macroscopic plant remains from mediterranean caves and rockshelters: avenues of interpretation

Geoarchaeology: An International Journal, Vol. 16, No. 4, 401–432 (2001)� 2001 John Wiley & Sons, Inc.

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Macroscopic Plant Remains from

Mediterranean Caves and Rockshelters:

Avenues of Interpretation

Julie Hansen

Department of Archaeology, Boston University, 675 Commonwealth Avenue,

Boston, Massachustts 02215

Macrobotanical remains from caves and rockshelters in theMediterranean provide substantialinformation about past human use of the sites as well as the surrounding environment. Themodes of deposition of both fresh and carbonized plant material in the past are varied and itis not always possible to distinguish among the geogenic, biogenic, and anthropogenic pro-cesses. Once deposited, seeds and other plant parts may be preserved through desiccation,mineralization, or, most commonly, carbonization, depending on the environment of the caveor rockshelter as well as human and other animal activities. It is assumed that large quantitiesof carbonized remains are the result of human activity, and such dense deposits can be usedas a measure of the intensity of occupation of the site. Where sufficient remains of woodcharcoal are recovered from stratified deposits, it is possible to identify the local vegetationand changes in the surrounding plant communities through time. When compared to otherenvironmental information, such as pollen studies, it is possible to obtain a more completepicture of the environment and to identify refugia for Mediterranean plants during the Pleis-tocene. In some cases, plants that are underrepresented or not at all represented in pollenspectra from lacustrine deposits may be recovered from the archaeological sites.� 2001 JohnWiley & Sons, Inc.

INTRODUCTION

A number of issues should be addressed when considering the interpretation ofmacroscopic plant remains from caves or rockshelters. Key among these is thevariety of pathways of deposition. In general, palaeoethnobotanical reports tendto concentrate on the human exploitation of the plants recovered from archaeo-logical sites, often with the implicit assumption that the remains are primarily theresult of anthropogenic activity. Macroscopic plant remains, however, can be in-troduced into cave and rockshelter sediments through a variety of pathways, bothnatural and anthropogenic. It is important to understand the different taphonomicprocesses that result in seeds, wood, or other plant parts being deposited andpreserved in these sites in order to assess more accurately their relevance to thehuman occupation of the area and their value as an indicator of paleoclimate.Another issue is the extent to which the plant remains can be directly linked to

past human activity. In some instances the density of remains can be an indicationof intensity of occupation of the site. Dense deposits of carbonized wood and other

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plant material are more likely the result of repeated burning events over time thana few or brief episodes of cave use.Lastly, it is important to examine the issue of environmental indicators from the

plant remains. Palynological studies from lacustrine sites are the primary sourceof data on past vegetation of an area, but may not provide sufficient evidence ofplants located near the cave or rockshelter under study, especially if the pollencoring area is a significant distance from the archaeological site. Both seeds andwood charcoal can provide valuable environmental information that can be usedto supplement palynological and other environmental data. Macroscopic remainsfrom a cave or rockshelter provide evidence of local vegetation and may includespecies not represented in lacustrine pollen spectra, and, in the absence of pollen,these data provide the primary source of information on local past vegetation.This article discusses the pathways of deposition and preservation of macro-

scopic plant remains in caves and rockshelters using data from several prehistoricMediterranean caves that have yielded stratified material. Having identified thevarious pathways of deposition, the relationship between density of anthropogenicremains and intensity of occupation will be examined. Finally, the value of collec-tion and analysis of these remains for environmental reconstruction of the areaaround a site will be illustrated.

PRESERVATION OF ORGANIC MATERIAL

Desiccation

Preservation of organic material through desiccation occurs because of the ex-ceptionally and consistently arid conditions since the time of their deposition. Thisprevents organisms such as molds, fungi, insects, or rodents from surviving andconsuming the organic material. Dry caves, such as Nahal Hemar in Israel (Kislev,1988; Figure 1), preserve a wide range of plant foods and artifacts in this way.Under desiccating conditions nearly all parts of a plant can be preserved, and fragilestructures such as fibers or fine hairs on fruits or seeds can be important for ac-curate identification of the species.

Mineralization

Another type of preservation of plant material in caves and rockshelters isthrough mineralization. This occurs in sites where dissolved minerals, such asCaCO3 in groundwater, are absorbed by the organic material and replace the cel-lular structure upon evaporation. In other instances, high phosphate levels in thedeposits can result in the plant material becoming mineralized with calcium phos-phate (Carruthers, 1991; Green, 1979). Phosphatic mineralization typically occursin the presence of highly organic deposits, such as in latrines and middens wherehuman and other organic waste is concentrated. In a cave or rockshelter, birdguano and other animal feces may accumulate either during or in the absence ofhuman occupation and provide such concentrations of phosphate (Goldberg andBar-Yosef, 1998; Wattez et al., 1990).

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Figure 1. Map showing locations of principal sites mentioned in the text; 1: Nahal Hemar; 2: Kebara; 3: Hayonim; 4: Zas Cave; 5: Franchthi Cave;6: Klithi; 7: Konspol Cave; 8: Grotta Azzurra, Grotta Caterina, Grotta dell’Edera; 9: Fontbregoua; 10: La Salpetriere; 11: Laroque II; 12: Abeurador;13: Dourgne; 14: Margineda; 15: Cova del Frare; 16: Arene Candide; a: Lake Gramousti pollen core; b: Lake Ioannina pollen core.

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Figure 2. Pathways of deposition of carbonized plant material in caves and rockshelters.

