Palaeoenvironmental reconstruction and flora exploitation at the Palaeolithic cave of Theopetra,...

ORIGINAL PAPER

Palaeoenvironmental reconstruction and flora exploitationat the Palaeolithic cave of Theopetra, central Greece:the evidence from phytolith analysis

Georgia Tsartsidou & Panagiotis Karkanas &Gilbert Marshall & Nina Kyparissi-Apostolika

Received: 7 October 2013 /Accepted: 18 February 2014 /Published online: 13 March 2014# Springer-Verlag Berlin Heidelberg 2014

Abstract This paper presents the results of phytolith analysiscarried out on sediments from Theopetra Cave in Thessaly,Central Greece. Theopetra is one of the most important latePleistocene sites in the region, with occupation spanning theMiddle Palaeolithic to the end of the Neolithic. The aim of thisstudy is to contribute to our understanding of the nature ofhuman occupation in the cave during the Palaeolithic andMesolithic. Palaeoenvironmental issues are also addressed inorder to understand the climate and vegetation around the caveduring that time. Twelve layers of anthropogenic and geogenicorigin which mark distinct occupation episodes have beensampled. The anthropogenic layers consist of combustionfeatures and are valuable indicators of human activity withinthe cave, providing information on the types of vegetationcollected for everyday activities and consumption. Thegeogenic sediments are mostly of natural origin and markintervals during which the site was mostly unoccupied. Theyprovide evidence for the climate and plant communities grow-ing around the cave. The results point to intensive occupationof the cave during the transition from the penultimate glacialto the last interglacial, a period of mild climate, high

precipitation and rich vegetation in the catchment area.Sporadic use of the cave is implied during the last glacial,followed by more frequent visits towards the end of thePleistocene. A range of plants were used for fuel, food andother day-to-day activities. Theopetra is discussed in compar-ison with Klissoura, a key Palaeolithic cave site in southernGreece. A number of conclusions are drawn concerning life atthe two sites and their surroundings, based on similarities anddifferences in phytoliths and other key environmental anddietary indicators.

Keywords Phytoliths . Palaeolithic . Mesolithic . Greece .

Cave . Combustion features

Introduction

The full complexity of Palaeolithic andMesolithic subsistencestrategies in Greece is not well understood. Most of the betterinvestigated sites are located in the regions of Epirus and thePeloponnese (Harvati et al. 2009). Moreover, with the excep-tion of the Mesolithic site of Maroulas (Sampson 2011),studies focusing on macro-remains (charcoal and seeds) havemostly been carried out in caves and shelters: Lakonis,Cyclope, Franchthi, Klisoura, Theopetra, Schisto and Boila(Hansen 1991; Litynska-Zajac 2010; Kotzamani 2009;Mueller-Bieniek 2010; Ntinou 2010). Phytolith studies havebeen extensively used as a tool for understanding the use ofplants in antiquity, although the bulk of these have focused onthe Neolithic, Bronze and Iron Ages (Albert et al. 2008;Horrocks et al. 2003; Madella 2001; Madella et al. 2009;Rosen 2001; Roberts and Rosen 2009; Ryan 2011; Shahack-Gross et al. 2005; Shahack-Gross and Finkelstein 2008;Shillito 2011; Tsartsidou et al. 2009). Only a few have con-centrated on the Palaeolithic and Mesolithic (Albert et al.1999, 2000, 2003, 2006; Cabanes et al. 2010; Henry et al.

Electronic supplementary material The online version of this article(doi:10.1007/s12520-014-0183-6) contains supplementary material,which is available to authorized users.

G. Tsartsidou (*) : P. Karkanas :N. Kyparissi-ApostolikaEphoreia of Palaeoanthropology–Speleology of Southern Greece,Ardittou 34b, 11636 Athens, Greecee-mail: [email protected]

P. Karkanase-mail: [email protected]

N. Kyparissi-Apostolikae-mail: [email protected]

G. MarshallThe Wiener Laboratory, ASCSA, Souidias 52-54, Athens, Greecee-mail: [email protected]

Archaeol Anthropol Sci (2015) 7:169–185DOI 10.1007/s12520-014-0183-6

http://dx.doi.org/10.1007/s12520-014-0183-6

2004; Karkanas et al. 2002; Madella et al. 2002; Rosen 2003;Schiegl et al. 2004; Zurro et al. 2009). In Greece, there hasonly been one study targeting the Palaeolithic specificallyAlbert’s (2010) report on the material from Klissoura Cave 1in the northeastern Peloponnese.

Phytoliths have the potential to provide information on theplants brought into a site by humans, as well as the structure oflocal palaeovegetation (Piperno 2006). As opposed to char-coal and seeds, they need not be charred in order to bepreserved (Harvey and Fuller 2004; Piperno 2006; Weiner2010). They also provide information about the specific partsof the plants in use, which is more difficult or impossible withcharcoal and seed studies. Moreover, they provide evidencefor plants that were growing in the vicinity of the cave asopposed to pollen which gives a geographically much broaderview. Phytoliths along with the rest of the archaeobotanicaldataset, including charcoal, pollen and seeds, can be used toinvestigate human choice and to reconstruct thepalaeoenvironment.

The present study focuses on the sediments from thePalaeolithic and Mesolithic occupation of Theopetra Cave.Phytolith analysis is used to investigate the types of plants inuse as well as their relative frequency within the sediments.The study focuses on samples recovered from anthropogeniccombustion features and those from the natural geogenicsediments of the cave. Combustion features reflect the specificcircumstances of firing events, and their plant assemblagescan provide clues as to why the fire was set, the season ofoccupation and the types of vegetation being collected.Following on from this, human choice and action are consid-ered, as reflected in the types of plants brought into the cavefor fuel and other activities. The results also provide a pictureof the types of plants growing in the vicinity of the site and cantherefore be used to investigate the local palaeoenvironment.For a more representative evaluation of plants growing aroundthe cave, phytoliths from the geogenic sediments are analysed.These are the result of natural geological processes with littleanthropogenic input (Karkanas 2001). They are waterborneand contain the remains of plants growing close to the site.Some may contain eroded material from the underlying com-bustion features (see taphonomical implications), but most aremade up of plant material from the humus horizon of the soiloutside the cave. The small size of the hill in which the cave islocated, along with shallow and rapid karst drainage, meansthat the remains of plants in the geogenic layers will not havebeen derived from any great distance and have not been storedin the karstic system for long time.

The study is complemented by the results from otherarchaeobotanical studies, including wood charcoal (Ntinouand Kyparissi-Apostolika 2008) and seeds (Kotzamani2009). Pollen analysis is still underway, and there are unfor-tunately no other studies available from the area aroundTheopetra. Therefore, the results from three other regions

(Fig. 1), Xinias lake, the Kopais Basin in central Greece, andthe Ioannina basin in northwestern Greece to the west ofPindus mountain chain, are used (Bottema 1979; Frogleyet al. 1999; Tzedakis 1999). Given that the oldest 14C datefrom Xinias lake is 40 kyr cal BP (Bottema 1979), only unitIV from Theopetra has been compared.

The site of Theopetra

Theopetra Cave is located 5 km south of the town ofKalambaka in northern Thessaly (Fig. 1). The mouth of thecave faces north overlooking the Peneios River where it exitsfrom the Pindus mountain chain and flows eastwards acrossthe Thessalian plain. The cave has an occupational area ofapproximately 500 m2 and was excavated by Nina Kyparissi-Apostolika of the Ephoreia of Palaeoanthropology andSpeleology of southern Greece (Kyparissi-Apostolika 1999,2006). The excavations produced a stratigraphic sequence of6.4 m in depth, documenting a long occupational history fromat least 130,000 years ago until recent times (Valladas et al.2007). The site has been intensively studied, providing asubstantial body of information regarding stratigraphy, min-erals, micromorphology, lithics, bones, human skeletons, ce-ramics and archaeobotanical remains (Adam 1999; Karkanas1999, 2001; Karkanas et al. 1999; Kotzamani 2009; Ntinouand Kyparissi-Apostolika 2008; Panagopoulou 1999;Stravopodi et al. 1999).

The stratigraphy is divided into six major units (Fig. 2),from unit I at the bottom to unit VI at the top (Karkanas 2001).Unit II has been further subdivided into 12 layers, described asanthropogenic and geogenic. The anthropogenic layers (II2,II4, II6, II11 and unit IV) consist of laterally continuoussuperimposed combustion zones. These are often made upof lenses of white, black and orange burnt materials, as well asbrown sub-layers. Combustion sequences such as these arecreated during repeated burning episodes within hearth fea-tures (Meignen 2007). The hearths are flat, not constructed inany way and represent mostly undisturbed and intact ashaccumulations produced during several burning episodes.The high degree of calcination of the ash components andtheir grey to white colour point to deliberately set fires in linewith reporting of similar features at other sites (Meignen2007). The geogenic layers (II1, II3, II5, II7–II10 and II12)are made up mostly of waterborne deposits and range fromstratified silty loam to sandy sediments. Theopetra is well-known for post-depositional chemical alteration of sediments,a process related to local hydrology and the accumulation oflarge quantities of guano during the Palaeolithic (Karkanaset al. 1999; Karkanas 2001). Diagenesis has resulted in thedissolution of bones in the central and rear parts of the cave,leading to the formation of authigenic phosphate minerals.Significantly though, the phytoliths have not been affectedby these processes (Karkanas et al. 1999, 2000).

