Preliminary investigation of the plant macro-remains from Dolni Vestonice II, and its implications for the role of plant foods in Palaeolithic and Mesolithic Europe.

Mason, Sarah L.R. ; Hather, Jon G. ; Hillman, Gordon C. 等

For the most part the Pleistocene, and even the earliest post-glacial, is a blank when it comes to evidence of humans eating plants. No wonder the old men's stories, of chaps who hunt great mammals and eat their meat, still dominate our unthinking visions of hunter-gathering in that period. Some real evidence, slight though it is, from a classic European Upper Palaeolithic site provides a more balance view.

Introduction

The role of plant foods in the diet of European hunter-gatherers during the Palaeolithic and Mesolithic is the subject of active debate, but little serious study. Despite analyses of plant remains from European Mesolithic sites (e.g. Price 1987; papers in van Zeist et al. 1991; Zvelebil in press) the results are generally sparse and restricted to very few taxa. For the Palaeolithic the evidence is still sparser, even if one includes the plant food remains recovered from Franchthi Cave, Greece (Hansen 1991) and Mezhirich in the Ukraine (Adovasio et al. 1992).

This paucity of data contrasts with evidence for the significant role of plant foods in early hunter-gatherer diet in other parts of the world with a similar vegetal resource base. In parts of North America, a range of wild plant foods have been recovered from sites of both Archaic hunter-gatherers (e.g. papers in Neusius 1986; and in Phillips & Brown 1985) and, increasingly, Palaeo-Indian hunter-gatherers (e.g. Frison 1991; Meltzer 1993). In the Near East rich assemblages of plant-food remains have been analysed from occupations of equivalent age to the European Upper Palaeolithic, and even earlier (Akazawa 1987; Garrard et al. 1988; Goren-Inbar et al. 1992; Hillman 1975; 1989; work in progress; Hillman et al. 1989a; 1989b; Kislev et al. 1992; McLaren in press; Potts et al. 1985; van Zeist & Bakker-Heeres 1984; van Zeist & Casparie 1968).

In addition to such direct evidence, heavy dependence on plant foods by pre-agrarian populations is also indicated by ethnoecological modelling (Hillman 1989; Hillman et al. 1989a; 1989b). A similar probability is implied by nutritional evidence suggesting that human energy requirements are unlikely to be fulfilled throughout the year by a largely meat-based diet, except in coastal and arctic habitats (Speth 1991; Speth & Spielmann 1983; notwithstanding the critique of Bunn & Ezzo 1993).

The gaps in current knowledge are, however, becoming more widely acknowledged, and van Andel (1990: 28) has suggested that determining the nature and use of the plant resource base for the Upper Palaeolithic is a 'major challenge', while Zvelebil (in press) says much the same of the European Mesolithic. There is clearly a need for empirical research from sites throughout Europe if the question of pre-agrarian plant use is to be properly addressed. This paper presents the results of one such study, and discusses some of the reasons for the paucity of data so far obtained.

Dolni Vestonice

Dolni Vestonice, in the Czech Republic, consists of two site clusters: Dolni Vestonice, I, excavated 1924-1979, and Dolni Vestonice, II, excavated in a rescue operation from 1985 to 1987. The sites form part of an Upper Palaeolithic Moravian group which appear to have been occupied during a warm phase or phases preceding the last glacial maximum. The Dolni Vestonice sites are located near the confluence of two rivers, on the lower slopes of the Pavlov Hills, an outcrop of Jurassic limestone rising c. 500 m above the surrounding lowlands, offering the occupants a variety of potential resource environments. Both site clusters consist of settlements with hearths and pits, and with large accumulations of mammoth bones near by, together with the bones of several other animal species (Klima 1963; Svoboda 1990; 1991; Vandiver et al. 1989). It has been suggested that Dolni Vestonice and the other Moravian sites share a number of features, including subsistence practices, with those of the Ukraine (including the site of Mezhirich) occupied roughly ten millennia later (Soffer 1985: 486).

The palaeoenvironment of Dolni Vestonice

Pollen analyses of samples from Dolni Vestonice II have been interpreted as indicating a forest-steppe environment (Svobodova 1991a; 1991b), with grasses (Poaceae/Gramineae), sedges (Cyperaceae), mugworts (Artemisia L.), and other members of the Asteraceae/Compositae family, together with chenopods (Chenopodiaceae), and members of the deadnettle (Lamiaceae/Labiatae) and buttercup (Ranunculaceae) families, and scattered junipers (Juniperus L.). There were also islands of conifers (pine -- Pinus L., spruce -- Picea A. Dietr., latch -- Larix Miller) and deciduous trees (oak -- Quercus L., lime -- Tilia L., hornbeam -- Carpinus L., beech -- Fagus L., hazel -- Corylus L., birch -- Betula L., willow -- Salix L., alder -- Alnus Miller), including both thermophilous elements and species which were probably concentrated in the river valleys. Wood charcoal analyses undertaken in the 1950s from Dolni Vestonice I have provided further detail on tree species utilized in the area, including several species of pines, fir (Abies Miller) and elm (Ulmus L.) (Klima 1963).

