) 450 Vegetation History and Archaeobotany (2019) 28:449–463 Introduction approach to the collection of wild plant foods. Though contradictory, the arguments of both Weiss et al. (2004) The collection of wild plant foods and Savard et al. (2006) are based on the assumption that the plant remains present on early sites were primarily The emergence of agriculture in the Pre-Pottery Neolithic collected as food. (PPN) of southwest Asia marked a major change in human However, it has long been acknowledged that the recog- subsistence strategies, with a shift from communities nition of wild plants as foods is problematic (e.g. Dennell based on the gathering of wild plants to those reliant pri- 1976; van Zeist and Bakker-Heeres 1985; Willcox et al. marily on the cultivation of eight founder crops (einkorn, 2008). First, there are other means by which seeds from emmer, barley, lentil, pea, chickpea, bitter vetch and flax). wild plants may have inadvertently arrived on archaeologi- It is widely accepted that a broad spectrum of plant foods cal sites, including their unintentional collection together was exploited by late Palaeolithic hunter-gatherers and that with wild food plants (e.g. Savard et al. 2003, p 102) or with there was a narrowing of this spectrum with the advent of woody material (Arranz-Otaegui et al. 2018), their arrival domesticated crops and the emergence of agriculture (e.g. on site as weeds of crops (e.g. Hillman 1984, 2000), or their Weiss et al. 2004; Savard et al. 2006; Willcox et al. 2008; inclusion in animal dung (e.g. Miller and Smart 1984; Miller Colledge and Conolly 2010). 1984, 1996; Spengler 2018). Secondly, even when it can The rise and fall of a broad spectrum of food resources be established that a plant was deliberately brought onto has been linked to notions of optimal foraging theory site, there are numerous reasons other than its use as food whereby, in the face of resource depletion due to popu- why it may have been collected, e.g. as building material, lation increase and/or environmental change, foragers bedding or fuel. The aims of this paper are therefore (1) to increasingly exploited lower ranked resources which sub- distinguish the suite of wild plants for which there is strong sequently declined in importance with the advent of higher evidence that they were (a) deliberately collected and (b) ranking resources such as domesticated plants and animals were intended to be used as food by pre-agricultural or early (Flannery 1969; Stiner et al. 2000; Stiner 2001; Stiner and agricultural communities, and (2) to re-evaluate the evidence Munro 2002; Winterhalder and Kennett 2006; Gremillion for the exploitation of a broad spectrum of wild plant foods, et al. 2014). Following this view, Weiss et al. (2004) have and the extent to which this changed during the development argued that a broad range of small-seeded grasses con- of early agriculture. tributed to the forager diet in the Upper Palaeolithic but that their contribution declined in comparison with that Contextual and compositional evidence of wild cereals until the PPNA (early PPN), after which their significance was negligible. They see the flourishing Two criteria have been advocated by archaeobotanists as of a broad range of plant foods in the Upper Palaeolithic a means of distinguishing the deliberate collection of wild as indicating a temporary switch to low-ranked foods due plants from their unintentional arrival on site: archaeologi- to pressure on food resources. cal context, such as in a storage pit or vessel (van Zeist There are several reasons, however, why the exploita- and Bakker-Heeres 1985, p 247) and high concentrations tion of a broad spectrum of foods might persist, including of seeds of the same type (e.g. Willcox et al. 2008, p 317). buffering against the risk of food shortages or a reluctance These criteria have been used in conjunction as in the to abandon culturally preferred wild foods. An alterna- case of concentrations of seeds in plaster bins at Çatal- tive view is that, rather than reflecting an increased use höyük (Bogaard et al. 2009; Twiss et al. 2009) and Hacılar of lower ranked taxa, the exploitation of a broad spec- (Helbæk 1970), in proximity to food processing areas at trum of foods was an opportunistic response to plentiful Ohalo II (Weiss et al. 2008) and Jerf el Ahmar (Willcox and diverse resource availability (Smith 2007, 2011a, b), 2002), and beneath a granary at Gilgal (Weiss et al. 2006). which facilitated more permanent settlement, and included Arranz-Otaegui et al. (2016) have recently commented, managed or domesticated plants alongside collected wild however, on the overall lack of detailed analyses of sample foods (Zeder 2012, 2015, 2016). Savard et al. (2006) pro- composition of PPN archaeobotanical assemblages and the vide some support for this view. Using a different quanti- poor recording of contextual associations between plant fication method to that used by Weiss et al. (2004), they remains and archaeological features. Here we address suggest that the use of small-seeded grasses did not fall these issues by considering both the botanical composition sharply after the Epipalaeolithic period in the eastern Fer- of individual samples and, where known, their archaeo- tile Crescent, but rather that these grasses or other non- logical context, in order to achieve our first aim of estab- grass species continued to form a significant component of lishing a robust link between the archaeological evidence the diet throughout the PPNA, suggesting an opportunistic and its interpretation in terms of deliberate collection and use. Approaches based on the overall frequency/ubiquity 13 Vegetation History and Archaeobotany (2019) 28:449–463 451 or abundance of wild species in a site (or period) assem- Methodology blage are unable to achieve this, and can merely indicate the availability of wild species. We therefore consider the Archaeobotanical database sample-level approach adopted here as a necessary precur- sor to our second aim, an investigation of the spectrum of We systematically reviewed the published and, where pos- wild plant foods at PPN sites and changes in wild food use sible, unpublished archaeobotanical data for all PPN and through time. earlier