In some sites, such as Franchthi Cave in Greece (Figure 1) (Hansen, 1991), min-eralized seeds were recovered from the same deposits in which carbonized remainsof the same species were found. It is difficult to explain this mixture of preservationwithin a relatively small area (2 � 2 m trench). One possible explanation is thatsome of the seeds are not in situ, perhaps having been exposed to mineralizingconditions in a different deposit and being displaced by anthropogenic, biogenic,or geogenic processes. Thus, it is critical to have an understanding of the sedimentsources in which the remains have been found (Woodward & Goldberg, this issue).Another reason for the appearance of both mineralized and carbonized remains

in cave and rockshelter deposits may be microenvironmental variations such aswater dripping from the cave roof, or deposition of animal or other waste in par-ticular areas. In the Upper Palaeolithic layers at the entrance to Kebara cave inIsrael (Figure 1), Goldberg and Bar-Yosef (1998:112) note the presence of “localizedmm-size impregnations of CaCO3 that is presumably derived from water drippingbeneath the brow of the cave.” Although no mineralized plant remains were recov-ered from these sediments, material deposited in this area would have been subjectto mineralization under these conditions. At Konispol cave in southern Albania(Figure 1), Schuldenrein (1998:513) has identified Stratum 7 as having been depos-ited under wet conditions when a “ponded basin developed under anaerobic con-

MACROSCOPIC PLANT REMAINS FROM MEDITERRANEAN CAVES AND ROCKSHELTERS

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ditions,” and he notes the presence of microvegetal remains in the stiff clays. Theseare mineralized silica skeletons of, as yet unidentified, plant parts. The wood char-coal from these levels is coated with clay, and some of the pieces are also impreg-nated with minerals to some extent.

Carbonization

The most common means of preservation of organic remains in all types of sites,and one with which this article is primarily concerned, is through carbonization.Carbonization (also referred to as charring) results when organic material is ex-posed to heat in the range of approximately 200–400�C, or to higher temperaturesin the absence of oxygen (such as when the material is smothered in ash), to theextent that it is reduced to about 60% elemental carbon (Hillman, 1981:139; Lopinotand Brussell, 1982). It is generally the harder, denser parts of the plants such asseeds, wood, and nutshells that are more readily preserved in this way, althoughsuch soft organs as grapes, pear fruits, or tubers have been recovered in a carbon-ized form (Bookidis et al., 1999; Hansen, 1991; Hillman et al., 1989, respectively).Carbonized remains are much less subject to destruction by the variety of orga-nisms that would otherwise consume the fresh organic material. Plants can becarbonized through natural means outside the cave as a result of lightning strikesor through anthropogenic processes such as campfires, field burning, or forestclearing (Figure 2). Inside the cave or rockshelter, they are most likely to be car-bonized as the result of deposition in a hearth fire (Figure 3).

DEPOSITION OF ORGANIC MATERIAL

Plant material, fresh or carbonized, can be deposited in caves or rockseltersthrough natural or anthropogenic processes, and it may be difficult or impossibleto distinguish the mechanism of deposition from examination of the remains them-selves. Figure 2 illustrates the various pathways for botanical material carbonizedoutside the cave to be deposited inside, and Figure 3 shows how fresh plant ma-terial could enter the cave or rockshelter and subsequently be preserved throughcarbonization, desiccation, or mineralization. It will be useful to examine the nat-ural and anthropogenic processes separately.

Natural Processes

Eolian

Wind-blown seeds, fruits or plant debris, either part of the natural “seed rain” orresulting from human activity such as crop processing, can be deposited throughthe cave mouth or other openings. In dry caves these remains are preserved throughdesiccation like plants brought in by inhabitants of the site. They may also besubject to mineralization through the processes discussed above, while some maybe incorporated into hearths and become carbonized. In most sites, however, muchof the fresh material will be consumed by microorganisms, insects, or rodents.

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Figure 3. Pathways of deposition and preservation of fresh plant material in caves and rockshelters.



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Wind will also pick up small carbonized remains from outside the cave and scatterthem through the cave mouth, sink holes, or other openings. Once mixed withanthropogenic material, it is impossible to identify the original mode of deposition.For this reason, palaeoethnobotanists working on cave and rockshelter sites in theMediterranean have not attempted to distinguish plant material deposited by eolianprocesses from those deposited through other means. One way to do this wouldbe through micromorphological examination of the sediments containing the car-bon.

Fluvial/Colluvial

Water percolating through the roof of the cave may pick up small, carbonized,or fresh seeds or other fragments and deposit them on the cave floor. In karsticsites, flowing water in the cave can also bring in allochthonous material. Inwashingthrough the mouth of the cave or rockshelter is another potential source of plantmaterial, and flood waters may also bring larger quantities of material into the site.In some cases the mechanism for deposition may be recognized through micro-morphological examination of the sediments as demonstrated by Weiner et al.(1998) for Layer 10 at Zhoukoutien, China, dated to about 400,000 yr B.P. This finelylaminated sediment with organic matter “is indicative of accumulation in quietwater” and suggests that the site was more open and subject to ponding at thattime (Weiner et al., 1998:252). As mentioned above, similar ponding was identifiedat Konispol cave in AlbaniaCarbonized or fresh material deposited through fluvial or colluvial processes will

often show the effects of abrasion and rolling. At Franchthi Cave, Greece, corngromwell (Lithospermum arvense) was predominant in the earliest Upper Palaeo-lithic (ca. 35,000–15,000 yr B.P.) levels excavated, with seeds numbering in thethousands in some excavation units (Hansen, 1991). Fresh seeds of Lithospermum

have highly sculptured seed coats with raised tubercles, but the specimens fromthe basal levels at Franchthi Cave were worn down, leaving a smooth, sometimespitted surface. They were found in a reworked terra rossa sediment (Farrand, 2000)and may have been part of the sediment prior to its deposition in the cave. Thiswould account for the abrasion of the seeds as the sediment moved via slumping,sliding, or as a thick slurry through the mouth or roof of the cave (William Farrand,personal communication).