A similar suite of plant communities is suggested by the pollen and macroscopic remains in a core from nearby Bulhary (Rybnickova & Rybnicek 1991). This indicates that park-forest (coniferous trees with more sparsely-scattered deciduous species) formed a complex with grassland-steppe vegetation. Tall herbs occupied flood plains and other riverine habitats; and aquatic vegetation occupied pools, lakes and other areas of standing water. Sub-alpine communities are suggested to have covered the upper slopes and cliffs of the Pavlov Hills, and another community of herbaceous plants characteristic of springs and seepages has been identified. Altogether a wide range of plant resources would have been available to local people.

Climatic reconstructions based on these data, particularly noting the relatively high levels of arboreal pollen and several thermophilous species, imply a climate which was probably continental and relatively cool, but not characterized by extremely low temperatures, permafrost or tundra vegetation. Conditions seem to have been more temperate than in periods before and after occupation of the sites, and to have coincided with one of the warmer oscillations of the Wurmian interpleniglacial (i.e. an interstadial within early |O.sup.18~ stage 2). The environment is suggested to have been of high primary productivity, rich enough in plant and animal resources to support permanent or semi-permanent occupation (Rybnickova & Rybnicek 1991; Svobodova 1991a; 1991b).

Dolni Vestonice II -- analysis of charred plaint remains

Only one sample from Dolni Vestonice II was recovered for the examination of plant remains. The sample, comprising 280 ml of the deposit, was taken from Hearth D, containing charcoal with a radiocarbon determination of 26390|+ or -~270 b.p. (ISGS 1744).

The charred plant material was separated using small-scale bucket flotation, and the floating and suspendable fractions collected in a sieve with mesh size of 250 ||micro~meter~. The sunken residue, collected in a 1-mm sieve, has been retained but not yet examined. The dried riot was subsequently passed gently through a nest of sieves (2-mm, 1-mm, 500-||micro~meter~ and 250-||micro~meter~) to assist sorting under a low-power reflected-light microscope (magnification range from 6x to 50x). As few fragments larger than 2 mm were present, these were amalgamated with the 1-2-mm fraction, and this comprises the only fraction so far examined.

The sample can be divided into four categories of plant remains: wood charcoal, which constitutes the bulk; fragments of vegetative parenchyma (soft storage tissue) which forms the major part of organs such as roots and tubers (Hather 1991; 1993); seed remains; and a final disparate category of varied materials, most of plant origin, but morphologically indeterminate. The material in general is highly fragmented, but much is sufficiently well-preserved for cells to be clearly visible on newly-fractured surfaces.

Wood charcoal

Scanning electron microscopy (SEM) was used to identify wood charcoal, the manipulation of very small fragments of charcoal being difficult using epi-illuminated light microscopy. The three taxa so far identified -- Larix Mill. sp. (larch), Juniperus L. sp. (juniper) and Taxus baccata L. (yew) -- support the previously-cited palaeoenvironmental evidence that the locality of Dolni Vestonice was at least partially forested; yew and juniper are characteristic of limestone topography. Regarding climatic inferences it should be noted that although some Eurasian junipers and larches thrive in mesic or warm conditions, other species can survive in areas of extreme winter cold: L. dahurica Turcz., L. sibirica Ldb. and L. sukaczewii N.Dyl. can grow even on top of permafrost (Dylis 1947). Yew has not previously been recorded from either pollen or wood charcoal in the area. It is strongly hygrophilous, and is generally confined to relatively mesic conditions, with a more-or-less oceanic distribution which extends east as far as central Europe. It is claimed to be particularly intolerant of winter cold, and is not found in areas of extreme continental climate (Ellenberg 1974: 77; Godwin 1956: 28, 275; Srodon 1975), though Krol (1975) records it growing to 1450 m in the Carpathians, and even 1800 m in Macedonia. Its presence might be interpreted as implying a climate in the locality of Dolni Vestonice slightly milder and moister than suggested by previous reconstructions; or alternatively, it might indicate long-distance transport of wood to the site, possibly by human agency. Only analyses of a larger number of pollen or wood charcoal samples from the area can resolve this question.