sites (primarily 12000–5000 cal bc) in southwest Contexts associated with storage or processing are Asia (central Anatolia, Cyprus, the southern and northern most likely to indicate the deliberate collection and use Levant, and the eastern Fertile Crescent) at which plant of plants. Storage contexts may include ceramic vessels, remains have been found (75 sites, Fig. 1). Archaeobot- clay bins, pits and receptacles composed of more perish- anical records of charred plant remains were entered into able materials, such as reed baskets (Hillman et al. 1989; a database as individual samples in all cases for which Özdoğan 2009, p 23). Pots and bins are good storage indi- sample-level data were available (3,162 samples from 52 cators but are rarely found in the PPN, and even these sites; see ESM Table A which also lists the sources of data may have received secondary deposits of plant material and ESM Text A which provides details of an online ver- after they had gone out of use as storage devices. Pits are sion of the database). This provides the finest resolution ambiguous indicators of storage, as they were often used possible for published records and site archives, and there- for other purposes such as disposal of refuse. Storage in fore the closest approximation of individual “behavioural perishable containers is usually only identifiable from con- episodes” (Jones 1991). centrations of plant material (usually in burnt destruction To prepare the database for analysis, it was necessary to deposits), and such concentrations are particularly indica- standardise both the plant nomenclature and the method of tive of storage if found in association with (e.g. in the quantification. Taxonomic synonyms were therefore com- same room as) less perishable storage containers, as at Jerf bined to give aggregated counts under a single taxonomic el Ahmar (Willcox 2002) and Çatalhöyük (Bogaard et al. name. For crops, the nomenclature proposed by Miller 2009; Twiss et al. 2009). Processing of plants for use may (1992) was adopted. This standardisation reduced the be indicated by association with processing facilities or by original 844 taxonomic entries to 698 unique taxa. The proximity to hearths or ovens in which they may have been most common method of quantification used in the source accidentally charred. For all contexts, however, re-use and literature was a count of the number of plant items. Where the introduction of secondary plant material can only be other quantification methods were used, such as weight, excluded through analysis of botanical composition. scales of abundance or presence, values were converted Two compositional characteristics of an archaeobotani- to counts as far as possible. Some authors provide conver- cal sample that have been used to indicate storage are den- sion factors for converting weight or abundance scales into sity and purity (Dennell 1976; Jones 1991). The greater the counts, but in the case of simple presence a count of one density of plant remains [number of plant items − usually was attributed to each reported taxon. Samples for which diaspores (hereafter referred to as seeds) per unit volume of the data were originally recorded as counts, or where data sediment], the more likely it is that they were deposited as could be reliably converted to counts, constitute 92% of a single entity. A sample of high purity (e.g. dominated by the total samples. one species) is likely to be the result of a depositional event relating to the dominant species within the sample. The combination of high density and high purity is thus a good Contextual evidence indication of storage of the dominant species, and concen- trations of many hundreds of seeds of the same species are The quality of contextual information in published reports often considered unambiguous indicators of storage. Even is highly variable. Detailed contextual descriptions for indi- in such extreme cases, samples are unlikely to be absolutely vidual samples are rarely provided (but see Miller 2002; pure due to the presence of contaminants introduced during Fairbairn 2007), and it is more common to find key words collection, storage or disturbance, or the storage of several relating to context. Where possible, we assigned samples to species together. Some plant material originally deposited one of eight context categories (containers, burnt destruc- in storage contexts and so originally dense and pure may tion deposits, pits, internal fire installations, external burnt have been subsequently disturbed or re-deposited, result- areas, refuse deposits, internal spaces and external spaces) ing in decreased density and purity. The recognition of such as in Table 1. Some of these relate to potential storage (e.g. store-derived samples may thus increase the likelihood of containers) and some to the disposal of refuse, whilst others successfully recognising deliberately collected plant taxa, are more ambiguous (e.g. internal spaces). as could the identification of processing for use. 13 452 Vegetation History and Archaeobotany (2019) 28:449–463 Fig. 1  Map of the Pre-Pottery Neolithic sites in southwest Asia Dhra 2, 32: el-Hemmeh, 33: Wadi Fidan A, 34: Wadi Fidan C, 35: with archaeobotanical remains included in the database. Black sym- Basta I, 36: Tell Ghoraifé, 37: Tell Aswad, 38: Dhuweilla and 39: bols indicate sites for which sample-level data was available; white Azraq 31. Northern Levantine sites—40: Tell Ain el-Kerkh, 41: Tell symbols indicate sites for which sample-level data was unavailable. Ras Shamra, 42: Tell Qaramel, 43: Tell Abr, 44: Dj’ade, 45: Halula, Central Anatolian sites—1: Hacılar, 2: Erbaba, 3: Çatalhöyük East, 46: Jerf el Ahmar, 47: Mureybet, 48: Abu Hureyra, 49: Douara Cave, 4: Can Hasan III and 5: Aşikli Höyük. Cypriot sites—6: Mylouthkia, 50: Cafer Höyük, 51: Gritille, 52: Nevali Çori, 53: Göbekli Tepe, 7: Ais Yiorkis and 8: Kastros. Southern Levantine sites—9: Hayonim 54: Tell Sabi Abyad II, 55: El Kowm I & II, 56: Çayönü and 57: Tell Cave, 10: Yiftahel, 11: Nahal Oren, 12: Kebara Cave, 13: Atlit- Bouqras. Sites of the eastern Fertile Crescent—58: Hallan Çemi, 59: Yam, 14: Gilgal, 15: Netiv Hagdud, 16: Jericho, 17: Nahal Hemar, Demirkoy, 60: Kortik Tepe, 61: Tell Maghzaliyeh, 