170 Archaeol Anthropol Sci (2015) 7:169–185

Most of the archaeological layers were formed during theMiddle and Upper Palaeolithic, the Mesolithic and Neolithic(Kyparissi-Apostolika 2006). The largest part of the sequenceat Theopetra is attributed to the Middle Palaeolithic (layersII1–II11) and is therefore important to our understanding ofthis period in Greece. Hearths from layers II2 and II4 havebeen thermoluminescence (TL) dated to the transition fromMIS6 toMIS5, the penultimate glacial to last interglacial (124±16 and 129±13 kyr BP, respectively) (Valladas et al. 2007).The lithic industry from these layers is dominated by use ofthe Levallois technique (Panagopoulou 1999). Hearths inlayer II11 have been TL dated (57±6 kyr BP) to the lastglacial (Valladas et al. 2007). This layer contains some spo-radic lithic material, mostly non-diagnostic although generally

of Palaeolithic type (Valladas et al. 2007). Hearth layers datedfrom 20 to 12 kyr cal BP have been assigned to the UpperPalaeolithic. An ashy layer correlated laterally with the upperpart of II12 has been 14C dated to 19 kyr cal BP (Karkanas2001), whilst the lower part of unit IV has been dated to 15 kyrcal BP. The end of the Upper Palaeolithic and the Pleistocenehas been dated to around 13 kyr cal BP from the middle part ofunit IV. The lithic material from the Upper Palaeolithic fallsunder the broad heading of Epigravettian (Adam 1999). Theupper part of unit IV is attributed to the Mesolithic and hasbeen dated from approximately 11.2–9.2 kyr cal BP(Facorellis et al. 2001; Karkanas 2001 and unpublisheddata). The lithic material from the Mesolithic is similar to thatfrom phase VII at Franchthi cave, with a lack of backed

Fig. 1 Map of Greece showing the location of Theopetra Cave as well as the other sites mentioned in the text

Archaeol Anthropol Sci (2015) 7:169–185 171

bladelets, geometric microliths or evidence for the microburintechnique. Instead, the assemblage is dominated by flakedebitage and a broad range of flake-based retouched tools(Adam 1999). Unit IV is preserved along the eastern andwestern walls as well as in front of the cave, but is not ascomplete as those from the Middle Palaeolithic. Combustionfeatures of unit IV consist mostly of dark black sediments.These are likely to represent the charcoal-rich base of hearthsthat have been preserved, with the white and grey upper ashpartly eroded or dissolved. The Neolithic layers are mostlyreworked and lack intact combustion features and were there-fore not included in this analysis.

Materials and methods

Sediment samples were collected after cleaning of thesections, from anthropogenic (II2, II4, II6 and II11) andgeogenic (II1, II3, II5, II7, II8, II9, II10 and II12) layersfrom unit II, as well as anthropogenic parts of unit IV(Online Supplementary Table 1; Fig. 2). Combustionlayers were sampled in each stratigraphic sequence ac-cording to depth, to determine phytolith compositionthrough time and in relation to different types of ashsediment, upper white, middle grey and lower black.The brown sediments between each of the individual

Fig. 2 The stratigraphy of thecave


hearth lenses were sampled separately. These probablyrepresent a mixture of geogenic and anthropogenic burntmaterials. The combustion sequences were also sampled hor-izontally at several points along their length, as well as withinthe same stratigraphic layer in different profiles.

The samples from the geogenic sediments were col-lected in two ways: from different points (vertically withdepth recorded) within the layer and then as a verticalcomposite sample from top to bottom of each layer.Since geogenic processes remain broadly constant duringthe formation of each layer, the second approach provid-ed a more representative picture of the layer as a whole.However, both approaches were used when possible.Five control samples were collected from outside of thecave in order to compare the modern vegetation profilewith those from the archaeological deposits within thecave. The first sample was collected from the upperhumus layer (A-horizon) outside of the entrance, and afurther four from trenches dug at 300 m and 1 km away.Two samples were collected from each trench: one from4 to 5 cm below the surface (A-horizon) and one from 20 to30 cm. In total, 102 samples were collected, including thosefrom within the cave and the control samples from outside(see Online Supplementary Table 1).

The analytical methods used follow those described inTsartsidou et al. (2008). The numbers of phytoliths wererelated to 1 g of sediment as well as 1 g of acid-insolublefraction (AIF), in order to allow comparison with othercaves as shown in Albert et al. (1999). The percentage ofeach phytolith type (or taxon) was calculated as a relativefrequency of the total with consistent morphology persample. When possible, a minimum of 200 phytolithswere counted per slide, though in most cases more than400 with consistent morphology were counted per sample.Samples with less than 50 phytoliths with consistent mor-phology were not included in the morphological analysisbecause of large statistical error. All assemblages weremorphologically classified following the InternationalCode for Phytolith Nomenclature (Madella et al. 2005),when possible.

A rich comparative collection of phytoliths of modernplants from Greece (Tsartsidou et al. 2007) was used toevaluate the sediment assemblages. This comparativecollection was expanded by the inclusion of two archae-ological samples of macrobotanical remains from thecave, seeds from juniper (Juniperus sp.) and gromwell(Lithospermum arvense). The aim was to explore thepossibility of identifying characteristic phytoliths fromthese two species, the most common macrobotanicalfinds from the cave. It should be noted that these seedswere already carbonized as they had been collected fromcombustion layers, and therefore, the number ofphytoliths was related to 1 g of ash.

Results

Taphonomic implications

The geogenic sediments formed as follows: Material fromoutside of the cave was transported in by water via the karstsystem and deposited on top of the underlying combustionfeatures. During this process, it is possible that some materialfrom the underlying combustion features may have beenreworked and thus included within the overlying geogeniclayers. Phytoliths in the geogenic sediments may thereforehave two origins, from within reworked combustion featuresand/or plant material from outside of the cave that wastransported in by natural geogenic processes. Comparisonbetween combustion layers and the overlying geogenic sedi-ments may help to resolve this issue. Hearths as opposed togeogenic sediments are in situ features with secure anthropo-genic context. But if geogenic and hearth sediments containsimilar phytolith assemblages, it would suggest that materialfrom the hearths had been reworked and incorporated withinthe natural geogenic sediments during their formation.Alternatively, if the hearths and geogenic layers were signif-icantly different, it would point to their natural origin fromoutside of the cave. If this is the case, then the geogenic layerswould constitute reliable indicators of the vegetation from thesurrounding area.

Geogenic layer II1 varied markedly from combustion layerII2, quantitatively and qualitatively. Quantitatively it producedfar fewer phytoliths, whilst qualitatively it contained morewood phytoliths and much lower frequencies of grass huskphytoliths. It also produced minor quantities of dicot leafphytoliths and almost no sedge phytoliths, in contrast to layerII2. Comparing geogenic layer II3 with the combustion layersII2 and II4, it was poorer and the two did not share manyqualitative similarities (Figs. 5, 8 and 9). Geogenic layer II5shared many similarities with combustion layer II4 apart fromdicot fruit phytoliths. A further difference was the muchhigher silica skeleton/single phytolith ratio. This suggests thatit does not contain much reworked material although thepossibility cannot be ruled out for certain. Geogenic layersII7–II10 were very poor and did not share any similarities withthe combustion layers. Conversely although quantitativelymuch poorer, geogenic layer II12 was qualitatively very sim-ilar to combustion layer II11. The silica skeleton/single phy-tolith ratio, dicot leaves and especially the wood phytolithfrequencies were very similar. On the other hand,Cyperaceae and dicot fruit phytoliths were present at muchlower frequencies than in II11. This suggests that II12 doesinclude some ash from the underlying combustion layers ofII11, along with waterborne geogenic sediments.

It is worth noting that silica skeleton/single phytolith ratiosare in general higher in the combustion sediments than thegeogenic ones. The lower ratios amongst the geogenic layers


could be explained taphonomically. Geogenic sedimentsmade up of material from outside of the cave, or erodedhearths, would be expected to contain mostly single and lightphytoliths that are more easily transported as opposed toarticulated ones. This is due to part of the silica skeletonhaving been removed through abrasion. As has been shown,these types of structures are fragile and sensitive to mechan-ical post depositional processes (Cabanes et al. 2009, 2010;Jenkins 2009; Shillito 2011). However, since there were noconsistent differences between the geogenic and anthropogen-ic layers in terms of the silica skeleton/single phytolith ratio(for example, geogenic layer II5 had a higher ratio thancombustion layer II6), we can use it as an environmentalindicator, but only supplementary to other proxies.