Biological

Insects and rodents living in a cave bring in seeds that may be redeposited in ahearth through human activity that displaces sediments. Birds may introduce plantmaterial for nesting or food. At the cave site of Fontbregoua in southern France(Figure 1), microlaminated layers dated to the Epipalaeolithic (10,000–8,000 yrB.P.) were found to be the result of the “slow accumulation of bird excreta asso-ciated with various organo-mineral residues also produced by birds; feather re-mains, finely divided wood pieces (twigs), bone fragments, seeds” (Wattez et al.,

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1990:432). At the site of Abeurador (Figure 1), also in southern France, micromor-phological examination of a sediment sequence identified three distinct layers: awhitish yellow layer derived from burning deciduous wood; a dull orange layercomposed of phosphatic aggregates containing burnt plant remains and bird bonefragments; and an underlying dark brown layer containing rotted plant residues.The dark brown layer has been identified as fragments of plants brought in by birds“forming a kind of litter which was subsequently humified” (Wattez et al., 1990:436). Birds of prey, such as owls, may also deposit seeds from the gut of their kill(MacPhail et al., 1997). For example, many owl pellets were collected from themodern surface at Franchthi cave and proved to contain seeds of several taxa.Organic material deposited through these means may be consumed by insects orother animals, or destroyed by bacteria and fungi. However, some organic materialmay be preserved through mineralization because of the high phosphate contentof the bird guano. Organic material may become desiccated if deposited in verydry caves, or become carbonized if accidentally incorporated into hearth depositsthrough human activity.Some plant remains may also be introduced in dung from herbivores utilizing

the site. Caves and rockshelters used by shepherds for lambing or for shelteringflocks during inclement weather can have a considerable build-up of dung overtime. For example, at Franchthi Cave, a thick layer of sheep and goat dung on thesurface was routinely harvested by the inhabitants of the area for fertilizer prior to1969 (Jacobsen, 2000). Similarly, the site of Klithi in Epirus, Greece (Figure 1), hada layer of recent goat dung 10 cm thick covering the prehistoric deposits (Baileyand Woodward, 1997:62). While most of the plant material in sheep and goat dungis finely comminuted, some whole or partial seeds or other identifiable plant partsmay be present. The high phosphate content of dung and the high calcium contentof karstic caves and rockshelters in the Mediterranean may result in some plantmaterial being mineralized as calcium phosphate. Microscopic examination of thesediments of Facies 3 at the site of Arene Candide in Italy (Figure 1) revealed alayer of ash containing burned fragments of sheep/goat coprolites with “maceratedtissue of possible woody Quercus material and dicotyledonous leaves” (Macphailet al., 1997:56), as well as cattle coprolites. Twig, branch, and leaf remains wereidentified in some layers, and, in other layers, abundant phytoliths indicate theburned remains of grass or straw. The alternation in composition of the layerssuggests a seasonal difference in plant availability and the similarity between thesedeposits and the surrounding ashes suggest that the plants were brought into thecave as fodder (Wattez et al., 1990:437).Dung present on the floor of a cave could be kicked into the hearth where the

plant material could become carbonized. The dung may also be deliberately burnedas fuel or refuse. One method of identifying burned dung is through micromor-phological examination of the sediments. Deposits at Grotta Azzurra, Grotta Ca-terina, and Grotta dell’Edera in the Trieste region of Northeastern Italy (Figure 1)provide substantial evidence of burned dung (Boschian, 1997; Boschian and Mon-tagnari-Kokelj, 2000). In addition to ash, micrite, and plant oxalates transformed



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to CaCO3 by combustion, “the occurrence of coprolitic aggregates, phytoliths, andcalcareous spherulites is evidence of the presence of ruminants in the cave” (Bos-chian and Montagnari-Kokelj, 2000:343). These could be primarily sheep and/orgoats but possibly also cattle based on the morphology of some of the coprolites.At Konispol Cave in Albania, Paul Goldberg has identified spherulites in hearthdeposits from the Neolithic levels (Goldberg, personal communication). It is inter-esting that the faunal study suggests that the cave was used for lambing during theNeolithic period (Nerissa Russell, personal communication; Harrold et al., 1999).Dung may have accumulated on the cave floor and been deliberately or accidentallyincorporated into the hearths.

Anthropogenic Processes

In addition to the natural processes of deposition through wind, water, or bio-logical activity, seeds and other plant parts will be brought into the cave by humansfor some purpose. The refuse from plant processing, as well as wood and otherbotanical material, may be used as fuel, thus providing the bulk of the macroscopiccarbonized remains that are recovered from archaeological sites. Refuse from plantprocessing and food preparation may also be disposed in the hearth or accidentallydropped in. There is no way to distinguish these different pathways with any cer-tainty, however.Although it is not always possible to identify the mode of deposition as either

anthropogenic or natural, when large quantities of carbonized plant remains arefound it can be assumed that they are primarily the result of human activity. Indeed,the relative density of such remains may be an indication of the intensity of oc-cupation of the site over time.

DENSITY OF ORGANIC MATERIAL AS A MEASURE OF INTENSITY

OF HUMAN OCCUPATION

The density of organic material on the floor and/or in fill deposits of a cave orrockshelter may be taken as an indication of the intensity of human occupation ofthe site. Year-round or seasonal occupation of the site will result in accumulationof carbonized material either in the hearths, on the floor, or concentrated in dumpsor pits. Occasional use of the site, on the other hand, will produce less buildup ofcarbonized remains. With deep, stratified deposits representing long periods of caveuse, it is possible to demonstrate variation in the density of carbonized plant re-mains. This information may be used to infer changes in the intensity of humanoccupation through time. Unfortunately, few published reports of plant remainsfrom Mediterranean caves and rockshelters provide this level of information. Atvarious sites in France, Spain, Andorra, and Portugal, carbonized plant remainswere collected through flotation from 1 m squares excavated in 5 cm levels (Badelet al., 1991). Thus each level would be approximately 0.05 m3 (5 L) of sediment,not including stones and artifacts. Floated samples that yielded more than 250macroscopic fragments (�5 mm) of wood charcoal were considered to be fairly

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Figure 4. Grams of flot (predominantly botanical remains) per 10 L of excavated sediment in trenchFAS at Franchthi Cave, Greece.