62: Qermez Dere, 18: Wadi Faynan 16, 19: Shkarat Msaied, 20: Beidha, 21: Ayn Abu 63: Yarym Tepe, 64: Nemrik 9, 65: Mlefaat, 66: Jarmo, 67: Chogha Nukhayla, 22: Tell Ramad, 23: Gesher Benot Yaaqov, 24: Ohalo Golan, 68: Sheikh-e Abad, 69: Chia Sabz, 70: Tepe Ali Kosh, 71: II, 25: Gesher, 26: Wadi al-Hammeh 27, 27: Iraq ed-Dubb, 28: Ain Ganj Dareh Tepe, 72: Tepe Abdul Hosein and 73: Chogha Bonut Ghazal, 29: Wadi el-Jilat 13, 30: Wadi el-Jilat 6 & 7, 31: Zahrat adh- Compositional analysis of deliberate collection (cf. Halstead 1994, where the same numerical cut-off was used to indicate the deliberate cultiva- Sample density could not be used routinely in our analysis tion of crops). because the volume of sediment was often not recorded in Sample purity can be quantified using a diversity index, archaeobotanical reports. As an alternative, therefore, we based on the number of species and the evenness in their used absolute counts of plant seeds. While this is problem- abundance. For our purposes, the Simpson index D (Simp- atic, because total counts are partly determined by sample son 1949) is appropriate because, unlike the well-known density but also by the size of the sample taken, seed counts Shannon index (Shannon and Weaver 1948), it is primar- are almost always available. In addition, samples with a total ily influenced by evenness, such that the number of species count of less than 50 seeds were excluded from our analyses making up the minority component of the sample does not as they were considered too small to be reliable indicators substantially alter the score (Morris et al. 2014; see also 13 Vegetation History and Archaeobotany (2019) 28:449–463 453 Table 1  Context categories and the types of contexts included in each the lower end of the range, and dividing the ­log10 value by 4 category (the number of ­log10 steps between 1 and 10,000) to arrive at Context category Context types a scale of 0 to 1. This can be summarised as follows: Containers Clay bins, ceramic vessels log10 N C= Burnt destruction deposits Houses, rooms, internal spaces; with 4 evidence of extensive burning Pits Both internally and externally located where, C = normalised logarithmic count, N = raw counts. Internal fire installations Ovens, hearths, rake out, interior burnt For each sample, the new normalised logarithmic count areas (C) was combined with the Simpson index (D), by calculat- External burnt areas Fire-spots, burnt/ash deposits, exterior ing the Euclidian distance between a sample and the maxi- hearths mum normalised logarithmic score of 1 (i.e. 10,000 or more Refuse deposits Middens, dumps, trash/rubbish seeds) and the maximum Simpson index of 1 (i.e. only one Internal spaces Occupation layers/deposits, floors, taxon in the sample). This provides an indicator of deliber- houses, rooms; not extensively burnt ate collection (DC score), normalised to a scale of 0 to 1, External spaces Not further specified as follows: √ (1 − D)2 + (1 − C)2 DC=1 − ESM Fig. A for examples based on synthetic data). On a 0 √ 2 to 1 scale, the Simpson index assigns a high score to samples with low diversity (high purity) and vice versa. It is given by: Results ∑ ( n )2 D= The relationship of botanical composition N to archaeological context where, D = Simpson index, n = count for a particular taxon, N = count for all taxa. To investigate the relationship of compositional data to In addition, archaeobotanical material is often identifiable archaeological context, we examined “abundance versus only to genus or family. This may result in several species purity” plots of the count of seeds in each sample (plotted being included under a single taxonomic identification, giv- on a logarithmic scale) against the Simpson index. Many of ing the semblance of greater purity. Conversely, a sample the samples are neither rich in plant remains nor composi- may appear less pure when the same species is identified tionally pure, and are therefore concentrated in the lower left at different taxonomic levels as, for example, when some part of Fig. 2a. These small mixed samples are least likely to seeds are identified to species and others (usually less well represent deliberate collection and were therefore excluded preserved) are identified only to genus. The Simpson index from further analysis (ESM Equation A, for definition of should therefore be taken only as an estimate of a sample’s small and mixed). The resulting dataset (comprising 41 sites purity. and 477 samples) is restricted to samples that are either rich (in the upper part of Fig. 2a, b) or pure (to the right side of Data analysis Fig. 2a, b) or both (upper right of Fig. 2a, b). To incorporate archaeological context into this composi- For quantified samples with at least 50 seeds (1,381 sam- tional plot, samples in the restricted dataset were re-coded ples from 49 sites), we examined the relationship between according to the eight context categories in Table 1 (see the count of seeds (as a proxy for density) and the Simpson also ESM Table B). It is clear from these plots (Fig. 3 and index (as a estimate of purity), first in order to investigate the ESM Fig. C) that some context categories tend to be con- relationship of compositional data to archaeological context centrated in particular areas of the plot. Samples from con- and, secondly, to derive a measure of the likelihood of delib- tainers, for example, are primarily concentrated to the right erate collection. For these purposes, the counts were capped of the plot (Fig. 3a) because of their purity, and especially at 10,000 to provide a range with fixed start and end points, towards the top right, due their large size. These samples and then converted to a scale of 0 to 1 to make them com- almost certainly represent the remains of stored seed. A parable with the Simpson index. The count of 10,000 was group of three samples from containers are more mixed in selected as the upper limit because this encompasses most composition, clustering towards the left of the plot, which of the sample sizes in our database (ESM Fig. B). These raw may indicate secondary deposition in these containers. Sam- counts were then transformed by taking logs to the base ten, ples from burnt destruction deposits show a similar