The phytolith assemblages from within the geogenic sedi-ments in the cave are a good reflection of those in thePalaeolithic soils, as is the modern soil surface from thecatchment of Theopetra. Control sample TH98 was from themodern humus layer, and the results were consistent with themodern environment (arboreal vegetation and wet climate).The rest of the control samples contained very few phytoliths.In open environments, pedogenic processes lead to poor phy-tolith preservation, compared to those in archaeological envi-ronments (Cabanes et al. 2011). In effect, phytoliths from thecomparatively protected environment of caves have a muchbetter chance of survival than those from more openenvironments.

Plant phytolith assemblages

Of the two seed taxa analysed, Juniperus sp. produced nocharacteristic phytoliths and therefore could not be identifiedin the sediments. On the other hand, L. arvense seeds pro-duced almost 900,000 phytoliths/g ash, with characteristicspiny and bulbous shape and morphology (Fig. 3a). As such,they are of taxonomic importance.

Sediment phytolith assemblages

The results of the analysis of the cave sediments and controlsamples from outside are listed in Online SupplementaryTable 1. The numbers of phytoliths are expressed per gramsediment as well as per gram AIF.

The control samples collected from the area outside of thecave (TH98–TH102) produced extremely low phytolith con-centrations, ranging from just a few to 12,000/g sediment. Therichest (TH98) was collected from the upper humus layer (A-horizon) close to the cave. The number of phytoliths droppedsignificantly in the samples collected from the deeper hori-zons. Since the four control samples (TH99, TH100, TH101and TH102), as well as those from the brown geogenic sedi-ments between the ashy combustion lenses (TH1, TH2, TH4,TH6 and TH15), produced less than 50 phytoliths with

consistent morphology per sample, they were not includedin the morphological charts. The four control samples werecompared only quantitatively with the cave sediments, whilstonly TH98 was compared both quantitatively and qualitative-ly. The low frequency of phytoliths in these samples high-lights their lack of anthropogenic input.

The cave sediments produced high phytolith concentra-tions, pointing to significant anthropogenic input of plantmaterial. The combustion layers were significantly richer thanthe geogenic ones, which were also much more variable. Thegeogenic sediments mostly contain phytoliths transported intothe cave by water. It is also possible that during the movementof this water, sediments already within the cave could bereworked resulting in the enrichment of the geogenic sedi-ments compared to the control samples, although in terms ofphytolith frequencies they would still be significantly poorerthan the combustion layers.

Geogenic layer II7 and most of the samples from layers II8and II9 produced only a few thousand phytoliths per gramsediment, as opposed to geogenic layers II1 and II3 withmillions. Combustion layers also varied markedly in termsof phytolith quantities, whilst there was no apparent

Fig. 3 Images of phytolith silica skeletons of: a Lithospermum arvenseseeds and b Celtis sp. fruits


correlation between the colour of the lenses and phytolithfrequency (Fig. 4). Both ashy (white and grey) and charcoalrich (black) lenses produced millions of phytoliths.Conversely, intermediate brown sediments preserved less than50,000 phytoliths per gram sediment. The richest combustionlayer was II2, with between 1 and 43 million phytoliths/gsediment. Those in layer II4 produced between 600,000 and4 million, whilst II11 produced between 500,000 and 5million/g sediment. Lower numbers were found in layer II6with up to 800,000 phytoliths/g sediment.

Figure 5 shows the ratio of multi-celled silica skeletons tosingle-celled phytoliths. As has been shown (Mithen et al.2008; Rosen and Weiner 1994), elevated silica skeleton num-bers reflect high levels of water. We therefore use high ratiovalues as a proxy for water availability. Lithospermumphytoliths cannot be used as an environmental indicator, dueto mineralization of its seeds by schlerenchymatisation(Pustovoytov et al. 2004; Roberts and Rosen 2009). LayersII2, II4, II5 and IV had the highest ratios (average 0.12–0.18).Layers II3, II11 and II12 were lower (average 0.06–0.1),whilst the remainder (II1, II6, II7, II8, II9 and II10) were allsignificantly lower than 0.05. Since farming was unknown atthe time, high precipitation is the most likely explanation forthe high ratios. This may suggest higher rainfall during theformation of layers II2–II5 and IVand possibly II11, with lessduring II1 and II7–II10. Moreover, cone and long-cell (rods)phytoliths which are produced by wetland plants (the formerby Cyperaceae and the latter by both Cyperaceae andJuncaceae) were found in the same sediments, mostly layersII2–II4 and IV (Fig. 6), with the highest frequencies (up to7 %) in II2. They were virtually absent in layers II7–II11. It isworth noting that silica skeletons characteristic of reeds werenot recovered in any of the sediments.

In terms of morphology and typology, phytoliths that couldbe assigned to wood (blocky morphotypes and those of vari-able morphology) were detected in all layers both anthropo-genic and geogenic (Fig. 7). Layers with higher frequenciesincluded II11 and II12 with 51 and 42 %, respectively. Wood

was the main fuel although the percentage of wood phytolithsidentified in some of the combustion lenses of II2 and II6 wasrelatively low. It has been shown (Albert 2000; Tsartsidouet al. 2007) that wood does not produce large amounts ofphytoliths and there are species with none at all. Interestingly,the geogenic layers at Theopetra produced relatively highfrequencies of wood phytoliths, probably due to dispersal ofash within the cave.

Grasses were the most common taxa (86 %) identified inthe majority of sediment samples, with Festucoids the pre-dominant subfamily. These are the tall wild grasses character-istic of the Mediterranean that thrive in cool temperate envi-ronments. Chloridoid, Arundinoid and Panicoid grasses wereidentified as well although at lower frequency (average 3–4 %). Stems and leaves were the most predominant elementidentified amongst the grasses. Husk phytoliths were presentat low frequency (between 1 and 5 %) in the geogenic sam-ples, lower than the control sample (TH98) collected fromoutside the cave, which produced 7 %. Conversely, the com-bustion layers ranged from 1 to 16% (Fig. 8), with the highestfrequencies (3–16 %) in layer II2, in which 15 out of the 19samples produced significantly more than 7 % huskphytoliths. Moreover, II2 was the only feature in which stemand leaf phytoliths were present at lower frequency thanhusks.

Figure 9 shows the frequencies of phytoliths (polyhedrals,sclereids, tracheary, dicot hairs) produced by the leaves ofdicotyledonous plants (trees and shrubs) and those (jigsawpuzzle phytoliths) that thrive mostly in deciduous forest envi-ronments. Combustion layers II2, II4 and IV, as well asgeogenic II5 and II12, produced larger quantities of phytolithsfrom the leaves of dicot plants. These ranged from 1 to 25 %(average 4.5–6.5 %) in contrast to the rest of the geogenicsamples (II1, II3, II7–II10) as well as the combustion layer inII6 which were all less than 3%.Moreover, the control sampleproduced less than 4 %. These results suggest the use of leafybranches, pointing to forest vegetation within the catchment ofthe site. Similar results were obtained from combustion layers

Fig. 4 Phytolith concentrations in the combustion layers of the cave. The different components of the combustion lenses are indicated as follows:Whitecolumns are white ash lenses, grey columns are grey ash lenses, black columns are charcoal rich lenses and dotted columns are the orange-coloured lenses


II2, II4 and IV, suggesting a comparable climate. ComparingFigs. 9 and 5, particularly in terms of average values, we seesimilar high frequencies in all three. In the same way, Fig. 6highlights the high frequency of wetland plants in these layers.

Combustion lenses from layer II11 produced extremelyhigh frequencies of phytoliths (up to 4.5 million/g sediment),whereas the brown geogenic sediments around the peripherycontained only a few thousand (3,600–4,000). Some of thelenses from II11 shared morphological similarities with thosefrom layers II2 and II4, including relatively high frequenciesof dicot leaf phytoliths (up to 9 %) and Lithospermum seeds(up to 60 %).

Phytoliths from seeds of fruits from dicotyledonous plantswere found in the majority of the cave layers. Two taxa wereidentified, L. arvense and Celtis sp. (Fig. 10a, b). Most of thelayers preserved less than 10 % phytoliths of dicot fruit seeds.The highest proportions were in combustion lenses of II4 (up

to 47 %), II11 (up to 60 %) and unit IV (up to 80 %). Thesebelongmostly to L. arvense. Celtis sp. was identified in almostall of the samples, and the frequencies in most did not exceed6%. The highest frequencies were in the upper parts of II4 andII6, the lower part of II11 and in most layers of unit IV. It isworth noting that none of the two taxa was recorded in thecontrol samples, pointing to the anthropogenic origin of theseeds from the cave.

Discussion

The phytolith assemblages from the Theopetra combustionfeatures provide evidence for high levels of anthropogenicinput. A variety of plants and parts thereof were selected forfuel, with wood an important component. Even waterbornesediments reworked by geogenic processes were relatively

Fig. 5 Ratio of multi celled silicaskeletons to single-celledphytoliths. aAll cave samples andthe control TH98 (C) areincluded, b average for thesamples per layer. Grey columnsare anthropogenic layers whilstblack columns are geogenic

Fig. 6 Phytoliths from sedgesand rushes (cones and rods)identified in the sedimentassemblages. Grey columns areanthropogenic layers whilst blackcolumns are geogenic. C is thecontrol sample (TH98)


rich in wood phytoliths, most probably a result of dispersal ofash throughout the site due to racking out, dumping andtrampling. Herbaceous plants as well as grasses were identi-fied in high quantities. Many of these may have been used askindling whilst others may have been thrown into the fireaccidentally or discarded intentionally after use.