rich. The problem with trying to assess the intensity of occupation from these datais that they are based on the absolute count of wood fragments. A group of 100fragments of the same species may have been derived from a single branch of atree, so this measure is difficult to interpret in terms of intensity of occupation.Another means of determining density of carbonized remains is to use the totalweight of all the carbon from a sample (not just macroscopic wood charcoal) in agiven volume of sediment. Data from Franchthi Cave, Greece, are presented hereas an example of how such information can provide insights into intensity of oc-cupation.All floated remains (flot), predominantly carbon, were weighed, and the number

of grams per liter of excavated sediment was determined (Hansen, 1991). Figure 4presents these data from Franchthi Cave trench FAS, where a nearly completesequence of occupation is preserved. The botanical zones were determined on thebasis of the assemblage of plants in contiguous excavation units (layers) (Hansen,1991). Zones I and II contained predominantly the Lithospermum mentioned



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Figure 5. Rate of deposition of botanical remains in trench FAS at Franchthi Cave, Greece.

above, along with two other species of the Boraginaceae family (Alkanna andAnchusa) and small amounts of wood charcoal. These seeds have a high silicacontent in their seed coat and are thus preserved without carbonization. The den-sity of flot (predominantly seeds) in Zone I is only 0.08 g/10 L of excavated sedi-ment, while that in Zone II is 0. 6 g/10 L, and the bulk of this is the Boraginaceaeseeds that were most likely deposited through geogenic rather than anthropogenicprocesses. The rate of deposition of the plant remains, including Boraginaceae,was less than 50 g per 100 years (Figure 5).While there is other evidence in the form of minute fragments of carbon, lithics,

and animal bones in Zones I and II to suggest human occupation, the origin of someof this material is questionable. Indeed, the carbon may have been depositedthrough geogenic processes such as colluviation, as suggested above for the Lith-ospermum, rather than as the remains of fuel from hearths. Based on the evidencefrom trench FAS, human occupation of Franchthi Cave was sporadic during theUpper Palaeolithic.In trench H1B in the center of the cave, there is less than one gram of flot per

10 L of sediment in Zones I and I/II, while in Zone II the amount increases to about6 g (Figure 6). Nearly all of this is comprised of seeds of Boraginaceae in the samerolled and worn condition as seen in FAS and presumably deposited in the sameway. The number of grams per 100 years (Figure 7) also indicates a very slow rateof deposition. The major difference between FAS and H1B is in the length of thehiatus identified in the lithostratigraphic analysis (Farrand, 2000). In FAS there is

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Figure 6. Grams of flot (predominantly botanical remains) per 10 L of excavated sediment in trenchH1B at Franchthi Cave, Greece.

Figure 7. Rate of deposition of botanical remains in trench H1B at Franchthi Cave, Greece.



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Table I. Correspondence of botanical zones and lithostratigraphic units in FAS.

Botanical ZoneExcavationUnit Stratum

ExcavationUnit

I 227–207 P 227–224Q 222–218R 217–210S1 208

II 206–205 U 206–203II/III 204–199 V 202–199III 198–176 W1 196–197IV 175–168 W2 192–163IV/V 167–160V 159–147 W3 159–153

X1 148–150V/VI 146 Xcut 151–146VI 145–136 X2 145–136VI/VII 135–129 Y1 128–133VIIb 128–109 Y2 126–84VIIc 108–73 Xcut 83–76

a hiatus of about 2400 years between Strata R and S1, and another longer hiatusof about 7300 years between Strata S1 and U. Stratum S2 is completely missing inFAS (Table I), but in H1B it roughly corresponds to botanical Interzone I/II (TableII). In H1B there are also hiatuses between Strata R and S1 of about 2000–2500years and between S1 and S2 of about 5000 years. Thus, there was differentialsedimentation within the cave, probably from varying anthropogenic activity aswell as geogenic processes. Whatever the type of habitation, it is clear from thepaucity of carbonized plant remains that occupation was not very intensive. Indeed,the scattered nature and small size of the carbonized wood fragments may indicatenatural rather than anthropogenic deposition, perhaps brought in through colluvialprocesses as suggested for FAS, at least in Zone I. There are carbonized seeds suchas lentils in Zone II in H1B that may indicate human activity.More consistent or long-term occupation of the cave is indicated in Zones III–

V, where the density of carbon in FAS reaches a peak of more that 25 g/10 L ofsediment. These zones correspond primarily to the Mesolithic period dating fromapproximately 10,200 to 9600 yr B.P. In H1B, the amount of carbonized plant re-mains is considerably less, on the order of only 5.0–5.5 g/10 L of sediment. Therewas also less carbon deposited per 100 years in H1B than in FAS. These data mayindicate that less burning and/or less dump and scatter of carbon occurred in thecenter of the cave than near the west wall where trench FAS is located. Perhapsthe area of FAS was where most of the hearth cleaning was deposited, thus leavingrelatively little in the center of the cave.There is a significant decrease in the density of plant remains in Zone V and

Interzone V/VI in FAS, the latest Mesolithic. A similar drop in other material suchas lithics, shell, and animal bone is also seen in these levels (Payne, 1973, 1975;

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Table II. Correspondence of botanical zones and lithostratigraphic units in H1B.

Botanical ZoneExcavationUnit Stratum Excavation Unit

I 215–171 P 215–214Q 213R 212–181S1 180–173

I/II 170–169 S2 172–159 e/172–166 wII 168–154 T 159–158/166–155 wII/III 153–152 U 154–151III 151–119 V 150–148

W 147–117X1 115

IV 118–106 X2 114–80

Perles, 1990; Shackleton, 1988). Plant remains were not recovered from corre-sponding levels in H1B, but other categories of material are reduced in this trenchas well. The paucity of remains suggests a decrease in intensity of occupation andutilization of the cave at this time. This may have been a prelude to the apparentabandonment of the site suggested by the depositional hiatus of about 600 yearsidentified in the lithostratigraphy between Strata X1 and X2 (Farrand, 2000). Thisbreak precisely corresponds to the end of botanical Subzone V/VI and the beginningof Zone VI, the early Neolithic.The earliest Neolithic deposits in Zone VI indicate a continued low level of use

of the cave. Carbonized plant remains were deposited at a rate of about 16 g/100years in these levels (Figure 5). There is a resurgence in Zone VII representing theMiddle, Late, and Final Neolithic. During the Neolithic period at this site, structureswere built outside the cave and the function of the cave may have changed at thistime. Perhaps some people continued to utilize the cave for seasonal habitation,as it remained cooler during the summer.It is interesting to compare the rate of deposition of carbon with the rate of

