New evidence for the processing of wild cereal grains at Ohalo II, a 23 000-year-old campsite on the shore of the Sea of Galilee, Israel.

Nadel, Dani ; Piperno, Dolores R. ; Holst, Irene 等

Introduction

The site of Ohalo II, located on the south-western shore of the Sea of Galilee in Israel, was a fisher-hunter-gatherer campsite dating to about 23 000 years ago (Nadel et al. 1995; Nadel 2002). Immediate and quick submergence of the site after it was abandoned provided remarkable preservation conditions (Belitzky & Nadel 2001; Tsatskin & Nadel 2003). The site was submerged for millennia, and thus a wide array of material remains were well preserved in situ. Of these, the most significant are the brush huts and the finds exposed on their floors.

As reported previously (Nadel & Werker 1999; Nadel 2002), the remains of six brush huts were identified during fieldwork, with several concentrations of hearths around them (Figure 1). Four of the brush huts were fully excavated, and two were comprehensively sampled. They all had a bowl-like cross-section, indicating that the inhabitants made shallow depressions in the soft bedrock before constructing the huts. Four huts were oval in general shape, while two had a kidney-like shape with an entrance from the east. The construction materials of Brush Hut 1, the largest at the site, were identified as thick branches of Tamarix (Tamarisk), Salix (Willow) and Quercus ithaburensis (Oak), covered by smaller branches of plants such as Atriplex/Seidlitzia (Orach/Seidlitzia) and Prosopis (Mesquite), as well as leaves and grasses (Nadel & Werker 1999: tab. 2).

Brush Hut 1 had three successive floors. Grass bedding was well preserved on the bottom floor; it was made of bundles of one species, Puccinellia (Nadel et al. 2004a). Evidence for other floors with grass bedding, badly preserved, was found on floor II and in other brush huts. Tiny fragments of twisted fibre were found in this brush hut, reflecting the possible use of cords, strings, nets etc. (Nadel et al. 1994). Wooden objects were found in several brush huts (Nadel et al. 2006).

Most remarkable are the charred and even uncharred seeds and fruit. So far a sample of c. 150 000 specimens has been studied (Kislev et al. 1992; Simchoni 1997; Weiss 2002; Nadel 2004). Most of these are well preserved, and they represent c. 150 taxa, many of which have been identified to species level. The floral assemblage directly informs the diet of the inhabitants, as well as the immediate mosaic of plant communities and the environment in more general terms. The wealth of organic remains also enabled the readings of over 40 [sup.14]C dates, which gave a calibrated range of 22 500-23 500 BP (Nadel et al. 1995, 2004b; Nadel 2002).

The skeleton of one adult male (c. 40 years old) was found in a shallow grave. The individual was about 1.73m tall and right-handed. He was buried on his back with hands folded on the chest, knees folded and head turned to the east (Nadel & Hershkovitz 1991; Hershkovitz et al. 1993, 1995). Other finds include a wealth of faunal remains, of which fish bones are particularly abundant (Zohar 2002). Within the mammals, gazelles are by far the most common, followed by fallow deer and smaller species (Rabinovich & Nadel 2005). More than 80 species of birds were identified (Simmons & Nadel 1998), and a range of micromammals was also studied (Belmaker et al. 2001). Within the brush huts and near the hearths we found a rich assemblage of flint artefacts (Nadel 2001), ground stone tools, worked bone tools (Rabinovich & Nadel 1994-95), as well as dentalium and columbella beads from the Mediterranean Sea (Bar-Yosef Mayer 2002).

Of pivotal interest is a flat basalt stone that was firmly set on the second floor (floor II) of Brush Hut 1 (Figure 1) on a patch of sand and supported by several cobbles (Figure 2; Nadel 2003). The stone has been studied, including through starch grain analysis (Piperno et al. 2004) as well as the distribution patterns of the seeds around it (Weiss et al. 2008). This analysis indicated that the stone had been employed in the grinding of wild cereals. Such is the importance of this finding for the Late Upper Palaeolithic and Early Epipalaeolithic economy and subsistence that further scientific examination of the stone and its neighbouring deposits was undertaken. The purpose of this paper is to report and assess the results of this second analysis, in particular the identification of traces of foodstuffs on previously untested areas of the artefact (e.g. its lower and certain sections of its top surface), and in the sediments surrounding it.