distribu- which gives greater weight to differences in sample size at tion to those from containers (Fig. 3b), but are rather more 13 454 Vegetation History and Archaeobotany (2019) 28:449–463 Fig. 3  Counts of plant items in archaeobotanical samples plotted on a logarithmic scale against Simpson index (small and mixed samples— see ESM Equation A—not plotted). Red (filled) symbols indicate samples from the following context categories: a containers; b burnt destruction deposits; c refuse deposits; d external burnt areas; e inter- nal fire installations; f pits. Contour lines indicate density estimations for each context category; multiple lines indicate increasing concen- trations of samples Fig. 2  Counts of plant items in archaeobotanical samples plotted on distribution to the samples from refuse deposits (Fig. 3d) a logarithmic scale against Simpson index. a All samples; b samples though some of these are of greater compositional purity, coded by DC score (small and mixed samples—see ESM Equation perhaps indicating the processing of plant material involving A—not plotted). The approximate location of the plant remains from fire at some of these locations. Internal fire installations are square F78c, Floor II, Hut 1 at Ohalo II is plotted as an asterisk in b even more widely distributed across the plot (Fig. 3e), with several particularly large pure samples located towards the dispersed. This is to be expected if they primarily represent top right of the plot. All of these are from ovens in burnt seeds stored within a building and charred during its destruc- destruction levels, so the plant remains probably derive tion, those samples to the right representing relatively pure from the final destruction of the building by fire rather than concentrations of stored products and those towards the left the earlier use of the ovens themselves. Other less specific resulting from mixing either between stored products or with context categories such as pits (Fig. 3f) and general clas- plant material lying on the floor of the building at the time sifications such as internal or external spaces (ESM Fig. C) of its destruction. also exhibit relatively wide distributions. Based on these At the other extreme, samples from refuse deposits tend results, we adopt a two-stage method for the recognition of to be concentrated towards the left of the plot (Fig. 3c), wild food plants: first we consider the likelihood that taxa reflecting their generally mixed composition, though some were deliberately collected, and secondly, from amongst the of these samples are very rich in plant remains. This clearly probably collected taxa, we distinguish those likely to have demonstrates that quantity alone is not a sufficient indication been collected as food from those more likely to have been of storage. Samples from external burnt areas show a similar collected for other purposes. At both stages, the quality of 13 Vegetation History and Archaeobotany (2019) 28:449–463 455 the evidence defines the certainty with which inferences Table 2  Deliberate collection (DC) scores for domesticated crops and about collection and use can be made and, inevitably, some wild progenitors with DC scores ≥ 0.5, grouped by domestication sta- tus. aPresumed status only inferences are more certain than others, as indicated below. Taxon Family DC The recognition of deliberate plant collection Domesticated crops Triticum monococcum Poaceae 1.00 The relationships between sample composition and archaeo- Lens culinaris Fabaceae 0.99 logical context clearly demonstrate that, as expected, likely Hordeum vulgare Poaceae 0.98 storage contexts tend to be associated with archaeobotanical Triticum dicoccum/monococcum Poaceae 0.86 samples that are both rich in plant remains and relatively Vicia erviliaa Fabaceae 0.81 pure in terms of species composition. It is also apparent that Hordeum vulgare var. nudum Poaceae 0.80 abundance of plant remains alone is as likely to be associated Hordeum vulgare distichum Poaceae 0.75 with rubbish deposits as with storage contexts. This allows Pisum sativum Fabaceae 0.74 us to assess the likely derivation of archaeobotanical sam- Triticum aestivum/durum Poaceae 0.70 ples using the Deliberate Collection (DC) score (Fig. 2b). Hordeum vulgare var. nudum/spontaneum Poaceae 0.67 The higher the score, the more rich and pure the sample, and Linum usitatissimum Linaceae 0.64 the more likely it is to represent storage; the lower the score Hordeum vulgare hexastichum Poaceae 0.61 the more likely it is that the sample represents discarded Triticum dicoccum Poaceae 0.57 material. Samples with intermediate DC scores may repre- Triticum spelta Poaceae 0.54 sent mixed or disturbed stored plant material or the debris of Domesticated crops/crop progenitors plant processing, though some may also derive from refuse Lensa Fabaceae 0.98 contexts. A concentration of plant remains from the northern Hordeum vulgare/spontaneum Poaceae 0.82 part of Hut 1 at Ohalo II, associated with a grinding stone Pisum/Vicia/Lathyrusa Fabaceae 0.69 (ESM Table C), has an intermediate DC score (0.57, Fig. 3), Cereal ­indeterminatea Poaceae 0.66 which is consistent with its interpretation as the remnants of Triticuma Poaceae 0.57 processing, as supported by starch grain analysis (Piperno Crop progenitors et al. 2004; Weiss et al. 2008). Pisum elatius Fabaceae 0.91 As well as recognising samples that represent deliberate Hordeum spontaneum Poaceae 0.85 collection, it is also necessary to distinguish which species Triticum boeoticum thaoudar Poaceae 0.84 in these samples were the objects of storage or processing, Triticum/Secalea Poaceae 0.83 while not overlooking the possible storage of mixed species or multiple episodes of processing. So, to allow the recogni- tion of up to three deliberately collected species per sample, taxa that comprise at least 30% of a sample indicated as rank taxa in order of importance, but rather by the quality deliberately collected are considered a potential component of the evidence for (and so the likelihood of) their deliberate of the stored or processed product (cf. Halstead 1994, where collection. the same percentage cut-off was used to indicate deliberate Alongside the wild species, we considered domesticated