sedimentation in the lithostratigraphic units identified by Farrand (2000) (Figures8 and 9). In these figures the rate of deposition of carbon from those excavationunits that are included in the lithostratigraphic units was calculated. As can be seenin Tables I and II, these units do not correspond exactly to the botanical zonesdiscussed above; hence, the graphs of grams/100 years based on the botanical zonesdo not exactly duplicate the graphs seen in Figures 8 and 9. The carbon is onecomponent of the sediment, but its rate of deposition is dependent on anthropo-genic factors that may not affect the other components of the sediment. Botanicalzones I and II correspond approximately to lithostratigraphic units P, Q, R, and S1(Tables I and II) (Farrand, 2000) and are not shown in Figures 8 and 9. Farrandnotes that the sedimentation rate in these strata is difficult to determine due to thelack of adequate radiocarbon ages. The tephra that comprises Stratum Q is dated



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Figure 8. Comparison of rates of sedimentation and deposition of botanical remains in trench FAS atFranchthi Cave, Greece.

between 33,000 and 40,000 yr B.P.,1 while the two ages from Stratum R are around25,800–26,850 yr B.P. The estimated rate of sedimentation for all the strata wasvery slow, with an average of about 1.25 cm/100 years from Q to the base of S1(Farrand, 2000:84–85).There are hiatuses, especially in FAS, within this sequence that may indicate

periodic abandonment of the cave and lack of anthropogenic deposits and/or ero-sion of sediment. Nonetheless, it is clear from the botanical data that very littlecarbon was being deposited during periods of sediment accumulation.In the latest Upper Palaeolithic and Mesolithic levels (Botanical Zones III– IV/V)

the intensity of occupation suggested by the significant increase in deposition ofcarbon per 100 years is corroborated by the rate of deposition in Stratum W thathas been estimated to be as much as 250 cm/100 years (Farrand, 2000:84). Abundantother cultural debris, including large quantities of land snail, account for much ofthis accumulation.The rate of sedimentation decreases dramatically in Strata X1(latest Mesolithic),

with a similar decrease in the rate of deposition of carbon. Farrand (2000:51) hasidentified “an unconformity at the top of this unit that may mark the Neolithic/Mesolithic boundary.” Utilization of the cave, at least in the area of trench FAS,continued at a fairly low level after this, as seen by the low rate of deposition inStrata X2 and Y1, corresponding to the early Neolithic. There is a dramatic increasein deposition of carbon in Stratum Y2, the Middle and Late Neolithic, while the rateof sedimentation does not increase significantly. This may indicate intensive utili-

1 All dates given in this article are in uncalibrated radiocarbon yr B.P.

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Figure 9. Comparison of rates of sedimentation and deposition of botanical remains in trench H1B atFranchthi Cave, Greece.

zation of the cave for activities involving plant processing and/or cooking duringthe later Neolithic with relatively little sediment deposited through cultural or nat-ural processes.

POST-DEPOSITIONAL PROCESSES

As with any category of material in a habitation site, plant remains are subjectto a variety of processes that can alter or displace them. Seeds and other smallplant parts can move up or down through cracks in the deposits or as the result oftectonic or biological activity. Carbonized remains can be crushed, dispersed, orremoved as a result of humans or other animals trampling and digging in the de-posits. Goldberg and Bar-Yosef (1998) have shown how animal burrows have par-tially destroyed hearth deposits in the sites of Kebara and Hayonim in Israel (Figure1). In the Mediterranean region with its very hot, dry summers, carbonized remainsnear the surface may also dry out, near the mouth of the cave or rockshelter. Thisvery dry carbon tends to break up when exposed to percolating moisture duringthe wet autumn and winter. Over time this may result in the carbon being reducedto small flecks and dispersed in the sediment. This is demonstrated in the shallowshoreline (Paralia) deposits outside Franchthi Cave. Small flecks of carbonizedmaterial were seen during excavation, but all attempts to recover any identifiableremains failed due to the minute size of the fragments (Hansen 1991). However,micromorphological analysis can be a very powerful tool for identifying such small



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plant remains. At several cave sites in Israel Goldberg et al. (1994) have shownhow even individual cells or minute tissue fragments of carbonized plants can beidentified in thin sections of stratified blocks of sediment.

PLANT REMAINS AS A RECORD OF ENVIRONMENTAL CHANGE

Range of Species Represented

A growing number of studies of macroscopic wood charcoal fragments from cavesites in France and Spain have shown that the identification of plant species canbe valuable in reconstructing past vegetation communities near the site (e.g., Heinz,1991; Ros Mora and Vernet, 1987; Vernet et al., 1987a, 1987b). However, in orderto do this, several assumptions must be made. As discussed above, it is assumedthat large quantities of carbonized plant remains in an archaeological deposit areprimarily the result of anthropogenic activity, having been brought into the cavefor a variety of purposes and either accidentally or deliberately burned. It alsoseems logical to assume, in the case of wood brought in for fuel, that it was col-lected within the site catchment (generally about 2 h walking distance) rather thana greater distance away. Although this does not preclude the possibility that moreexotic plant species may be deposited, they are unlikely to be dominant membersof the assemblage. A site located in an ecotone where several different vegetationcommunities may be accessible within easy walking and transport distance maybe expected to provide evidence of plants from more than one community. Lastly,it is assumed that in addition to obvious hearth deposits, carbonized plant remains,including wood charcoal, will be scattered around the hearth and floor surfaces.At Hayonim and Kebara caves in Israel, for example, Goldberg and Bar-Yosef (1998:121) note that “it has become clear that ashes were spread outward from the heartharea to cover a larger surface, which would perhaps be adequate for sleeping” (seealso Weiner et al., 1995). Some charcoal would also be present among these ashes.Additional large concentrations of carbonized remains may also be found as dumpsof hearth cleaning, either on a floor surface or in pits (Goldberg and Bar-Yosef,1998).Whether a site was occupied year-round, seasonally, or only occasionally, hearth

deposits will contain a record of the more recent burning events while carbonscattered on the floor or dumped in a heap will represent the remains from peri-odically cleaning out the hearth. Depending on the pathways for deposition andcarbonization of plant material in a cave or rockshelter, the result can range froma thin scatter of plant fragments to dense concentrations of material. Hearth de-posits will represent the most recent burning episodes and, therefore, may have amore restricted range of species than the dumps or scattered carbon on the cavefloors and in fills (Heinz, 1991; Perles, 1977). These latter deposits are the result ofcarbon from repeated fires and will contain a more representative range of speciesused over time. This has been demonstrated by Christine Heinz (1991) at the siteof Abeurador in western Languadoc, France, located 590 m above sea level (asl)on a limestone promontory overlooking the Aude valley (Figure 1). The site today