[FIGURE 1 OMITTED]

New starch grain studies at Ohalo II

When uncovered, the stone appeared to have been carefully positioned on a small patch of sand supported by small pebbles, suggesting it functioned as an anvil or grinding stone. The original starch grain study utilised a modern reference collection of plants that included all of the approximately 15 grass genera, and all but one grass species identified by Ehud Weiss from the macrobotanical remains found on floor II of Hut 1, and from immediately around the stone itself (Piperno et al. 2004 for lists and descriptions). This collection represents virtually all of the grass taxa found throughout the site. Additionally, modern collections of starch grains from non-grass species commonly found on floor II and other large reference collections described in other studies, together with the published literature, were used in the study (see Piperno et al. 2004). The analysis of a sample from the upper stones' surface resulted in the recovery of 150 starch grains, of which 127 were from grasses. Seventy-eight out of the 127 grass grains had the size and morphological characteristics found to be shared only by Triticurn, Aegilops, and/or Hordeum, members of the Triticeae tribe of grasses, leading us to propose an AHT starch group (Piperno et al. 2004). Twenty-three grains out of the 78 in the AHT group had morphological features found only in Hordeurn and this finding, together with the absence of specific grain types characteristic of either Triticurn or Aegilops, led us to suggest that seeds from these two genera were probably not processed with the stone. In sum, the evidence indicated that most of the identifiable grass starch recovered from the stone was probably from Hordeum, and the processing of selected wild grasses was taking place well before their domestication in south-west Asia.

[FIGURE 2 OMITTED]

Sediments from Brush Hut 1 were not analysed in the first study, and the underside of the stone not tested for starch presence. In this study we looked at the bottom surface and an additional area of the surface of the upward-facing facet of the stone (Figure 3) that had yielded 150 starch grains in the first study (Table 1). In addition, six sediment samples from two successive floors from Brush Hut 1 were analysed (Table 2). These include the second floor (II), where the grinding stone was set, and the third floor (the bottom one, III). These samples serve as control samples aiming to test whether starch granules from the stone simply represented a general distribution of starch within the site.

[FIGURE 3 OMITTED]

Starch extraction and identification procedures followed those in Piperno et al. (2004), with the exception that ultrasound was not employed as a step in starch isolation. Since our original study, research has been carried out on modern plants from the Middle East and Europe that included: 1) other genera of the Triticeae tribe such as Elymus spp., Leymus spp., and Secale spp.; 2) additional species of Tridcum, Hordeum, Aegilops, and Secale; and 3) grass genera from other Poaceae tribes (e.g. Henry & Piperno 2008; Henry et al. 2009, 2011). This work confirms that starch grains from the AHT taxa differ in size and morphology from other members of the Triticeae as well as other Poaceae, and that individual genera in the AHT group may be identified and/or ruled out from representation in archaeological samples using a starch grain assemblage approach (such as the one employed in our first study). For this study, we also carried out additional modern reference work that involved comparing and contrasting more closely a type of starch grain found in some grasses, compound polyhedral forms, in order to determine if another important genus of grass, Avena, could be identified. As will be discussed in more detail below, it appears that Arena starch grains do possess diagnostic attributes not seen in other grasses studied.

Results

In the six studied control sediment samples, two contained one starch grain each, and no starch was recovered in the others (Table 2). Plant remains such as pollen, spores, phytoliths, tissue fragments, and charcoal were, however, common. This finding supports our initial conclusion that starch recovered from the upper surface of the stone primarily represents plants processed on the stone, and not contamination from associated sediments.

It should also be remembered that a variety of field observations and taphonomic studies clearly indicate that the remains on floors in general, and in Brush Hut 1 in particular, were mostly in situ, sometimes hardly having moved since the site was abandoned and inundated. These records include the charred contour of the floors (Nadel & Werker 1999; Nadel 2003), thin-section studies (Tsatskin & Nadel 2003), the setting of the grinding stone, and the non-random distribution patterns of flints (Nadel 2001), fish bones (Nadel et al. 1994) and especially seeds (Weiss et al. 2004a, 2004b, 2008, and see below). The taphonomic background is further support for the conclusion that the starch grains found on the upper surface of the stone represent the stone's function as a grinding implement.