Parenchymatous tissues -- possible evidence for use of root foods

Although, with experience, most parenchymatous fragments are usually distinguishable from wood charcoal at the sorting stage using low-power microscopy, the certain determination of some fragments as parenchyma, and, in almost all cases, the precise identification of this category of material, requires the use of SEM (Hather 1991; 1993). All the parenchyma appears to derive from fleshy taproots. Several fragments contain secretory cavities, a characteristic of the family Asteraceae/Compositae, which contains many species with edible roots, several characteristic of steppe environments. In interpreting these remains several possibilities must be considered. The charred fragments could derive from plant roots remaining in the ground when a firepit was first made, or from plants used as fuel, either directly or in sod-burning. However, in such cases a wider range of root and stem tissues should be preserved, including the tough inedible rhizomes of grasses, sedges and other plants. From the presence solely of fragments of fleshy root tissue it is possible to infer that these are most likely to be either remnants of food discarded on to the fire or material dropped accidentally during food preparation.

Seeds and fruits

The only intact specimen was a single, extremely distorted, seed. Features of the testa were too damaged to assist identification, and its internal tissues were fused, thus preventing the use of embryo and endosperm anatomy to refer it to family (Hillman & Mason, work in progress; Martin 1946).

Morphologically indeterminate fragments of plant origin

This category of plant material is very diverse. Generally SEM has proved essential for its examination, as many features are barely visible under low-power microscopy. Some material may be severely charred parenchyma, with all cell structures destroyed. Other fragments bear superficial similarity to remains from Epipalaeolithic Tell Abu Hureyra in Syria (Hillman et al. 1989a) and Late Palaeolithic Wadi Kubbaniya in Upper Egypt (Hillman 1989; Hillman et al. 1989b) which have been tentatively identified as the charred faeces of infants fed on finely-ground plant food. On close examination, some Dolni Vestonice fragments appear to be charred amalgamations of mineral matter and plant fragments, in one case recognizable as woody tissue. However, other fragments appear, like some of the Near Eastern specimens, to consist of a matrix of finely-comminuted plant material, sometimes with inclusions, of which one has a possible epidermal surface. Preliminary comparison has been made with an experimentally-charred plant-food 'mush' (of acorns), which under SEM examination consists of a more-or-less featureless matrix. Within the matrix are fragments of testa, together with small fragments of cotyledon, some of which have epidermal surfaces. The interpretation of Dolni Vestonice material as a plant-food mush is not inconsistent with this preliminary comparison. Although acorn mush was used for comparison -- and the pollen record suggests oaks in the region -- mushes can be made from many food plants, and from plant organs including fruits (and nuts), seeds, roots and tubers, or 'cambium' (inner bark) tissues. Determining whether any material is of faecal origin will require the use of chemical criteria (Hillman et al. 1993; Wales & Evans in press). Additionally, some less-textured fragments might derive from other organic substances containing occasional fragmentary plant inclusions; recent research by Wales (pers. comm.) has suggested that some of the Abu Hureyra specimens might be denatured bitumen.

Future work

Further examination of parenchyma from Dolni Vestonice may enable closer identification of the Compositae parenchyma. The 'comminuted plant material' is particularly challenging, as this has been previously reported only from the few Near Eastern sites cited. Even if source taxa are unidentifiable, a demonstration that processing of potential starch staple plant foods was undertaken, with its implications for technology and dietary strategies, would be of great significance at a site of such antiquity as Dolni Vestonice.

Plant foods: a missing element in European Palaeolithic/Mesolithic research

Our preliminary investigation demonstrates that evidence of the probable use of plant foods is recoverable even from sites of the antiquity of Dolni Vestonice. Evidence of this type can begin to redress the imbalance in existing perceptions of European pre-agrarian subsistence. Despite the long history of research into the European Palaeolithic and Mesolithic the role of plant foods in this period has been given surprisingly little consideration. Even for the Mesolithic, the lack of good evidence has ensured that Clarke's (1976) self-confessedly speculative paper, detailing the potentially dominant role of plant foods as staples providing carbohydrates, proteins and fats remains the most comprehensive and often-cited statement on the subject (e.g. Price 1989; papers in Zvelebil 1986; Zvelebil in press). The paucity of evidence from both Upper Palaeolithic and Mesolithic has allowed a view to persist, explicitly or implicitly expressed, that plant foods played an insignificant, or undetectable, role in subsistence.

There are several reasons for these residual misconceptions, and for the failure to invest in the systematic recovery and analysis of remains of plant foods from early sites in Europe.