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lies in the meso-mediterranean vegetation zone dominated by evergreen oak. Strat-ified cultural deposits from the Late Palaeolithic to the Middle Neolithic have beenexcavated (Vaquer and Barbaza, 1987). Mesolithic hearth deposits show only twoprimary species while the floor and fill deposits from the same level provide a morecomplete range of species, including those found in the hearths (Figure 10). Thus,while it is important to identify all of this material, for the purposes of environ-mental reconstruction, floor and fill deposits should form the primary source ofdata and plant remains should be collected from these levels.

Environmental Reconstruction

Most of the macroscopic carbonized plant remains in a cave or rockshelter arepresumed to be anthropogenic in origin. Although, as discussed above, geogenicand biogenic processes may also deposit plant remains in these sites, the quantityof such material that is ultimately preserved, either through carbonization, miner-alization, or desiccation, will be minimal compared to the larger amounts of seeds,fruits, and wood brought in by humans for food and fuel. These resources may bebrought in from a considerable distance away, such as when the site is used as ashort-term shelter or for other purposes by people who have brought plant foodsand fuel with them. A good example is the Late Neolithic–Early Bronze Age siteof Zas cave on the Island of Naxos, Greece (Figure 1), where the macrobotanicalremains consist of a variety of crop plants such as barley and legumes (Zachos,1990). The area around the site would have been unsuitable for growing such crops,and it is likely that they were produced in the valley and brought up to the caveby people on a hunting trip or by shepherds tending their herds and using the caveas a temporary shelter. Where sites were occupied for longer periods as a primaryhabitat or even a seasonal habitat occupied for several months, plant food re-sources and fuel may have been gathered from the areas closer to the site, althoughthis does not preclude the possibility of more distant resources being exploited.With respect to fuel, however, it seems more likely that trees and shrubs growing

within a walking distance of 1–2 h would have been exploited first before moredistant sources were utilized. Such plants would, therefore, comprise the bulk ofthe species of macroscopic charcoal remains found in the deposits and could be avaluable supplement to pollen data from lake sediment records that may representmore regional vegetation. At the site of Dourgne in the Aude valley, Languedoc,France (Figure 1), Guilaine et al. (1987, p. 551) note that “the palynological andcharcoal data allow confirmation of the presence in the proximity of the site ofCorylus, lime (Tilia cordata), blackthorn (Prunus spinosa), (Pinus cf. sylvestris),Abies, and Juniperus at the end of the Preboreal,” suggesting that the species werecollected from the local vegetation.In the western Mediterranean, an intensive study of macroscopic fragments of

wood charcoal from caves and rockshelters in southern France and Spain hasprovided substantial environmental data for this part of the Mediterranean. In heranalysis of macroscopic charcoal fragments from floor and fill deposits at Abeu-



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Figure 10. Number of taxa carbonized wood from two hearths (a, b) and scattered fragments in habi-tation level C6 at Abeurador Cave, France (from Heinz, 1991:304, Figure 2, reprinted with permissionof the publisher).

rador, Heinz (1991) identified 28 taxa throughout the stratigraphic sequence thatshe divided into two phases on the basis of changes in species (Figure 11). Phase1, covering the Epipalaeolithic to the Middle Mesolithic, is subdivided into twosubphases; subphase 1a encompasses the Epipalaeolithic, and subphase 1b in-cludes the Middle Mesolithic.Subphase 1a represents dry mountain conditions in the region during the late

glacial, with conifers, including pine and juniper, dominating the floral assemblage.This vegetation suggests a forest-steppe linked to an open landscape where moun-tain influences and cool, dry climate prevailed. Species such as Prunus amygdalus

(almond) would have been present in sheltered areas where more moisture was

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Figure 11. Relative percentage of wood charcoal taxa from Aberuador Cave, Herault, France (from Heinz, 1991:309, Figure 6, reprinted with permissionof the publisher, Elsevier Science Ltd.).



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available. The appearance of almond in these early levels is paralleled at other LatePleistocene cave sites in the Languedoc region such as la Salpetriere and LaroqueII (Bazile-Robert, 1987) dated to around 13,000 yr B.P. These data also correspondwell with palaynological studies from lacustrine deposits in the region (Jalut, 1987).In subphase 1b (Middle Mesolithic) at Abeurador, there is a decrease in pine and

almond disappears from the samples. Juniper, on the other hand, increases andbecomes the dominant species. Trees such as deciduous oak, maple, elm, and haw-thorn would have been associated with a deciduous oak forest, while warmer Med-iterranean conditions are indicated by evergreen oak, buckthorn (Rhamnus), andpistachio. The interpretation of the expansion of juniper, according to Heinz (1991:319) is controversial in that it may represent either natural expansion of this speciesdue to an ameliorating climate or human disturbance of the vegetation. Clearingof the oak forest vegetation by humans would favor the spread of juniper. Heinznotes, however, that a peak in juniper pollen from lacustrine sites in southernFrance is dated to the Lateglacial, around 15,000 yr B.P., and a core from Padul,in Granada province, Spain shows a dominance of juniper between ca. 15,000 and9000 yr B.P. with a peak at 10,000 yr B.P., a period when forest clearance by humanswas less likely. This, in turn, would suggest that the spread of juniper was influ-enced by environmental conditions rather than human interference. Further anal-ysis of both palynological and palaeoethnobotanical samples is necessary to re-solve this issue.The Neolithic at Abeurador is represented by Phase 2 (8000–6000 yr B.P.) (Figure