Environmental issues

As shown by the taphonomic analysis, most of the geogeniclayers apart from II12 were not particularly affected by thehearths. They may have been slightly enriched, but theirphytolith assemblages reflect the environment rather thancontamination from the hearths. As such, they serve as anatural environment proxy and together with the anthropo-genic sediment phytoliths can be used to reconstructpalaeovegetation and palaeoclimate.

There is a large volume of papers that deal with pastenvironmental and especially vegetation conditions.Alexandre et al. (1997) first proposed a formalized versionof the ratio of arboreal to grass phytoliths during their work intropical Africa. Known as the Dicot/Poaceae (d/p) ratio, it isused to differentiate forested environments from open grass-land. Other researchers have similarly found that phytolithscan be used to distinguish between forested and savannahenvironments (e.g. Aleman et al. 2012; Alexandre et al.

1999; Barboni et al. 2007; Bremond et al. 2005, 2008).However, the use of the d/p ratio does not work for all areas(Strömberg 2004). Care must be taken in the northernMediterranean where the most common taxon, Quercus sp.,does not produce the spherical phytoliths needed for its cal-culation (Bremond et al. 2004). This is true for the Theopetraregion where as shown by anthracological analysis as well aspollen data,Quercus sp. was the predominant taxon during thelate Pleistocene. This may be the reason why sphericalphytoliths are almost completely absent from the Theopetraphytolith assemblages.

Within the Theopetra phytolith assemblages, the most ap-parent difference was between wet and forested environmentscompared to dryer and more open conditions (Table 1). Thisconclusion is based on the abundance of phytoliths fromdicotyledonous plants, the silica skeleton/single phytolith ratioand the presence of wetland species such as sedges and rushes.In parallel, charcoal and pollen data are discussed in order toidentify similarities or differences.

In beginning the discussion of the lower stratigraphic layers(II1–II4), the following is a brief summary of the currentlyavailable dating evidence. The TL dates from layers II2 andII4 are 124±16 and 129±13 kyr BP, respectively (Valladaset al. 2007). New tephrochronology results place the top oflayer II4 at the very beginning of the last interglacial(Karkanas et al., submitted for publication). Given that the

Fig. 7 Wood phytoliths (blockymorphotypes and variablemorphology phytoliths) identifiedin the sediment assemblages.Grey columns are anthropogeniclayers whilst black columns aregeogenic. C is the control sample(TH98)

Fig. 8 Grass husk and stem/leaf phytoliths identified in the sediment assemblages. Layers 2, 4, 6, 11 and IVare the anthropogenic layers.C is the controlsample (TH98)


two TL dates of II2 and II4 are statistically similar, they placethe formation of the hearths of these layers at broadly the sametime, at around the end of the penultimate glacial and

beginning of the last interglacial. Stratigraphically this wouldsuggest that underlying layer II1 was also formed before theend of the penultimate glacial. The results of the phytolith

Fig. 9 Phytoliths of dicot leaves(polyhedrals, sclereids, trachearyelements, dicot hairs and jigsawpuzzle phytoliths) identified in thesediment assemblages. a All thecave samples and the controlTH98 (C), b average for thesamples per layer. Grey columnsare anthropogenic layers whilstblack columns are geogenic

Fig. 10 Concentrations of phytoliths of dicot fruits identified in the sediment assemblages. a Lithospermum arvense, b Celtis sp. Grey columns areanthropogenic layers whilst black columns are geogenic


analysis from this bottom layer point to a cold climate withsparse vegetation. Conversely, the results from layers II2, II4,II5 and possibly II6 suggest milder conditions with higherprecipitation, as indicated by abundant remains of dicotyle-donous trees and shrubs which thrive in wet-forested environ-ments and C3 grasses. The presence of sedges also points toan increase in river flow due to higher precipitation. Togetherthis evidence suggests a shift from cooler conditions in layerII1, to milder ones in II2 and II4. The phytoliths from theselower layers correspond with the TL results, which place theformation of II2 and II4 around the end of MIS6 and thebeginning of MIS5e, respectively, with layer II1 earlier duringMIS6.

Similarly, analysis of wood charcoal suggested sparse veg-etation in layer II1, with a shift towards pioneer species andgallery forests in layer II2, some temperate taxa in layers II3and II4 and then temperate woodland in layers II5 and II6.High frequencies of thermophilous taxa such as Quercus sp.,Fraxinus sp. and Carpinus/Ostrya have been identified inthese layers (Ntinou and Kyparissi-Apostolika 2008).Similar results have been observed in pollen sequences fromGreece and elsewhere for the last interstadial of the transitionto the last interglacial (Frogley et al. 1999; Roucoux et al.

2011; Sánchez-Goñi et al. 1999; Seidenkrantz et al. 1996;Tzedakis et al. 2003). More specifically in pollen diagramsof the MIS6 to MIS5e transition in the Ioannina basin, anumber of climatic oscillations are observed with a shortstadial following a pronounced interstadial (Frogley et al.1999; Roucoux et al. 2011; Tzedakis et al. 2003). At thebeginning of the sequence (130 kyr), there was an increasein precipitation and expansion of deciduous Quercus sp.(Frogley et al. 1999). This may coincide with layer II2 atTheopetra. The small shift towards drier conditions and moreopen vegetation prior to 128,000 years ago may correspondwith layer II3, within which phytoliths suggest a cooler inter-val with a significant decrease in dicotyledonous trees andlower silica skeleton/single phytolith ratios. This supports theobservation by Karkanas (2001) that layer II3 was frost af-fected in contrast to layers II1 and II4–II6. The pollen evi-dence from the Kopais basin (Tzedakis 1999) also supportsthe results of the phytolith analysis. Last interglacial pollen,which in our study includes layers II2–II6, contained higharboreal pollen percentages and concentrations dominated byoak (Quercus sp.). Values of evergreenQuercus sp. along withPistacia sp.,Phillyrea sp. and Fraxinus sp. were higher duringthe earlier part of the interglacial whilstQuercus sp. and Pinus

Table 1 The correlation of phytoliths indicative of warm and wet conditions as well as the vegetation and climate inferred per layer and climatic period

Layer Phytoliths Vegetation, climate Climatic period

IV upper High frequencies of dicot leaves and riverineplants, and high silica skeleton/singlephytolith ratio

Arboreal vegetation andincreased humidity

Holocene

IV middle Relatively high frequencies of dicot leaves andriverine plants, and relatively high silicaskeleton/single phytolith ratio

Fluctuating arboreal vegetation Tardiglacial

IV lower Very low frequencies of dicot leaves and sporadicriverine plants, and relatively low silicaskeletons/single phytoliths ratio

Sparse arboreal vegetationand low humidity

Glacial

II12 High frequencies of dicot leaves and riverineplants, and high silica skeleton/single phytolithratio (may be intrusive)


Interstadial

II11 Moderate frequencies of dicot leaves and very lowfrequencies of riverine plants, along withmoderate silica skeleton/single phytolith ratio

Fluctuating arboreal vegetationand moderate humidity

Interstadial

II9–II10 Very low frequencies of dicot leaves and riverineplants, and very low silica skeleton/singlephytolith ratio

Sparse arboreal vegetation anddecreased humidity

Glacial

II7–II8 Very low frequencies of dicot leaves and an absenceof riverine plants, along with very low silicaskeleton/single phytolith ratio

Sparse arboreal vegetation anddecreased humidity

Glacial

II4–II6 High frequencies of dicot leaves and riverine plants,and high silica skeleton/single phytolith ratio


Last Interglacial

II3 Moderate frequencies of dicot leaves and riverineplants, and moderate silica skeleton/singlephytolith ratio

Reduced arboreal vegetation Penultimate Glacial/LastGlacial Transition

II2 High frequencies of dicot leaves and riverine plants,and high silica skeleton/single phytolith ratio


Penultimate Glacial/LastGlacial Transition

II1 Low frequencies of dicot leaves and riverine plants,and moderate silica skeleton/single phytolith ratio

Sparse arboreal vegetation andlow humidity

Penultimate Glacial


sp. were abundant during the latter, along with the presence ofAbies sp. and Betula sp.