With regard to the additional tests carried out on the grinding stone itself, analysis of the central area of the bottom side of the stone revealed a total of 13 starch grains. One of them is probably from Hordeum, another is from the AHT group, eight are bell-shaped grains, and three are compound grains from grasses. This result can be compared with the additional analysis of the upper side of the stone. In the first analysis, which yielded 150 starch grains, we sampled about two-thirds of its total area (Figure 3). In this study, the remaining, central portion of the stone was sampled by brushing it with a toothbrush, followed by collecting the residue through rinsing with water, and lastly floating the residue with caesium chloride to isolate the starch. This analysis resulted in the removal of a total of 49 starch grains. The large number of grains recovered from the upper surface of the stone in both analyses, together with the considerable difference in starch grain quantities between the top and bottom of the stone, further support our previous interpretation that the stone's function was to grind plants, and that this activity was carried out on its upper side. The starch grains recovered from the stone's bottom may represent plants lying on the ground surface before the stone was placed there, or the starch may have adhered to the stone when it was handled. Another possibility is that this facet was less frequently used as a grinding surface.

The taxa represented by starch grains from the second analysis of the stone were in close agreement with those previously identified. Out of the 49 grains recovered, five were discoidal (called "lenticular" in Piperno et al. [2004: 670]) shapes from AHT taxa, and three of these were identifiable as Hordeum (see Piperno etal. 2004 for a complete discussion). Nine grains were bell-shaped forms that can occur in a number of taxa, including AHT grasses, and the 35 other grains were compound, polyhedral forms from grass seed starches. As in the previous stud> none of the discoidal grains had the crater-like impressions characteristic of Triticum or lamellae or 'pleats' like those in Aegilops species, suggesting that most of the grains from the AHT group are from Hordeum. The Hordeum discoidal grains are once again most similar to those from H. spontaneum, H. glaucum, and H. bulbosum (see Piperno et al. 2004), but we cannot distinguish between the three. As in the first analysis, Hordeum species such as H. marinum can be ruled out on the basis of size characteristics.

We are now able to make a positive identification of an additional grass taxon that was not recognised in the previous study. Eleven out of the 35 compound, polyhedral grains appear to be from the genus Avena (Figure 4). We made this determination on the basis of a more detailed study of modern starch grains from this genus, and their comparison with other taxa. The modern Avena species studied were Arena barbara, A. byzantina, A. fatua, and A. sterilis, which were available to us from EW's collections and folders housed at the herbarium of the National Museum of Natural History (Table 3). Seeds from A. barbata and A. sterilis occurred at Ohalo II. Arena starch grains have numerous small and distinctive pressure facets due to the grains packing together very tightly in the amyloplasts during their formation (Figure 5). These types of compound polyhedral starches were also described for Arena by Henry & Piperno (2008) and Henry et al. (2009), who considered them a possible genus-specific trait. Neither we nor other investigators have described this type of grain in other taxa, and we now consider them to be diagnostic of Avena. A re-evaluation of the 56 compound polyhedral grains isolated from the stone in the first study indicates that two of these can be identified on a morphological basis as Arena. A size difference can be seen between the modern and archaeological Arena grains (Table 3). The latter had a mean length of 15.8 [micro]m (range: 10-18 [micro]m) and the modern grains including the domesticated exhibited a mean length of 6-12 [micro]m (range: 4-18 [micro]m) (Table 3). It may be that another wild species of Arena was utilised or, alternatively, that the smallest Arena grains are for some reason not well represented on the stone. The former is probably more likely. In summary, it appears that Arena was also processed on the stone, although only a few of its seeds were found on the hut floor. The remainder of the compound, polyhedral grains recovered can belong to any number of genera in the Poaceae, including Arena and the AHT taxa.

[FIGURE 4 OMITTED]

This new identification of Avena granules holds a special importance for our reconstruction of human diet at the Ohalo II camp. The amounts of oat grains present were meagre in relation to other grasses found at the site. For example, on the second floor in Hut 1, where the grinding stone came from, 35 oat grains were found out of a total of c. 16 000 grass grains; 19 Avena grains were from A. barbara, 6 were from A. sterilis, and 10 were identified as A. barbata/sterilis. The third floor, situated directly underneath the second floor and covered by mat-like bedding from Puccinelia cf. convoluta stalks (Nadel et al. 2004a), was richer in Arena: 357 oat grains were recovered with 100 from A. barbara, 99 from A. sterilis, and 158 from A. barbata/sterilis (out of c. 15 000 grass grains).

[FIGURE 5 OMITTED]

The Avena numbers are two orders of magnitude smaller than the total number of other grass grains from these two floors. However, as far as we are aware it represents the earliest direct evidence for the use of oats for human consumption. Moreover, we have already demonstrated (Piperno et al. 2004) that the starch granules from the grinding stone most probably represent the grinding of grass grains and the use of the flour-like product for baking. If so, oats were part of the diet at Ohalo II, together with other cereals and grasses (Weiss et al. 2004b, 2008).