11). The dominant plant is deciduous oak, with other species associated with amore dense closed canopy forest. This is paralleled at other sites in southernFrance such as Margineda in the Herault valley (Figure 1), as well as several Neo-lithic sites in Spain such as the Cova del Frare in the Matadepera region nearBarcelona (Figure 1) (Ros Mora and Vernet, 1987). Palynological studies in thePyrenees (Jalut, 1987) support this interpretation of an expansion of deciduous oakforest between ca. 8000 and 6000 yr B.P.Wood charcoal is the primary type of plant material recovered from Konispol

Cave, Albania (Hansen, 1999). The site is located about 400 m asl on a limestoneridge overlooking the Pavel valley (Figure 1). Figures 12, 13, and 14 show themacroscopic wood charcoal remains from trenches VIII, X, and XXI, respectively.The radiocarbon ages need some clarification here. An age of 26,370 � 180 yr B.P.(Beta-67799) in unit 42 of trench VIII is considered too early (Ellwood, 1996, 1997;Harrold et al., 1999); it is thought that the deepest deposits are no more than14,000–15,000 years old (Harrold et al., 1999). There is a reasonably consistent setof radiocarbon ages for Trench XXI with the earliest age at 8900 � 180 yr B.P.(Beta-80001). In this sequence, however, the age of 7410 � 80 yr B.P. (Beta-79999)is thought to be too young (Ellwood, 1996, 1997).The lowest levels of trenches VIII and X at Konispol cave contain a few Upper

Palaeolithic artifacts (Harrold et al., 1999) and can be correlated with Strata 7, 8,and 9 of the lithostratigraphic sequence in the center of the cave (Schuldenrein,1998). Stratum 9 consists of weathered yellow clay (terra fusca), possibly deposited

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Figure 12. Relative percentage of wood charcoal taxa from Trench VIII, Konispol Cave, Albania. Percentages are based on the sum of identifiedfragments in each unit.

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Figure 13. Relative percentage of wood charcoal taxa from Trench X, Konispol Cave, Albania. Percentages are based on the sum of identified fragmentsin each unit.

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HFigure 14. Relative percentage of wood charcoal taxa from Trench XXI, Konispol Cave, Albania. Percentages are based on the sum of identifiedfragments in each unit.



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by flowing water. Thus the carbonized plant remains in Stratum 9 may not havederived from human activity in this part of the cave but may be from a hearthdeposit elsewhere in the site, or perhaps from outside the cave. Several sinkholesin the cave roof would have provided an inlet for water and sediment. All of theunits in trench VIII (Figure 12) can be correlated to Stratum 8 identified by Schuld-enrein (1998) as redeposited and oxidized clayey sand, possibly the result of lo-calized spring seep. Hence, the wood charcoal from these units may not be in situ

and may have been derived from anthropogenic activity elsewhere inside or outsidethe cave, or is the product of natural fires, with the charcoal deposited with thesediment. Whatever the mode of deposition, the wood can give us a few clues aboutthe natural vegetation in the area of the site. The rest of the Upper Palaeolithicrepresented in the cave is correlated with Stratum 7, which consists of reducedorganic-rich clay with small plant remains. As already noted, Schuldenrein (1998)suggests this may have been deposited in standing water in the cave. The pondingof water may correlate with an increase in precipitation at the beginning of theHolocene. A radiocarbon age of 11,410 � 80 yr B.P. (Beta-56414) at the base of thisstratum (unit 28 in trench VIII) provides some temporal control.Juniper (Juniperus sp.) is present in the lowest levels of trench X, but almond

(Prunus amygdalus) predominates after unit 27, that is in later deposits, in thistrench as well as in trench VIII (Figure 13). Wood of maple (Acer), plane tree(Platanus), and buckthorn (Rhamnus) also appear in the early deposits. Planetrees may have been growing near the spring to the north of the cave or in thevalley along the river. The presence of almond also indicates a refuge where moremoisture was available during the relatively cold and dry phase of the early Lateg-lacial, between about 13,000 and 15,000 yr B.P.The wood charcoal from Konispol Cave can be compared to pollen data from

Epirus in northwestern Greece (Figure 1). From Lake Ioannina, at a similar ele-vation to Konispol, Bottema (1974) has shown that between 28,000 and 10,000 yrB.P. oak and pine predominate in the sparse arboreal pollen, withmaple and juniperalso present, as at Konispol (Figure 15). For the period from 13,000 to 10,000 yrB.P., pollen data from Lake Gramousti at an elevation of 400 m asl (Figure 1) showsthat trees were still sparse and deciduous oak and juniper would have formed anopen forest or steppe-forest (Figure 16) (Willis, 1997, 1992). Species such as al-mond, plane tree, buckthorn, and pistachio are barely or not at all represented inthese Late Pleistocene spectra. Almond trees are insect-pollinated and, therefore,produce little pollen that does not travel great distances when carried by the wind.Consequently, almond is not represented in pollen spectra, and the wood charcoalfrom sites such as Konispol Cave provide the only evidence of their presence inthe region.Almond continued to dominate the plant remains from the Mesolithic levels

(9000–7000 yr B.P.) at Konispol, but additional species occur at this time as well,such as ash (Fraxinus) and deciduous oak seen in trenches X (Figure 12) and XXI(Figure 14). These species suggest expansion of mixed deciduous woodland as aresult of the warmer and wetter climate. The Mesolithic levels correspond to Strata

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Figure 15. Pollen percentage diagram for Lake Ioannina, Epirus, Greece. Percentages are based on total pollen sum. Tick marks � 5% for solid curvesand 2% for open curves. Open curves represent pollen percentage �5 (from Bottema, 1974: Figure 31, reprinted with permission of the author).

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New Phytologist).