Different vegetation structures and inferred climatic condi-tions are indicated for layers II7–II10. The phytolith datasuggest that the last interglacial ended somewhere within layerII7. From this point onwards, lower frequencies of dicotleaves imply a more open landscape with sporadic broad leaftrees, probably tethered to water sources such as rivers orlakes. Low frequencies of sedges in II9 with a completeabsence in layers II7, II8 and II10 may suggest a decrease inprecipitation and harsher climatic conditions. However, a shiftaway from human use of sedges cannot be excluded. With adate from overlying layer II11 of 57±6 kyr BP (Valladas et al.2007), it is likely that layers II7–II10 were formed during thecold and arid conditions of the last glacial and in particularMIS4. Due to the lack of dates from this part of the sequence,we cannot be sure if the latter part of MIS5 is also included.Evidence of frost activity in these layers (Karkanas 2001)further reinforces the phytolith data, whilst the results ofcharcoal analysis also suggest harsh climatic conditions fromlayer II7 until II10. Open parkland vegetation is indicated bythe abundance of Juniperus sp. and Prunus sp., whilst treetaxa in gallery forests are suggested by Salix–Populus sp.(Ntinou and Kyparissi-Apostolika 2008). Analysis ofJuniperus sp., Salix sp. and Populus sp. by Tsartsidou et al.(2007) showed that they produce no more than a few hundredphytoliths per gram wood and just a few thousand per grambark. This explains why they are not a major component in thephytolith assemblages. The leaves of Populus sp. producehigher amounts of phytoliths (1.4 million/g dry plant) asopposed to Salix sp. and Juniperus sp. that produce no morethan 20,000 and 200,000 phytoliths/g dry plant, respectively(Tsartsidou et al. 2007). Populus sp. may thus be a significantcomponent of the leaf assemblages found in layers II7–II10.Phytoliths from combustion layer II11 suggest mild condi-tions, a small interstadial during the last glacial, possibly at thebeginning of MIS3. Anthracological analysis identified smallnumbers of temperate taxa in these hearths, includingQuercussp., Tilia sp. and Ulmus sp. (Ntinou and Kyparissi-Apostolika2008).

Pollen evidence from the Kopais basin (Tzedakis 1999)shows that the last glacial consisted of alternating warm andcold episodes, with shifts from more open conditions withgrasses, sagebrush and chenopods, to closed forest.Combustion episodes from unit IV which correspond to thetransition from the Pleistocene to Holocene repeat more or lessthe same pattern as that identified in layers II2 and II4, withoscillations between wetter-forested conditions and coolerintervals with more open woodland.

We should again discuss the available dates before theenvironmental proxies of unit IV. The first sample in thesequence (TH93) derived from a combustion episode datedto approximately 18,000 cal BP (unpublished data). Cold and

harsh conditions of the glacial maximum are indicated by lowfrequencies of dicot trees, the lack of sedges and the low silicaskeleton/single phytolith ratio. The final five samples of thesequence (TH94–97, TH50) are all dated to around 10,000 calBP, and the phytoliths in most point to vegetation rich indeciduous trees and bushes, suggesting humid and temperateconditions with high river flows towards the end of thePleistocene. Lower values in some of the samples could berelated to taphonomic issues. The 11 intermediate samples(TH43–49, TH82–83 and TH91–92) are dated to between15 and 13kyr cal BP (Karkanas 2001; Facorellis et al. 2001;unpublished data). Oscillation in the amount of tree coversuggests alternating cold and warm intervals. Comparing thedicot leaf frequencies with phytolith concentrations (seeOnline Supplementary Table 1), there was no apparent corre-lation between high frequencies of arboreal vegetation andrich phytolith assemblages which are indicative of more in-tense occupation.

Anthracological analysis has shown that most of the layersof unit IV contain evidence for a vegetation succession fromopen grassland with pioneer species and gallery forests of thelast glacial, to open woodland with temperate and thermoph-ilous taxa such as Quercus sp. and Fraxinus sp. (Ntinou andKyparissi-Apostolika 2008). The Xinias pollen diagrams alsopoint towards oscillations with harsher conditions up to 15 kyrcal BP years ago, after which they ameliorated. Towards theend of Pleistocene and beginning of the Holocene (around10 kyr cal BP), high tree pollen values with Quercus sp.,Fraxinus sp. and Pistacia sp. were recorded. Similarly, theKopais pollen data from the end of the Pleistocene and earlyHolocene suggest high arboreal percentages with Quercus sp.well represented along with Pistacia sp. and Juniperus sp.

Subsistence strategies

The exploitation of a wide variety of plants as identified in thephytolith assemblages, including trees, herbs, fruits, wildgrasses and sedges, suggests a broad diet and implies a varietyof activities undertaken in the cave. Phytoliths show thatgrasses of all types were being brought into the cave, includ-ing Festucoids which must have been present in abundance inthe vicinity (cool climate with tall Festucoid grasses) andChloridoids, Panicoids and Arundinoids which were collectedin small quantities. Dry grasses are excellent kindling materi-al, whilst their seeds are in many cases edible. Those fromPoaceae are calorie dense (Henry et al. 2011) and may havebeen included in the diet. The abundance of grass seeds, as hasbeen indicated by grass husk phytoliths, during the formationof combustion layer II2 in the Middle Palaeolithic, probablypoints to the collection of specific grasses for consumption.Given the hardness of wild grass seeds (Piperno et al. 2004), itis likely they would have needed to be processed prior to use.Although there is no evidence of any artefacts specialized for


use as grinding implements or storage features within thecave, wild grass consumption cannot be ruled out. The dryingof seeds for easier dehusking might have taken place by thefire, into which the empty husks would have been discarded.Other studies have suggested that wild grass seeds wereincluded as part of the Palaeolithic diet. Madella et al.(2002) have identified grass seeds as a component of the dietduring the Middle Palaeolithic at Amud cave in Israel. Henryet al. (2011) have reported a variety of plant foods includinggrass seeds (Triticeae) in the form of phytoliths and starchgrains recovered from dental calculus of Neanderthal skele-tons from Shanidar cave in Iraq and Spy cave in Belgium.Seed assemblages and starch remains from the UpperPalaeolithic site of Ohallo II suggest extensive collection ofseeds and the consumption of smaller species (Piperno et al.2004; Weiss et al. 2004). Finally, the phytolith data fromKlissoura Cave 1 also point to grass seed consumption(Albert 2010).

Cyperaceae plants identified in the cave may also havebeen consumed. Sedges’ rhizomes are a valuable source ofstarch and protein, whilst their presence points to exploitationof nearby wetlands. Sedges as well as grass stems and leavesmay also suggest fibre working on site for matting and bed-ding. The latter has been identified in a small number ofMiddle Palaeolithic sites. At Tor Faraj in Jordan, there was aclear spatial arrangement with dicot plants in the hearth areasand monocotyledonous phytoliths found on the periphery,interpreted as some sort of bedding (Henry et al. 2004;Rosen 2003). Nadel et al. (2004) also report evidence for grassbedding at Ohallo II. Although the Theopetra assemblages donot provide clear evidence for bedding, the high concentra-tions of stem/leaf grass phytoliths point to the use of at leastsome of these types of plants. Reeds were not identified in thesediments from Theopetra. Considering that reeds producehigh quantities of characteristic phytoliths, their absence prob-ably points to human preference for sedges and grasses.

The large quantities of seeds from dicotyledonous plants(Celtis sp. and L. arvense) identified during phytolith analysissuggest an emphasis on the collection of dicot fruits andespecially Celtis sp. (Fig. 3b) during layer II4, as well as mildintervals during the last glacial (rare in combustion lensesfrom layer II11), and L. arvense (Fig. 3a) towards the end ofthe Pleistocene (Unit IV). Celtis sp. is a widely grown ediblefruit. Its seeds have been found in ash deposits fromChoukoutien Cave in China (Brothwell and Brothwell1969), Middle Palaeolithic deposits at Douara cave in Syria(Matsutani 1987) and at Franchthi Cave in southern Greece(Hansen 1991). The large quantities of gromwell (L. arvense)seeds identified in the Upper Palaeolithic layers at Franchthi(Hansen 1991) were thought to be part of the sediment prior todeposition and therefore not the result of human activity.However, this was not the case at Theopetra, since the seedswere recovered from the ashy layers of the hearths.

Nevertheless, its use is puzzling since it is not obviouslyedible. It is an annual herb that thrives on hills, pastures,steppic rocky mountain slopes and the margins of cultivatedground (Pustovoytov et al. 2004). The Japanese eat the leavesof L. arvense as a boiled salad (Tanaka 1976 in Hansen 1991),and there are reports of the pharmaceutical use of the plant inAsiatic countries. In antiquity, species of Lithospermum havebeen identified at several sites, suggesting a range of possibleuses by the inhabitants of Theopetra. Over 600 perforatedseeds of Lithospermum officinale were recovered from theGulmenitsa Culture burial site in Bulgaria, which have beeninterpreted as beads (Renfrew 1973). Perforated seeds ofLithospermum were also identified at Neolithic sites else-where in Europe and interpreted as items of jewelry(Schlichtherle 1988). Gromwell seeds were used duringChinese antiquity for decoration on tubs (Jiang et al. 2007).In Poland, they were used for their supposed medicinal valuefor more than 3,000 years (Baczyńska and Lityńska-Zając2005). The roots of gromwell produce a red dye that is alsoconsidered to have therapeutic value. This red colour mayhave also been used during the Palaeolithic and Mesolithic.Matsutani (1987) argues for the use of Boraginaceae plants fordye at the Palaeolithic site of Douara cave in Syria, with roots,seeds, leaves and stems, together with the seeds of Celtis sp.after consumption of the edible fruit, used for fuel. It isinteresting to note the continuous presence of gromwell atTheopetra from the Middle Palaeolithic to the Mesolithic.Accepting that they were intentionally brought in, this couldsuggest that knowledge of its attributes and uses lasted for aconsiderable period of time.