cultivation of crop species). Some seeds are identified only crops because these are likely to have been brought onto a to higher taxonomic levels, so all taxa (species, genus or site deliberately, and can therefore be used as an independent family identifications) that make up ≥ 30% of at least one benchmark for the DC scores likely to indicate a deliberately sample with a relatively high DC score (≥ 0.5), are listed in collected taxon. All of the southwest Asian founder crops Table 2 (domesticates and their wild progenitors) or Table 3 except Cicer arietinum (chickpea) make up ≥ 30% of at least (other wild taxa). Whilst the DC scores for each taxon may one sample with a DC score ≥ 0.5, and Triticum monococ- be based on only one sample (ESM Table D), it nevertheless cum (einkorn), Lens culinaris (lentil) and Hordeum vulgare provides the best single piece of evidence for that taxon’s (barley) have very high DC scores (> 0.9). Triticum dicoc- deliberate collection. Taxa in the upper part of Tables 2 and cum (emmer) derives from a sample with a relatively low 3 (from samples with the highest DC scores) are those with score (0.57) but its true representation is probably masked the strongest evidence for deliberate collection and storage. because many glume wheat grains are identified only as T. Taxa in the lower part of the tables (from samples with pro- monococcum/dicoccum, which makes up ≥ 30% of a sample gressively lower DC scores) are successively less likely to with a DC score of 0.86. Chickpea, on the other hand, is be from a single stored product, and are more likely to result quite rare overall in the archaeobotanical record for the PPN from processing for use, mixed stored products, or domes- (Zohary et al. 2012, p 89) and so may not have been widely tic refuse. It should be noted that Tables 2 and 3 do not cultivated. Such rarely found species are relatively unlikely 13 456 Vegetation History and Archaeobotany (2019) 28:449–463 Table 3  Deliberate collection (DC) scores for wild plant taxa (excluding crop progenitors) with DC scores ≥ 0.5 13 Vegetation History and Archaeobotany (2019) 28:449–463 457 to be represented in stored deposits, which are themselves Tables 3 and 4 include several fruit and nut taxa (Pista- quite rare. cia, Ficus, Prunus/Amygdalus, Celtis, Lycium), which may We also considered wild crop progenitors because these well have been collected as food. The seeds of many of the must have been deliberately collected at some stage prior other wild taxa are also edible, including Poaceae, Fabaceae, to their domestication. Their representation is complicated, Brassicaceae, Cyperaceae, Atriplex and Polygonum/Rumex. however, by problems of identification (see, for example, Plants collected for their seed have a good chance of being Zohary et al. 2012). Wild and domesticated forms of Lens preserved archaeologically, especially if the seeds are spp. (lentil), Cicer spp. (chickpea) and Vicia ervilia (bitter parched during processing, as is common for Polygonum vetch) are very difficult to distinguish, and distinctions are (Hillman 2000, p 357). The leaves and shoots of several of often based on size alone (or even the date of the deposit). these taxa are also known ethnographically to be eaten as Lens orientalis and Cicer reticulatum, the wild forms of greens, as are the leaves and shoots of other taxa in Table 4 lentil and chickpea, and bitter vetch are rarely (if ever) whose seed is not normally eaten (e.g. Lactuca, Juncus, recorded as such in archaeobotanical reports. Identifications Malva sylvestris and Suaeda). The tubers of Bolboschoenus to genus are therefore very common and include both wild maritimus, and other Cyperaceae species such as Cyperus and domesticated forms. The problem of identification is esculentus, are another potential source of food (Simpson less severe for the cereals, but while the two-grained form and Inglis 2001; Dogan et al. 2004; Holec et al. 2014). These of wild einkorn (T. boeoticum thaoudar) is often recorded tubers are fully formed at about the time when plants are as such, the one-grained form (T. boeoticum aegilopoides) in seed (Jordan-Molero and Stoller 1978; Davis 1986), and is usually included in the more general category “T. boeoti- seeds may be deposited at settlements if, for example, the cum”. Despite these complications, crop progenitors are also above-ground parts were gathered during the uprooting of well represented in Table 2, by species that can be accurately below-ground organs. Some of the plants in Table 4 can also identified as wild—pea (P. elatius) and barley (H. sponta- be used for their oil or as flavouring, (e.g. Brassicaceae and neum), and potentially by Lens sp., all of which are found in Capparis). samples with a DC score > 0.8. T. boeoticum is also found As well as food, there is a variety of other ethnographi- in a sample with a DC score of 0.45. cally attested plant uses (Table 4). For example, many plants Overall, therefore, the representation of domesticated have been used for their medicinal properties, and those for crops and their progenitors suggests that taxa making up which their seeds are utilised in this way include Atriplex, greater than 30% of a sample with a DC score ≥ 0.5 provide Descurainia sophia and Ziziphora. Reeds and rushes are a reasonable measure by which to assess the likelihood of commonly collected for use as matting, bedding or in con- a taxon being deliberately collected for use. On this basis, struction but, as has been noted for B. maritimus and Jun- about 40 genera of wild plants (excluding crop progenitors) cus (Hillman 2000), they are often used before they come from about 20 families may be considered as potential can- into seed. Plants could also be used for dyeing (e.g. Arnebia didates for deliberate collection (Table 3). decumbens) or decorative purposes (e.g. Crucianella, Heli- anthemum and M. sylvestris), and there is one instance of Distinguishing wild plant foods from plants Helianthemum seeds stored in a pot within a shrine (at Çatal- collected for other uses höyük) for an unknown, but presumably socially significant, reason. Even when a case can be made for the deliberate collec- Although the taxa listed as potential food plants in tion of wild plants, it