Discussion and conclusion

The new analysis of the Ohalo II grinding stone, along with a study of the associated sediments and further consideration of starch morphology in modern grasses from the region, indicate that the central part of the stone had captured high numbers of starch grains, further supporting its interpretation as a grinding implement. In contrast, sediments in the six tested squares from earlier and adjacent contexts contained little starch, showing that starch grains on the stone do not represent part of a general occurrence. Grasses remain the dominant type of plant present in the starch record, but to these wild oats can now be added.

The variety of cereals that were collected, processed and consumed at Ohalo II now includes barley, wheat and oats. In terms of food gathering, these reflect a close knowledge of the environment and specific habitats in the region, as well as the diversity of edible species and their availability. Furthermore, they indicate that the inhabitants chose not to rely on one species, but to invest in all three. However, if retrieved seed numbers reflect the relative importance of each species, then oats were much less important in the diet than barley and wheat.

The finds discussed here all come from a sealed floor (the second in a sequence of three), within a well-delineated brush hut. Importantly, the grinding stone was set on the floor in a firm pre-planned manner. Furthermore, there was a unique pattern of cereal grain distribution around the stone (Figure 6). The numbers were high (thousands), and the pattern was unmistakable. It is noteworthy that starch of cereals was retrieved in two separate analyses from the upper surface of the implement. The bottom surface had much lower quantities, while the sediment around the stone was very poor in starch content. It should also be stressed that other organic finds were well preserved in the same brush hut, including grass bedding and wood objects.

[FIGURE 6 OMITTED]

The aspect of local diet 23 000 years ago is also of significance. One should remember that the local inhabitants were hunting a wide range of herbivores (with an emphasis on gazelles), procuring large numbers of fish (most probably by traps and nets) and catching an astonishingly wide variety of birds (over 80 species). The plant remains were preserved in very high numbers on brush hut floors and around fireplaces, reflecting their central role in the diet. The variety of edible species in general, and of the cereals in particular, indicates the strong reliance on these, at least in the relevant seasons. Furthermore, the numbers of grains, and especially the evidence from the stone implement and its immediate surroundings, clearly reflect a well-developed technology for processing such foods. It should be stressed that various large flat stone objects were found in other brush huts, and they are currently under study.

The presence of oats raises an interesting question regarding their use as a staple food. Oat grains, like most cereals, are packed with starch and also contain an appreciable amount of protein. We tend to include oat in the cereals--the group of large-grained annual grasses which serve as the principal crops for most Old World civilisations. However, in the archaeobotanical record, oats are quite rare in prehistoric times. The only evidence for oat cultivation in south-west Asia comes from an unusual assemblage of some 12 000 Arena sterilis grains, found at the Neolithic site of Gilgal (Lower Jordan Valley, Israel), radiometrically dated to c. 11 700-10 550 cal BE These were found with c. 260 000 grains of wild barley, which together most probably represent pre-domestication cultivation of barley fields with an admixture of oats (Hartmann 2006; Weiss et al. 2006). Except for this rare find, we are unaware of any sign of oat cultivation or domestication in Neolithic or Bronze Age sites in south-west Asia and the Mediterranean basin (Zohary et al. 2012).

The rarity of domestic oat remains in Neolithic and later sites, where plant domestication occurred, requires explanation. Wild sterilis and fatua forms are widely distributed over many parts of the region. Moreover, wild oats (Avena sterilis), wild emmer (T. turgidum spp. dicoccoides) and wild barley (Hordeum spontaneum) grow together and form natural 'fields of wild cereals' (Zohary et al. 2012). Only small numbers of oat remains have been retrieved from Neolithic sites in the Fertile Crescent and Europe, and they appear to represent wild or weedy sterilis or fatua forms. Remains of non-shattering sativa or byzantina varieties with their characteristic plump seed--clear indications for their domesticated status--appear first in Europe, in the fourth and third millennia BP (Willerding 1970: 345-6; Villaret-von Rochow 1971). It is assumed, therefore (Zohary et al. 2012), that oat was first a weedy type that infested wheat and barley cultivation. Later, under agricultural regimes, it evolved to be a crop, picked up by the farmers to be yet another principal food.