The great variety of plants brought into Theopetra was alsosuggested by the macrobotanical remains (Kotzamani 2009),with many different types discovered in the MiddlePalaeolithic sediments. These included the seeds ofJuniperus sp. and wild legumes (Vicia sp., Lathyrus sp.,Lens sp.), as well as those of wild dicot and monocot plants(Ficus sp., Rubus sp., Vitis sp., Hippophaes sp., Celtis sp.,Polygonum sp., Lolium sp., Eragrostis sp., Elleocharis sp.,L. arvense). Sediments from the end of the Upper Palaeolithicwere poor in terms of macrobotanical remains, compared tothe Mesolithic which was rich in seeds from species such asCeltis sp., Juniperus sp. and L. arvense.

In terms of diet through time, we notice some interestingdifferences at Theopetra. As macro- and micro-remains show,combustion layer II2 preserved the highest frequencies of wildgrass seeds although a forested environment was suggested bythe rest of the phytolith data. This may reflect deliberatechoice with a diet based also on wild grass consumptionduring the end of the penultimate glacial. During the forma-tion of layer II4, the diet was enriched with Celtis fruits. Theuse of these continued towards the end of Pleistocene incombustion layer IV. Wild grass seeds were also present atthis time but to a lesser extent.


Use of Theopetra Cave

The cave was intensively used during warm climatic periodsas indicated by high frequencies of phytoliths in combustionlayers II2 and II4. Unit IV also suggests frequent use of thecave around 15 and 13 kyr cal BP and then sporadicallyaround 10 kyr cal BP, with arboreal vegetation andwet climateindicated by the phytolith assemblages. There was significant-ly less use of the cave earlier during the glacial as indicated bythe lower overall frequencies of phytoliths per gram sediment,probably to do with the deteriorating conditions as indicatedby the rapid decline in phytoliths indicative of warm and wetconditions. One possible exception is combustion layer II11which points to occupation during a warmer climatic period(interstadial) of the last glacial. Cultural remains includinglithics and bones also point to sporadic use during the lastglacial (Panagopoulou 1999; Valladas et al. 2007). One obvi-ous explanation may be that the north facing wet- and frost-affected cave (Karkanas 2001) was unattractive during coldclimatic periods. It is possible that areas closer to the coastwere preferred. Large quantities of grass seeds point to habi-tation at least during the flowering seasons, spring, summerand early autumn, though storage and use during wintercannot be excluded, with the cave perhaps occupied all yearround.

Comparison with Klissoura cave

Klissoura Cave I in the northern Peloponnese preserves a richcultural sequence dated from the Middle Palaeolithic toMesolithic (Koumouzelis et al. 2001; Stiner et al. 2010). Themost current phytolith results only relate to the UpperPalaeolithic and Mesolithic layers (Albert 2010), and thus,comparison with Theopetra will be limited to unit IV which isattributed to the final Pleistocene.

Comparing the sediments of Theopetra with those fromKlissoura highlighted two main differences. The Klissouracombustion feature phytolith assemblages were poorer thanthose from the geogenic sediments. In contrast, the Theopetracombustion layers were much richer than the surroundinggeogenic sediments. The quantities per gram AIF recoveredfrom the combustion features of Klissoura ranged from10,000 to 240,000 phytoliths, whereas at Theopetra theyranged from 73,000 to 3 million. These results point eitherto significantly more anthropogenic input and more intenseuse of the hearths at Theopetra, or the use of different plants asfuel as discussed below.

There are also some important morphological differencesin the phytolith assemblages from the two caves. The UpperPalaeolithic and Mesolithic combustion features of Klissourashow high wood ash content, along with significant quantitiesof herbaceous plant and dicotyledonous leaf phytoliths (Albert2010). Conversely, those from unit IVat Theopetra preserved

fewer wood phytoliths, but larger quantities of those fromdicot leaves. The relatively lower overall frequencies ofphytoliths at Klissoura compared to Theopetra may be relatedto differences in the relative proportions of hard woods, leavesand other materials used for fuel at the two sites. Hard woodscontain comparatively few phytoliths, and their use wouldtherefore result in a small assemblage. Conversely, other typesof potential fuel including leaves are richer in phytoliths,although in terms of heat output they are relatively poor. Theabundance of dicot leaf phytoliths at Theopetra may suggestdeliberate human choice of these materials for fuel, but per-haps also different environmental conditions, in particulargreater aridity in the catchment area of Klissoura. The highfrequency of dicot fruits may also suggest that different sub-sistence practices were carried out by the inhabitants of thetwo caves.

In terms of grasses, C3 types prevailed at both sites. Therewere differences though in the frequencies of C4 grasses,which at Klissoura have only been recovered in theEpipalaeolithic layers dated to between 15 and 14 kyr calBP. Albert (2010) has suggested that the presence of C4Chloridoid grasses in these layers points to drastic changesin the environment and climate, with the expansion of opensteppic landscapes and the disappearance of most trees. Theanthracological study from Klissoura supports this interpreta-tion, with the identification of Prunus amygdalus/spinosa,which grows in relatively arid and more open environments(Ntinou 2010). Theopetra shows a different picture, with C4grasses present in almost all the layers but at low frequency(up to 4 %). Unit IV showed no abrupt change, and there wascontinuous evidence for broad leaf plants, suggesting noincrease in aridity.

Common to both caves was the presence of inflorescencephytoliths, suggesting collection of wild grasses for consump-tion during the flowering period. If both caves were occupiedduring the warmer parts of the year, then a general trendregarding the use of caves during the Palaeolithic andMesolithic could be suggested. This possibility needs to beexplored further.

Conclusions

The analysis of phytoliths from the sediments at Theopetraallows us to infer climatic conditions and vegetation structureduring the last 140,000 years at the site. The results point tointensive use of the cave during the Middle Palaeolithic and inparticular towards the end of the penultimate glacial andduring the last interglacial. The cave was also used duringmilder intervals of the last glacial and towards the end of thePleistocene. More sporadic use was indicated for cooler pe-riods during the last glacial. In terms of human subsistenceand day to day activities, a broad variety of plants were


exploited for food, as fuel and probably fibre working for themanufacture of matting and bedding, along perhaps withdecoration and possibly for their medicinal properties. A dietrich in dicot fruits and possibly seeds of wild grasses may havebeen adopted. The cave was in use during warmer periods ofthe year, which could perhaps indicate a preference towardsthe exploitation of resources frommountainous areas betweenspring and autumn, with the coast possibly in use during thewinter.

Acknowledgments This study was carried out during GT’s Fellowshipin Environmental Studies in 2010 at the Wiener Laboratory of theAmerican School of Classical Studies at Athens. She is grateful to Dr.Sherry Fox, the director of the Wiener Laboratory, for her support. Wealso thank the two anonymous reviewers for the detailed and constructivecriticisms of this manuscript.

References

Adam E (1999) Preliminary presentation of the Upper Paleolithic andMesolithic stone industries of Theopetra Cave, western Thessaly. In:Bailey GN, Adam E, Panagopoulou E, Perles C, Zachos K (eds) Thepalaeolithic archaeology of Greece and adjacent areas, proceedingsof the ICOPAG conference, Ioannina. British School at AthensStudies 3. British School at Athens, Athens, pp 266–270

Albert RM (2000) Study of the ash layers through phytolith analyses fromthe Middle Palaeolithic levels of Kebara and Tabun Caves.Dissertation, University of Barcelona

Albert RM (2010) Hearths and plant uses during the Upper Palaeolithicperiod at Klissoura cave 1 (Greece). The results from phytolithanalysis. Eurasian Prehistory 7(2):71–85

Albert RM, Lavi O, Estroff L, Weiner S (1999) Mode of occupation ofTabun cave, Mt Carmel, Israel during theMousterian period: a studyof the sediments and phytoliths. J Archaeol Sci 26:1249–1260

Albert RM, Weiner S, Bar-Yosef O, Meignen L (2000) Phytoliths in theMiddle Palaeolithic deposits of Kebara cave, Mt Carmel, Israel:study of the plant materials used for fuel and other purposes. JArchaeol Sci 27:931–947

Albert RM, Bar-Yosef O, Meignen L, Weiner S (2003) Quantitativephytolith study of hearths from the Natufian and MiddlePalaeolithic levels of Hayonim cave (Galilee, Israel). J ArchaeolSci 30:461–480

Albert RM, Bamford MK, Cabanes D (2006) Taphonomy of phytolithsand macroplants in different soils from Olduvai gorge (Tanzania)and the application to Plio-Pliostocene palaeoanthropological sam-ples. Quat Int 148:78–94

Albert RM, Shahack-Gross R, Cabanes D, Gilboa A, Lev-Yadun S,Portillo M, Sharon I, Boaretto E, Weiner S (2008) Phytolith-richlayers from the Late Bronze and Iron Ages at Tel Dor (Israel): modeof formation and archaeological significance. J Archaeol Sci 35:57–75