cannot be assumed that they were Tables 3 and 4 have met our criteria for deliberate col- collected for food. This must be inferred from the plant’s lection in at least one context, it does not necessarily fol- characteristics (e.g. whether edible, palatable or nutri- low that they arrived on site as foods wherever they were tious), ethnographic evidence and depositional context, found. There are also incidental routes by which potential such as association with evidence of food preparation or food plants may have reached a site as, for example, when consumption. Here we define wild food plants as those fruits or seeds were collected unintentionally with crops, consumed as staples, nutritional supplements or flavour- foraged wild species, woody material or animal dung. For ings, and Table 4 gives the recorded uses for food or other example, some mixed samples are very large (those in the purposes of all the wild taxa listed in Table 3 as poten- upper left of Fig. 2a, b), and it is tempting to conclude tial candidates for deliberate collection. The purpose of that these represent the remains of plant food processing, Table 4 is to assist in distinguishing taxa that are likely to especially when they derive from hearths or other areas have been collected as food from those more likely to have of burning. A closer examination of the botanical compo- been collected for other purposes, rather than to differenti- sition of such samples, however, suggests an alternative ate between different types of non-food use. interpretation. 13 Table 4  Recorded uses of the wild plant taxa listed in Table 3, taxonomically ordered. NB: no known recorded uses for Helianthemum (ledifolium), Aizoon (hispanicum) and Crucianella. 458 ● ➝Widely recognised use. ○➝Occasionally/rarely recognised use 13 Vegetation History and Archaeobotany (2019) 28:449–463 Vegetation History and Archaeobotany (2019) 28:449–463 459 Many of these large, mixed samples are from Neolithic grasses (Poaceae, especially Avena sterilis and Hordeum middens at Çatalhöyük, and the plant remains in these bulbosum/murinum), fruits and nuts (Prunus/Amygdalus, deposits are thought to derive partly from discarded house- Pistacia and Ficus), members of the Brassicaceae (espe- hold refuse and partly from in situ burning (Bogaard et al. cially Brassica/Sinapis) and other herbaceous species (B. 2013). A comparison of the charred plant remains in sam- maritimus, Lactuca, and Polygonum/Rumex species), of ples from one of these middens (Area 181) shows that both which Lactuca and Rumex are primarily eaten as greens and fire-spots and midden deposits exhibit a similar range of are less likely to be represented by their seeds. Of these, botanical compositions (see, for example, the adjacent sam- only Avena sterilis, Prunus/Amygdalus, Brassica/Sinapis, ples D69 and D72 in ESM Fig. D), suggesting that the plant B. maritimus and Polygonum came from a sample with a remains derive from the same source. Many of the dominant DC score of ≥ 0.7 and so from the area of the “abundance- taxa in these samples grow in saline habitats (Aeluropus, purity” plot associated with storage contexts (Fig. 3a, b). B. maritimus) or marshy areas (rushes), and others have B. maritimus and Polygonum/Rumex may also have been small or hard-coated seeds (small-seeded legumes), Che- deposited on some occasions as a result of the burning of nopodiaceae, Brassicaceae, Helianthemum (ledifolium) dung, whether from wild or managed animals. Though it has that survive passage through the ruminant digestive system been argued that the collection of dung from wild animals (Miller and Smart 1984; Charles 1996; Wallace and Charles is unlikely (Hillman et al. 1989, 1997; Savard et al. 2006), 2013). These, and several less dominant small-seeded ethnographic evidence and counter-arguments have been taxa (Sporobolus (saline habitats), Chenopodium cheno- presented to support the collection wild animal dung as a podioides, Juncus (marshy areas), Alopecurus, Artemisia source of fuel (Miller 1996, 1997). annua), are suggestive of grazing habitats, and the burning Using the same criteria, a greater number of wild taxa of dung fuel has been suggested as a major contributor of the were recognised as potentially collected in later PPN periods plant remains found in these deposits (Bogaard et al. 2013; (Early PPNB to PPNC), after the emergence of domesti- Filipović 2014). The Çatalhöyük samples are not unique in cated crops. These include Descurainia (sophia), Capsella this respect; for example, Epipalaeolithic samples from Abu and Sisymbrium species (and perhaps other members of the Hureyra, including a large mixed sample, which plots with Brassicaceae), fruits and nuts (Ficus, Pistacia and Celtis the large Çatalhöyük midden samples, are also composed (tournefortii)), wild grasses (including Taeniatherum caput- of taxa consistent with derivation from dung (Miller 1996). medusae, Poa, Aeluropus, Alopecurus, Crypsis, Agrostis, So, even taxa that were sometimes brought to site as foods Bromus and Lolium species), wild legumes (Fabaceae), both could, at other times and places, have arrived on site by large-seeded (Vicia/Lathyrus) and small-seeded (including other routes. These include taxa likely to survive ruminant Trigonella (astroites), Astragalus, and Melilotus species), digestion (e.g. Cyperaceae—especially B. maritimus—Jun- and a wide range of other herbaceous species, including cus, Atriplex, Polygonum, Rumex and small-seeded legumes) members of the Boraginaceae (Buglossoides arvensis, B. that are as likely to have been deposited in dung fuel as col- tenuiflora and Arnebia decumbens) (Table 3). Of these, only lected for food, especially with the development of animal the Brassicaceae and Boraginaceae species, Ficus, Celtis, management (Matthews 2016). Other plants, including some Taeniatherum caput-medusae, Poa, Lathyrus and the her- of the same taxa, may also have been collected for purposes baceous species, Helianthemum, B. maritimus, Atriplex, other than food, such as for building materials (Cyperaceae, Suaeda and Lycium came from a sample with a DC score especially B. maritimus and Juncus) or dye plants (Arnebia of ≥ 0.7, suggestive of storage (Fig. 3a, b). The use of dung decumbens). The