1 While it is true that the relatively fragile remains of most food plants are, in general, less abundant on Palaeolithic and Mesolithic sites than on later ones, because of the age and nature of sites, a proportion of them do commonly survive. However, the pessimistic view that plant remains are preserved only under 'exceptional' conditions (e.g. Price 1989: 49; Prinz 1987: 77), or even that they are 'not preserved in archaeological contexts' (Clark 1987: 301) may have deterred attempts at recovery -- large-scale sampling and retrieval has rarely been applied systematically to Mesolithic and Palaeolithic European sites.

2 The meat-biased view of hunter-gatherer subsistence has been supported by the supposedly limited availability of edible plant resources in late Pleistocene environments (Bailey et al. 1983; Gamble 1986: 98-103; Jochim 1976; Mussi 1990: 141; Straus 1990: 101). This view derives in part from the lack of good information, and consequent debate, regarding the range of possible floristic environments of the Late Pleistocene period (e.g. Hopkins et al. 1982; Softer 1985: 149-52), but also from a poor understanding of the types of plant foods likely to have been utilized by hunter-gatherers. However, ecological studies of present-day vegetation, ethnobotanical studies of wild plant-use by recent hunter-gatherer and agrarian peoples, and archaeobotanical evidence have demonstrated the richness of superficially resource-poor environments such as the semi-arid steppe of southwest Asia (Hillman et al. 1989a; Hillman, Steele & Leeds, work in progress). Many of the food plants concerned (or very near relatives) would also have been available to Palaeolithic and Mesolithic populations in contiguous areas of south and central Europe, and some in Mesolithic northern Europe as well. It is probable that many of these played a similar role in European diet.

3 The focus in studying plants in pro-Neolithic subsistence (e.g. papers in Zvelebil 1986; Zvelebil in press), remains the period of transition to agriculture, and this has detracted from a proper recognition of the fact that the basis of subsistence of most pre-agrarian peoples was probably provided by that majority of wild plant foods which were never eventually 'husbanded', cultivated or domesticated (Mason 1992: 46-8, 197-8, 203-4).

4 Archaeobotanists working on plant remains from early sites have often failed to recognize the remains of certain plant foods, some of which probably served as starch staples. Parenchymatous tissues derived either from vegetative plant parts (e.g. fleshy roots, tubers, stems, etc.) or from non-vegetative organs (storage tissues within seeds and fruits) can superficially be confused with poorly preserved wood charcoal, as can fragments of processed plant tissues. It is likely that such remains occur fairly frequently in archaeobotanical assemblages, but are usually dismissed as unidentifiable. These oversights are exacerbated by the fact that archaeobotanical assemblages are rarely analysed in a holistic way, which is essential to obtain the maximum amount of information on past plant foods (Hather in press). It is important to incorporate data from wood charcoal, especially from older sites, and others where the palaeoenvironment is poorly understood. When interpreted with care (allowing for the possibility of long-distance transport, and taking into account taphonomic processes, including human selection) this can provide useful information on probable arboreal dominants, and thence on the local resource-base. Additionally, the fruits, inner bark, leaves and shoots of trees can themselves be a direct source of food. The need to examine whole plant assemblages with regard to potential information on food plants is clear.

5 Identifications of food plants are traditionally based on gross external morphology, and are only appropriate for seeds and some fruits surviving relatively intact and originating from regions where the local flora is well known. Such methods suffice for well-preserved remains of cereals and their weeds, but are inadequate for other plant remains which might occur in both pre-agrarian and agrarian contexts, e.g. roots and tubers, leaf and stem foods, and 'cambium' or inner bark foods, as well as fragmentary seeds and fruits with external surfaces missing. The use of anatomical techniques, and frequently of SEM, such as used in the current study are essential for the identification of these categories of remains, and in some cases the methodologies and criteria required for identification require fundamental research.

Conclusions

This analysis of plant food remains from one small hearth sample at Dolni Vestonice hints at the untapped potential that exists for recovering information on plant foods from European Palaeolithic and Mesolithic sites, provided suitable sampling, recovery and identification techniques are applied. Once applied to a broad range of sites, the consequences of such investigations of direct archaeobotanical evidence for the use of plants as food may force a fundamental revision of our understanding of the subsistence strategies and diet of European hunter-gatherers during the late Pleistocene and early Holocene.

Acknowledgements. The authors would like to acknowledge the help of Olga Soffer, University of Illinois at Urbana-Champaign, USA, and Jiri Svoboda, AU CAV, Czech Republic, who provided the plant remains and associated information. Research by JGH is funded by the Science and Engineering Research Council.

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