Aleman J, Leys B, Apema R, Bentaleb I, Dubois MA, Lamba B,Lebamba J, Mart in C, Ngomanda A, Truc L (2012)Reconstructing savannah tree cover from pollen, phytoliths andstable carbon isotopes. J Veg Sci 23:187–197

Alexandre A, Meunier JD, Lezine AM, Vincens A, Schwartz D (1997)Phytoliths: indicators of grassland dynamics during the lateHolocene in intertropical Africa. Pal Pal Pal 136:213–229

Alexandre A, Meunier JD, Mariotti A, Soubies F (1999) Late Holocenephytolith and carbon-isotope record from a latosol at Salitre, south-central Brazil. Quat Res 51:187–194

Baczyńska B, Lityńska-Zając M (2005) Application of Lithospermumofficinale L. in early Bronze Agemedicine. VegHist Archaeobot 14:77–80

Barboni D, Bremond L, Bonnefille R (2007) Comparative study ofmodern phytolith assemblages from inter-tropical Africa. Pal PalPal 246:454–470

Bottema S (1979) Pollen analytical investigations in Thessaly (Greece).Palaeohistoria 21:19–40

Bremond L, Alexandre A, Villa E, Guiot J (2004) Advantages anddisadvantages of phytolith analysis for the reconstruction ofMediterranean vegetation: an assessment based on modern phyto-lith, pollen and botanical data (Luberon, France). Rev PalaeobotPalynol 129:213–228

Bremond L, Alexandre A, Hély C, Guiot J (2005) A phytolith index as aproxy of tree cover density in tropical areas: calibration with LeafArea Index along a forest–savannah transect in southeasternCameroon. Glob Planet Chang 45:277–293

Bremond L, Alexandre A, Wooller MJ, Hély C, Williamson D, SchäferPA, Majule A, Guiot J (2008) Phytolith indices as proxies of grasssubfamilies on East African tropical mountains. Glob Planet Chang61:209–224

Brothwell D, Brothwell P (1969) Food in antiquity. Thames and Hudson,London

Cabanes D, Burjachs F, Expósito I, Rodríguez A, Allué E, Euba I, VergésJM (2009) Formation processes through archaeobotanical remains:the case of the Bronze Age levels in El Mirador cave, Sierra deAttapuerca Spain. Quat Int 193:160–173

Cabanes D, Mallol C, Expósito I, Baena J (2010) Phytolith evidence forhearths and beds in the late Mousterian occupations of Esquilleucave (Cantabria, Spain). J Archaeol Sci 37:2947–2957

Cabanes D, Weiner S, Shahack-Gross R (2011) Stability of phytoliths inthe archaeological record: a dissolution study of modern and fossilphytoliths. J Archaeol Sci 38:2480–2490

Facorellis Y, Kyparissi-Apostolika N, Maniatis Y (2001) The cave ofTheopetra, Kalambaka: radiocarbon evidence for 50,000 years ofhuman presence. Radiocarbon 43:1029–1048

Frogley MR, Tzedakis PC, Heaton THE (1999) Climate variability innorthwest Greece during the Last Interglacial. Science 285:1886–1889

Hansen J (1991) The Palaeoethnobotany of Franchthi cave. Excavationsat Franchthi cave, Greece, Fascicle 7. Indiana University Press,Bloomington

Harvati K, Panagopoulou E, Runnels C (2009) The palaeoanthropologyof Greece. Evol Anthropol 18:131–143

Harvey EL, Fuller DQ (2004) Investigating crop processing using phy-tolith analysis: the example of rice and millets. J Archaeol Sci 32:739–752

Henry DO, Hietala HJ, Rosen AM, Demidenko YE, Usik VI, ArmaganTL (2004) Human behavioural organization in the MiddlePalaeolithic: were Neanderthals different? Am Anthropol 106:17–31

Henry AG, Brooks AS, Piperno DR (2011) Microfossils in calculusdemonstrate consumption of plants and cooked foods inNeanderthal diets (Shanidar III, Iraq; Spy I and II, Belgium).PNAS 108(2):486–491

Horrocks M, Irwin GJ, McGlone MS, Nichol SL, Williams LJ (2003)Pollen, phytoliths and diatoms in prehistoric coprolites fromKohika, Bay of Plenty, New Zealand. J Archaeol Sci 30:13–20

Jenkins E (2009) Phytolith taphonomy: a comparison of dry ashing andacid extraction on the breakdown of conjoined phytoliths formed inTriticum durum. J Archaeol Sci 36:2402–2407

Jiang H, Li X, Liu CJ, Wang YF, Li CS (2007) Fruits of Lithospermumofficinale L. (Boraginaceae) used as an early plant decoration(2500 years BP) in Xinjiang, China. J Archaeol Sci 34:167–170

Karkanas P (1999) Lithostratigraphy and micromorphology of TheopetraCave deposits, Thessaly, Greece: some preliminary results. In:Bailey GN, Adam E, Panagopoulou E, Perles C, Zachos K (eds)


The palaeolithic archaeology of Greece and adjacent areas, proceed-ings of the ICOPAG conference, Ioannina. British School at AthensStudies 3. British School at Athens, Athens, pp 240–251

Karkanas P (2001) Site formation processes in Theopetra Cave: a recordof climatic change during the Late Pleistocene and Early Holocenein site formation processes in Theopetra Cave. Geoarchaeology16(4):373–400

Karkanas P, Kyparissi-Apostolika N, Bar-Yosef O, Weiner S (1999)Mineral assemblages in Theopetra, Greece: a framework for under-standing diagenesis in a prehistoric cave. J Archaeol Sci 26:1171–1180

Karkanas P, Bar-Yosef O, Goldberg P, Weiner S (2000) Diagenesis inprehistoric caves: the use of minerals that form in situ to assess thecompleteness of the archaeological record. J Archaeol Sci 27:915–929

Karkanas P, Rigaud JP, Simek JF, Albert RM,Weiner S (2002) Ash bonesand guano: a study of the minerals and phytoliths in the sediments ofGrotte XVI, Dordogne, France. J Archaeol Sci 29:721–732

Kotzamani G (2009) From gathering to cultivation: archaeobotanicalresearch on the early plant exploitation and the beginning of agri-culture in Greece (Theopetra Schisto, Sidari, Dervenia).Dissertation, University of Thessaloniki, Greece (in Greek)

Koumouzelis M, Ginter B, Kozlowski K, Pawlikowski M, Bar-Yosef O,Albert RM, Litynska-Zajac M, Stworzewicz E, Wojtal P, Lipecki G,Tomek T, Bochenski ZM, Pazdur A (2001) The early UpperPalaeolithic in Greece: the excavations in Klisoura cave. JArchaeol Sci 28:515–539

Kyparissi-Apostolika N (1999) The palaeolithic deposits of TheopetraCave in Thessaly (Greece). In: Bailey GN, Adam E, PanagopoulouE, Perles C, Zachos K (eds) The palaeolithic archaeology of Greeceand adjacent areas, proceedings of the ICOPAG conference,Ioannina. British School at Athens Studies 3. British School atAthens, Athens, pp 232–239

Kyparissi-Apostolika N (2006) Twelve years of excavation and research,proceedings of the international conference (Trikala, 6–7 November1998), 2nd edn. INSTAP and Ministry of Culture, Athens

Litynska-Zajac M (2010) Plant material from Klissoura cave 1 in Greece.Eurasian Prehistory 7(2):87–90

Madella M (2001) Understanding archaeological structures by means ofphytolith analysis: a test from the Iron Age site Kilise Tepe-Turkey.In: Meunier JD, Colin F (eds) Phytoliths applications in earthscience and human history. Balkema, Lisse, pp 173–182

Madella M, Jones MK, Goldberg P, Goren Y, Hovers E (2002) The exploi-tation of plant resources by Neanderthals in Amud cave (Israel): theevidence from phytolith studies. J Archaeol Sci 29:703–719

Madella M, Alexandre A, Ball T (2005) International code for phytolithnomenclature. Ann Bot 96:253–260

Madella M, Jones MK, Echlin P, Powers-Jones A, Moore M (2009) Plantwater availability and analytical microscopy of phytoliths: implica-tions for ancient irrigation in arid zones. Quat Int 193:32–40

Matsutani A (1987) Plant remains from the 1984 excavations at Douaracave. In: Akazawa T, Sakaguchi Y (eds) Paleolithic site of Douaracave and paleogeography of Palmyra basin in Syria. Part IV: 1984excavations, bulletin no. 29. The University Museum, University ofTokyo Press, Tokyo, pp 117–122

Meignen L (2007) The hearths at Kebara cave and their role in siteformation processes. In: Bar-Yosef O,Meignen L (eds) Kebara cave,Mt. Carmel, Israel. The Middle and Upper Palaeolithic archaeology,part I, American School of Prehistoric Research Bulletin 49.Peabody Museum of Archaeology and Ethnology, HarvardUniversity, Cambridge, pp 91–122