use of these taxa as food cannot therefore fuel is also well documented in this period (Matthews 2016), be universally inferred. which may account for some of the occurrences of taxa such as Helianthemum, B. maritimus, Atriplex and Suaeda. The chronological and geographical distribution Few taxa were recognised as likely to have been delib- of deliberately collected wild food plants erately collected in more than one of the five geographical regions in Fig. 1 (central Anatolia, Cyprus, southern and We compared the wild plant foods recognised in pre-domes- northern Levant, and the eastern Fertile Crescent). Those tication periods with those following the advent of domes- that do occur widely tend to be taxonomically diverse ticated crops in order to evaluate the evidence for a narrow- groupings, such as small-seeded legumes, Brassicaceae and ing of the wild plant food spectrum during the transition to small-seeded grasses (Table 3). The exceptions to this are agriculture. For samples dated to the PPNA or earlier, and B. maritimus, Polygonum and/or Rumex, which are found so before the emergence of domesticated crops, several taxa in multiple regions (ESM Table D). Moreover, within each were identified that best meet our criteria for recognition region it is unusual for a taxon to be recognised as deliber- as wild plant foods, based on their composition—(Table 3) ately collected at more than one or two sites. Indeed, only and known use as food—(Table 4). These include wild 12 wild taxa excluding crop progenitors were found at more 13 460 Vegetation History and Archaeobotany (2019) 28:449–463 than one site, and many of these are those likely to be over- in the form of storage. On the other hand, samples with represented due to preservation bias—such as Boraginaceae intermediate DC scores provide more ambiguous evidence species, the nutlets of which are encased in a siliceous outer of deliberate collection, as they may derive from a number coat, or fruits represented by their robust waste product (fruit of different sources, including plant processing and mixed stones). Rather than indicating genuine regional patterns in storage, but also refuse or animal dung. This ambiguity also plant use, this probably reflects the difficulty of finding reli- applies to the large but mixed samples commonly recovered able evidence for deliberate collection. from refuse deposits, where such large quantities of charred remains are more likely to result from the deliberate burning of dung fuel than the accidental burning of food processing Discussion remnants (Miller and Smart 1984; Miller 1984). Many, if not most, of the taxa from these samples may therefore derive The recognition of deliberately collected food plants from the use of dung fuel. So, while both storage and processing constitute equally When recognising wild plant collection, there are two valid evidence for plant collection, storage can be more reli- aspects that need to be considered: first, what constitutes ably indicated from compositional evidence (purity and den- strong evidence for deliberate collection and secondly, how sity) alone whilst the recognition of processing must remain reliably can we recognise this evidence? Both storage for uncertain unless backed up by strong contextual evidence future use and processing prior to use can be considered such as their association with ground stone tools. Such evi- strong evidence of collection. However, for pre-agricultural dence can be difficult to find, however, so while it is possible communities exploiting (and possibly cultivating) a range to recognise taxa that were deliberately collected it is more of wild plants available in different seasons, storage may difficult to say which taxa were not. have been a less essential (and, if these communities were also mobile, even impractical) element of plant usage than it is for agricultural communities based on grain crops avail- The broad spectrum of plant food collection? able in one season only. At the Upper Palaeolithic site of Ohalo II (Weiss et al. 2008) and the EPPNB site of Tell We have brought together the botanical composition of sam- Qarassa (Arranz-Otaegui et al. 2016), for example, there ples, their archaeological context and the potential use of is contextual evidence for wild plant food processing but the taxa dominant in samples, to systematically recognise not for storage. Evidence for feasting in the form of large- those wild plants for which a good case can be made for their scale food processing and cooking also increases through deliberate collection as food. On this basis, relatively few of the PPN period (Twiss 2008), which provides greater oppor- the wild taxa found at pre- and early agricultural sites can tunity for plant material to be preserved through charring. be confidently recognised as contributing to the human diet Moreover, taxa that are primarily represented by their waste at one time or another during the PPN, though we acknowl- products, such as fruit stones and nutshell, are more likely to edge that this group represents the minimum number of wild be found in processing than storage contexts. For this reason, taxa used as food. In addition, some foods, such as fruits our approach has focused on recognising both the storage and nuts, may be overlooked because they are usually rep- and processing of wild plant material in the archaeological resented by the waste by-products of consumption, such as record. seeds and shell, rather than by the product consumed. With regards to reliability, however, while contextual evi- We can now re-evaluate the spectrum of wild food plants dence of plant processing is occasionally found, as at Ohalo at pre-agricultural and early agricultural sites. One of the II (see above; Weiss et al. 2008), the most unambiguous largest groups of wild taxa represented on pre-agricultural evidence for deliberate collection of a wild plant is usually (PPNA and earlier) sites is the grasses, and these are usu- evidence of its storage. Our comparison of contextual evi- ally assumed to have been collected for food. Our evidence dence and sample composition across southwest Asia has indicates, however, that of the 40–50 genera of grasses rep- indicated that, where good contextual