The oat grains found at Ohalo II, A. barbara and A. sterilis, were of a wild type, which raises the question of dehusking. In both cases, the distribution unit includes one (A. barbara) or two (A. sterilis) fruits/grains, but not the glumes that remain on the plant. However, the paleas (the inner bracts of the spikelet, holding the grain) are thickly hairy and hold long kinked awns (Figure 7). To prepare the grains for grinding, these had to be removed. However, so far no such remains have been found in Ohalo II. For this reason, we assume dehusking took place somewhere outside the hut and clean grains were brought in for grinding. Unlike wheat and barley, dehusking of oat is a relatively simple procedure. We assume that singeing the spikelets and rubbing them in bare hands is all that is needed to separate the grains from the paleas.

[FIGURE 7 OMITTED]

At the time of our first analysis, the Ohalo II stone was the earliest implement shown to have been used directly to process plant materials. During the past five years, research carried out on even older Upper Paleolithic artefacts from Italy, Russia and the Czech Republic has demonstrated (mostly by starch grain analyses) that a variety of plants were ground with stone implements (Aranguren et al. 2007; Revedin et al. 2010). It should be pointed out, however, that in those earlier sites the stones were not regular, and usually not as large or flat as the Ohalo II specimen. The Ohalo and the European Upper Paleolithic cases demonstrate the value of starch analysis, as starch grains may routinely survive on stone artefacts from Palaeolithic periods, providing detailed information on previously little-understood patterns of plant exploitation.

Acknowledgements

The Ohalo II project was generously supported by the Irene-Levi Sala CARE Archaeological Foundation, the Israel Science Foundation (Nos. 831/00 and 711/08), the Jerusalem Center for Anthropological Studies, the L.S.B. Leakey Foundation, the Stekelis Museum of Prehistory in Haifa, the MAFCAF Foundation, the National Geographic Society and the Israel Antiquities Authority. We thank the Smithsonian National Museum of Natural History and the Smithsonian Tropical Research Institute for their support of the starch grain research. We also thank two reviewers for their useful advice. Figure 6 is reproduced by kind permission of Elsevier; Figure 7 is reproduced by kind permission of Oxford University Press.

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Received: 13 December 2011; Accepted." 16 March 2012; Revised: 10 April 2012

Dani Nadel (1), Dolores R. Piperno (2,3), Irene Holst (3), Ainit Snir (4) & Ehud Weiss (4)

(1) The Zinman Institute of Archaeology, The University of Haifa, 31905 Mt. Carmel, Israel (Email: dnadel@research.haifa.ac.il)

(2) Program in Human Ecology and Archaeobiology, Department of Anthropology, National Museum of Natural History, P.O. Box 37012 Smithsonian Institution, Washington, DC 20013-7012, USA (Email: pipernod@si.edu)

(3) Smithsonian Tropical Research Institute, Apartado Postal 0843-03092, Balboa, Republic of Panama (Email: holsti@si.edu)

(4) Institute of Archaeology, The Martin (Szusz) Department of Land of Israel Studies and Archaeology, Bar-Ilan University, 52900 Ramat-Gan, Israel (Email: snira@netvision.net.il; eweiss@biu.ac.il)

Table 1. Starch remains from the grinding stone, floor 11
(Brush Hut 1).

 Starch
Face Study remains Remarks

Upper First (2004) 150 See text
Upper Second (2011) 49 Similar composition of taxa
 (Hordeum; AHT group but no
 Triticum or Aegilops-types) as in
 the first analysis (see text)
Lower Second (2011) 13 One probable Hordeum grain (see
 text)

Table 2. Sediment samples from Brush Hut 1 analysed for starch.

Sample Square Height Floor Remarks

1 D81d 212.34-36 II ash
2 E79 212.40 II
3 E79c 212.42 II white ash
4 F80b 212.30 II white ash
5 D80d 212.45 III
6 E78d 212.50 III

Table 3. Dimensions of modern and ancient (Ohalo 11) Avena starch
grains.

 Mean
 length Range
Species Origin (micro]m) [micro]m)

Avena barbata ssp
 barbata Pott ex
 Link E. Weiss 2003 12 6-18
Avena barbata Pott NMNH2119068 9.5 6-16
Avena barbata Pott Kadesh stream, E. Weiss 8.7 4-12
Avena sterilis L. NMNH 1109491 7.7 4-12
Avena sterilis L. Mt. Hermon 1994, E. Weiss 8.6 4-14
Avena byzantina
 Koch Gatersleben 1.93, E. Weiss 9.1 4-14
Avena fatua L. Gatersleben 1.93, E. Weiss 6 4-10
Archaeological Ohalo 11 stone tool
 Avena grains (n = 11) 15.8 10-18