Mithen S, Jenkins E, Jamjoum K, Nuimat S, Nortcliff S, Finlayson B(2008) Experimental crop growing in Jordan to develop methodol-ogy for the identification of ancient crop irrigation. World Archaeol40(1):7–25

Mueller-Bieniek A (2010) Archaeobotany in the Mesolithic settlement ofMaroulas/Kythnos. In: SampsonA, KaczanowskaM, Koslowski JK

(eds) The prehistory of the island of Kythnos (Cyclades, Greece) andthe Mesolithic settlement at Maroulas. The Polish Academy of Artsand Sciences—The University of the Aegean, Krakow, pp 141–142

Nadel D, Weiss E, Smchoni O, Tsatskin A, Danin A, Kislev M (2004)Stone age hut in israel yields world's oldest evidence of bedding.PNAS 101(17):6821–6826

NtinouM (2010)Wood charcoal analysis at Klissoura Cave 1 (Prosymna,Peloponnese): the Upper Palaeolithic vegetation. EurasianPrehistory 7(2):47–69

Ntinou M, Kyparissi-Apostolika N (2008) The Pleistocene–Holocenecharcoal record from Theopetra Cave, Thessaly, Greece: implica-tions for vegetation, climate and human use. In: Damblon F, Court-Picon M (eds) Programme and abstracts, 4th International Meetingof Anthracology, Brussels, 8–13 September 2008 (p 105). RoyalBelgian Institute of NATURAL Sciences, Brussels. ISSN: 0378-0902

Panagopoulou E (1999) The Theopetra Middle Palaeolithic assemblages:their relevance to the Middle Palaeolithic of Greece and adjacentareas. In: Bailey GN, Adam E, Panagopoulou E, Perles C, Zachos K(eds) The palaeolithic archaeology of Greece and adjacent areas.Proceedings of the ICOPAG conference, Ioannina. British School atAthens Studies 3. British School at Athens, Athens, pp 252–265

Piperno DR (2006) Phytoliths: a comprehensive guide for archaeologistsand paleoecologists. Altamira, Lanham

Piperno DR, Weiss E, Holst I, Nadel D (2004) Processing of wild cerealgrains in the upper Palaeolithic revealed by starch grain analysis.Nature 430:670–673

Pustovoytov KE, Riehl S, Mittmann S (2004) Radiocarbon age of carbon-ate in fruits of Lithospermum from the early Bronze Age settlementof Hirbet ez-Zeraqon (Jordan). Veg Hist Archaeobot 13:207–212

Renfrew JM (1973) Palaeoethnobotany. The prehistoric food plants of thenear east and Europe. Methuen, London

Roberts N, Rosen A (2009) Diversity and complexity in early farmingcommunities of Southwest Asia: new insights into the economic andenvironmental basis of Neolithic Çatalhöyük. Curr Anthropol 50(3):393–402

Rosen AM (2001) Phytolith evidence for agro-pastoral economies in theScythian period of southern Kazakhstan. In: Meunier JD, Colin F(eds) Phytoliths applications in earth science and human history.Balkema, Lisse, pp 183–198

Rosen AM (2003) Middle Palaeolithic plant exploitation: themicrobotanical evidence. In: Henry DO (ed) Neanderthals in theLevant. Behavioral organization and the beginnings of human mo-dernity. Continuum, London, pp 156–171

Rosen AM, Weiner S (1994) Identifying ancient irrigation: a new methodusing opaline phytoliths from emmerwheat. J Archaeol Sci 21:125–132

Roucoux KH, Tzedakis PC, Lawson IT, Margari V (2011) Vegetationhistory of the Penultimate Glacial period (Marine isotope stage 6) atIoannina, north-west Greece. J Quat Sci 26(6):616–626

Ryan P (2011) Plants as material culture in the Near Eastern Neolithic:perspectives from the silica skeleton artifactual remains atÇatalhöyük. J Anthropol Archaeol 30:292–305

Sampson A (ed) (2011) The cave of the Cyclops. Mesolithic andNeolithic networks in the northern Aegean, Greece. Bone toolindustries, dietary resources and the paleoenvironment andarchaeometrical studies. Prehistory monographs 31 vol. II.INSTAPAcademic, Philadelphia

Sánchez-Goñi ME, Eynaud E, Turon JL, Shackleton NJ (1999) Highresolution palynological record off the Iberian margin: direct land–sea correlation for the Last Interglacial complex. Earth Planet SciLett 171:123–137

Schiegl S, Stockhammer P, Scott C, Wadley L (2004) A mineralogicaland phytolith study of theMiddle Stone Age hearths in Sibudu cave,Kwazulu-Natal, South Africa. S Afr J Sci 100:185–194

Schlichtherle H (1988) Neolithische Schmuckperlen aus SamenundFruchtsteine. In: Küster H (ed) Der PrähistorischeMensch und seines


Umwelt. Forschungen und Berichte zur Vor- und Frühgeschichte,Baden-Würtemberg 31. Theiss, Stuttgart, pp 199–203

Seidenkrantz MS, Bornmalm L, Johnsen SJ, Knudsen KL, Kuijpers A,Lauritzen SE, Leroy SAG,Mergeal I, Schweger C,VanVliet-LanoëB(1996) Two-step deglaciation at the oxygen isotope stage 6/5e transi-tion: The Zeifen-Kattegat climate oscillation. Quat Sci Rev 15:63–75

Shahack-Gross R, Finkelstein I (2008) Subsistence practices in an aridenvironment: a geoarchaeological investigation in an Iron Age site,the Negev highlands, Israel. J Archaeol Sci 35:965–982

Shahack-Gross R, Albert RM, Gilboa A, Nagar-Hilman O, Sharon I,Weiner S (2005) Geoarchaeology in an urban context: the uses ofspace in a Phoenician monumental building at Tel Dor (Israel). JArchaeol Sci 32:1417–1431

Shillito LM (2011) Simultaneous thin section and phytolith observation atfinely stratified deposits from Neolithic Çatalhöyük, Turkey: impli-cations for paleoeconomy and early Holocene paleoenvironment. JQuat Sci 26(6):576–588

Stiner M, Kozłowski J, Kuhn S, Karkanas P, Koumouzelis M (2010)Klissoura Cave 1 and the Upper Paleolithic of Southern Greece incultural and ecological contexts. Eurasian Prehistory 7(2):309–321

Stravopodi E, Manolis S, Kyparissi-Apostolika N (1999)Palaeoanthropological findings from Theopetra Cave in Thessaly:a preliminary report. In: Bailey GN, Adam E, Panagopoulou E,Perles C, Zachos K (eds) The palaeolithic archaeology of Greeceand adjacent areas, proceedings of the ICOPAG conference,Ioannina. British School at Athens Studies 3. British School atAthens, Athens, pp 271–281

Strömberg CAE (2004) Using phytolith assemblages to reconstruct theorigins and spread of grass-dominated habitats in the Great Plainsduring the Late Eocene to Early Miocene. Pal Pal Pal 207:239–275

Tsartsidou G, Lev-Yadun S, Albert RM, Rosen AM, Efstratiou N, WeinerS (2007) The phytolith archaeological record: strengths and weak-nesses evaluated based on a quantitativemodern reference collectionfrom Greece. J Archaeol Sci 34:1262–1275

Tsartsidou G, Lev-Yadun S, Efstratiou N, Weiner S (2008)Ethnoarchaeological study of phytolith assemblages from an agro-pastoral village in Northern Greece (Sarakini): development andapplication of a Phytolith Difference Index. J Archaeol Sci 35:600–613

Tsartsidou G, Lev-Yadun S, Efstratiou N, Weiner S (2009) Use of spacein a Neolithic village in Greece (Makri): phytolith analysis andcomparison of phytolith assemblages from an ethnographic settingin the same area. J Archaeol Sci 36:2342–2352

Tzedakis PC (1999) The last climatic cycle at Kopais, central Greece. JGeol Soc 156:425–434

Tzedakis PC, Frogley MR, Heaton THE (2003) Last Interglacial condi-tions in southern Europe: evidence from Ioannina, northwestGreece. Glob Planet Chang 36(3):157–170

Valladas H, Mercier N, Froget L, Joron JL, Reyss JL, Karkanas P,Panagopoulou E, Kyparissi-Apostolika N (2007) TL age-estimatesfor the Middle Palaeolithic layers at Theopetra Cave (Greece). QuatGeochronol 2:303–308

Weiner S (2010) Microarchaeology. Beyond the visible archaeologicalrecord. Cambridge University Press, New York

Weiss E, Wetterstrom W, Nadel D, Bar-Yosef O (2004) The broadspectrum revisited: evidence from plant remains. PNAS 101:9551–9555

Zurro D,MadellaM, Briz I, Vila A (2009) Variability of the phytolith recordin fisher–hunter–gatherer sites: an example from the Yamana society(Beagle Channel, Tierra del Fuego, Argentina). Quat Int 193:184–191


Palaeoenvironmental reconstruction and flora exploitation at the Palaeolithic cave of Theopetra,...

Documents

Transcript of Palaeoenvironmental reconstruction and flora exploitation at the Palaeolithic cave of Theopetra,...