information is avail- resented in southwest Asia, there is strong evidence for the able, it is only likely storage contexts (such as containers deliberate collection of only ten genera, each of which may and burnt destruction deposits) that are associated with represent no more than one collected species. On the other archaeobotanical samples that are both rich and pure; other hand, our evidence has provided support for the deliberate known archaeological contexts (such as refuse deposits and collection of some taxa that have previously been only tenta- external burnt areas) have not produced such samples. Thus tively suggested as plant foods on the grounds of their over- compositional evidence alone (a high DC score indicating all abundance or ethnographic use, for example B. maritimus a combination of seed abundance and sample purity) pro- (Savard et al. 2006, p 189), Atriplex (de Moulins 1997, p 93) vides a reliable indication of deliberate collection, usually and Polygonum (Hillman 2000, pp 357–358). 13 Vegetation History and Archaeobotany (2019) 28:449–463 461 To some extent, therefore, our results have called into context, with the aim of establishing a robust link between question the exploitation of a wide range of plant species archaeological evidence and its interpretation in terms of by pre-agricultural communities in southwest Asia in the deliberate collection and use of plants as food. This has put period leading up to domestication. Moreover, comparing the recognition of wild plant foods on a firm footing, and has the evidence from pre-agricultural sites, for which few wild led to the recognition of a suite of wild plant taxa for which food plants have been confidently recognised, with that from there is strong evidence for their exploitation as plant foods later proto- and early agricultural sites (EPPNB to PPNC), at pre-agricultural and early agricultural sites in southwest there is evidence for the collection of a broader range of Asia. This has shown that relatively few of the wild taxa wild plant foods at the latter, though this may reflect the found at pre- and early agricultural sites can be confidently greater quantity of evidence and the more frequent occur- recognised as contributing to the human diet. The approach rence of unambiguous storage contexts and cooking or other adopted here has resulted in a re-evaluation of the evidence processing involving fire. For these reasons, while we do not for the exploitation of a broad spectrum of wild plant foods interpret this contrast as indicating an increase in the diver- at pre-agricultural sites, and its supposed narrowing during sity of wild plant foods collected at these later PPN sites, the early development of agriculture. This has implications we have found no evidence for a narrowing of the plant food for how we understand the processes leading to the domes- spectrum during the adoption of agriculture. tication of crops, and points towards a mutualistic relation- Rather, our results, based on taxa for which there is reli- ship between people and plants as a driving force during the able evidence of their deliberate collection as food, suggest development of agriculture. little change in the variety of wild plant foods exploited between pre-agricultural and proto-/early agricultural peri- Acknowledgements This research was funded by a European Research Council (ERC) grant ‘The Evolutionary Origins of Agriculture’ ods, despite the availability of potentially higher ranked (Grant no. 269830-EOA, PI Glynis Jones) and a Natural Environment domesticated crops in the later period. This implies a per- Research Council (NERC) grant ‘Origins of Agriculture: an Ecologi- sistence in opportunistic foraging throughout the PPN, and, cal Perspective on Crop Domestication’ (Grant no. NE/H022716/1, if it occurred in resource-rich environments and encouraged PI Colin Osborne), both at the University of Sheffield. The database used builds on an earlier database compiled by Sue Colledge during increased sedentism, would have provided ample opportu- ‘The Origin and Spread of Neolithic Plant Economies in the Near East nity for experimentation in new exploitation techniques, dur- and Europe’ project (AHRB, PIs Stephen Shennan and James Conolly, ing which time the cultivation of a range of different species University College London) and the ‘Domestication of Europe’ project could be trialled (Smith 2007, 2011b, 2016; Zeder 2012, (NERC, PI Terry Brown, University of Manchester). We would like to thank the following for access to unpublished archaeobotanical data: 2015, 2016). Andy Fairbairn, Dragana Filipović, Rainer Pasternak, Manon Savard, An important consequence of this interpretation of the Chantel White, Jade Whitlam and George Willcox. We are also grate- archaeobotanical evidence is that it does not require an ful to Amy Bogaard and Paul Blackwell for comments and advice on explanation for the emergence of agriculture to be based on aspects of the research. an externally-driven demographic or environmental “push” Open Access This article is distributed under the terms of the Crea- model, whereby foragers were forced to exploit lower ranked tive Commons Attribution 4.0 International License (http://creat​iveco​ plant species (in response to resource depletion) that sub- mmons​.org/licen​ses/by/4.0/), which permits unrestricted use, distribu- sequently declined with the advent of domesticated crops. tion, and reproduction in any medium, provided you give appropriate Rather, continued opportunistic foraging may have provided credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. a context for the development of a mutualistic relation- ship between people and certain plants, with both taking advantage of favourable conditions, a diverse and plentiful resource base for people, and a rich anthropogenic environ- References ment for plants—that is to the evolutionary benefit of both and led ultimately to crop domestication and a dependence Arranz-Otaegui A, Colledge S, Ibañez JJ, Zapata L (2016) Crop hus- bandry activities and wild plant gathering, use and consumption at on agriculture (Rindos 1980, 1984; Smith 2007, 2011b; the EPPNB Tell Qarassa North (south Syria). 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