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6c–d (Schuldenrein, 1998). Stratum 6c is composed of organic sand and silt withcultural debris in the form of lithics, and Stratum 6d is culturally sterile clay. Theclear evidence of human activity in Stratum 6c, especially in the western part ofthe cave (trench XXI), would lend support to the interpretation that the woodcharcoal in these levels is anthropogenic in origin. In Epirus, the pollen data fromIoannina (Bottema, 1974; Figure 15) and Lake Gramousti (Willis, 1992, 1997; Figure16) indicate that during the early post-glacial (ca. 11,000 yr B.P.) deciduous oakexpanded and formed a mixed deciduous forest with a variety of species not rep-resented in the Konispol remains.The Early to Middle Neolithic remains in trench XXI and IX at Konispol show a

spread of Mediterranean species, such as buckthorn and pistachio, that suggestsome open vegetation and somewhat drier conditions. However, deciduous oakand elm are also present in these levels, as seen in Trench X, so a mixed woodlandmay be a more accurate reconstruction here. The deposits are identified as Strata6a–b for the Early Neolithic and Stratum 5 for the Middle Neolithic, and are largelyanthropogenic in origin, with a mixture of ash lenses, hearths, cultural debris, andanimal bone. There are several radiocarbon ages in trench XXI that place the Earlyto Middle Neolithic cultural deposits between about 7000 and 6000 yr B.P.Willis (1997) notes a change in the vegetation in the region of Lake Gramousti

beginning around 7000 yr B.P. At this time oak begins to decline and fir (Abies),hornbeam (Carpinus orientalis/Ostrya), and hazel (Corylus) increase. Subse-quently, all arboreal pollen decreases and herbaceous species increase. Willis(1997) attributes this change to human activity in the form of clearing woodlandfor animal husbandry. At Konispol, it is interesting to note that cereal agricultureis evidenced at this time by the presence of emmer wheat (Triticum turgidum

dicoccum) in the Neolithic levels. At the same time, herding was also being prac-ticed, as indicated by the sheep and goat bones. There is nothing in the plantremains, however, to suggest any land clearance in the area of Konispol at thistime.It is clear that macroscopic plant remains can provide evidence for local vege-

tation in the area of a cave or rockshelter. These data can also be used to supple-ment palaynological studies of lacustrine deposits to detect species not representedin pollen spectra. It is also useful to correlate the plant remains with the lithostra-tigraphy of the site, both to aid in determining the origin of the deposits and toexamine the microenvironment of the remains.

CONCLUSION

Macroscopic plant remains from caves and rockshelters can be of considerablevalue not only for identifying possible food and fuel resources, but also for detect-ing vegetation change around the site and, thus, provide some indication of climatechange. In order to interpret the remains correctly, however, it is important tounderstand the mechanisms by which they have entered the site. Fresh seeds andother plant parts can be deposited in cave sites through a variety of pathways and



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subsequently become preserved in the sediments of the site. It is not always pos-sible to determine what the ultimate source of these remains was, however. Seedsand twigs can be deposited by birds nesting in the site and birds of prey will depositseeds from the gut content of their prey. These plant materials can become pre-served through desiccation, mineralization, or carbonization as a result of the en-vironmental conditions in the site and/or incorporation into a hearth. In addition,birds provide some of the mineralizing phosphate in the guano that is depositedon the cave floor. Other plant material may be brought in by insects or other ani-mals, or be washed or blown into the site and become preserved through the samevariety of processes.The most common method of preservation of plant remains in caves and rock-

shelters (indeed, on all archaeological sites) is through carbonization. Botanicalmaterial can be burned outside the cave as a result of lightning strikes or fromhuman activity and subsequently be blown into the cave or brought in as a com-ponent of colluvial or fluvial sediments. Such remains will often be very small andbroken up and may also be rolled and abraded. Plant material will also be carbon-ized inside the cave or rockshelter when deposited in a hearth by humans, whetheraccidentally or deliberately as food or fuel. It is not always possible to distinguishthe anthropogenic remains from those brought in by some other vector, althoughit is assumed by palaeoethnobotanists that large quantities and dense concentra-tions of plant remains are usually the result of human activity.The density of plant remains in cave deposits can be one indication of the inten-

sity of occupation of the cave by humans. Piles or dumps of ash and carbon, aswell as defined areas of burning surrounded by carbon scatter, can be interpretedas hearth cleaning, and plant material should be collected from these features.When a hearth is raked out, carbonized remains will often be spread over the floorsurface, and these remains will represent species from repeated firings, while theremains within the hearth may be from the last one or two uses of the feature.Thus, for environmental reconstruction, it is necessary to sample the floor and filldeposits to obtain the maximum variety of species over time.A stratified sequence of wood charcoal can provide a substantial amount of in-

formation on the local vegetation of a cave or rockshelter through time. These datacan be compared to palynological studies from lacustrine sediments in order todevelop a more accurate reconstruction of the environment. Some species, suchas almond, may occur as wood charcoal in archaeological deposits but will not befound in pollen cores because they are insect-pollinated. Thus the macroscopicremains can broaden our interpretation and provide information on refugia forspecies that would otherwise go undetected.This article has attempted to illustrate the value of macroscopic plant remains

and show how they can be used in conjunction with sediment analyses and otherenvironmental data to interpret the use of caves and rockshelters. The increasingnumbers of such studies in the Mediterranean have substantially broadened ourunderstanding of past human interaction with the environment from the Late Pleis-tocene through the early Holocene. However, more studies are needed in parts of

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the eastern Mediterranean where there are many cave sites but little botanical data.Further work on the processes of deposition and analysis at the microscopic levelmay help to increase our ability to recover botanical data from such sites.

I am grateful to Paul Goldberg, Jamie Woodward, and an anonymous reviewer for their comments onvarious drafts of this paper. I would also like to express my thanks to Karl Petruso, Frank Harrold, andJoe Schuldenrein for information on aspects of the Konispol sequence, and to Bill Farrand for infor-mation on the Franchthi sequence. Any errors in the final publication are my responsibility.

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Received July 1, 2000

Accepted for publication November 24, 2000

macroscopic plant remains from mediterranean caves and rockshelters: avenues of interpretation

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