klissoura cave 1 and the up per paleolithic of south ern greece in cul tural and ecological...

7/27/2019 KLISSOURA CAVE 1 AND THE UP PER PALEOLITHIC OF SOUTH ERN GREECE IN CUL TURAL AND ECOLOGICAL CONT

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tional mixing at the MP-UP layer contact is indi -cated.

The formation of the Upper Paleolithic throughMesolithic stratigraphic sequence was dominated byanthropogenic processes. Constructed hearths of di-verse forms, ash dumps, raked-out ash features,and trampled ash remains are very common (Kar-kanas, this issue). The ear liest Upper Paleolithiclayer (V) truncates the top of the Middle Paleo-lithic sedimentary series. As of this writing, LayerV represents the only Uluzzian occupation docu-mented in Greece (though unpublished finds fromKephalari Cave may also be Uluzzian), and its in-dustry resembles essentially contemporary indus-tries in It aly. The Aurig nacian layers have a gen -

erally discrete appearance, and the earliest Auri-gnacian layer (IV) truncates the underlying de-posits.

The Aurignacian components in Klissoura 1are distinguished from all other cultural layers bythe remarkable complexity of in situ hearths,which range from simple small basin or stackedforms to clay-lined types (Karkanas et al., 2004).Remnants of a small structure are demarcated inthe ear liest Aurignacian layer (IV) by a roughlyoval scatter of large stones, a dis crete organic

stained area within, and an exceptionally denseconcentration of perforated shell beads.

Following the upper Aurignacian layers aretwo enigmatic cultural horizons. The definition ofthese horizons suffers in part from the lack of in-formation for the region and period, but also fromambiguities concerning the behavioral causes ofvariation in the lithic industries. Hypotheses forexplaining this variation include ethnic differencesand distinct traditions of tool manufacture and, al-ternatively, differences in the circumstances of oc-cupation by essentially similar groups by season.We offer some preferred explanations below butacknowledge that resolution of these questions ul-timately requires more information from as yet un-discovered Upper Paleolithic sites in the Pelopon-nese.

Mod erately rich faunal and lithic assem-blages were obtained from the Epigravettian andMesolithic layers. These cul tural layers ex pe ri-enced frequent truncations and considerable dis-

turbance, mainly from human activity, and diffuseinterfaces characterize the contacts between them.The second major hiatus in the stratigraphic series

corresponds to the LGM, after which appears thetypical middle (or early-middle) Epigravettianwith conspicuous links to south-central Italy. Thethird hiatus sep arates the Epigravettian (layersIIa-d) from the Mesolithic (layer 5a). Here, how-ever, cultural continu ity can be seen despite the

presence of depositional hiatus. The occurrenceof the Terminal Paleolithic marked by a Epigra-vettian tradition in neighboring caves of the Klis-soura Gorge indicates that Epigravettian groupsdid not abandon the region at this time, but ratheroccupied different caves and shelters in the area.

The rich and varied archaeological record ofKlissoura Cave 1 provides an unprecedented andfor the moment unique body of information about

the Upper Paleolithic of southern Greece. Coher-ent lithic, bone, shell and osseous tool assembla-ges and many fea tures and spa tial data were re-corded and studied in this collaborative research

project. This detailed record of Upper Paleolithicactiv ities yields several of surprises and insightson Upper Paleolithic behavior and cultural diver-sity in Eurasia, including the great age of the earli-est Upper Paleolithic occupation and the contextsof on-site activities throughout the Upper Paleo-lithic.

CULTURAL SEQUENCE BASED

ON LITHIC ASSEM BLAGES

The sequence of cultural horizons and lithicassemblages does not document an uninterruptedlo cal evolution of Paleolithic cultures (Kacza-nowska et al., this is sue). Instead, it is character-ized by cultural and occupational discontinuities.The Upper Paleolithic begins with the Uluzzian(layer V), replaced by the Aurig nacian as repre-sented by layer IV and layers IIIg-a. Layer IIIcontains a non-Aurignacian Upper Paleolithic in-dustry. The Gravettoid component in layer IIIcontains what we have described as a Mediterra-nean backed bladelet/blade in dus try. The tech-nological character and stylistically distinct na-ture of the Gravettoid assemblage rules out a localevolution, suggesting instead origins from differ-ent regions of the mid-northern Mediterranean re-

gion. Layers 6, 6a, and 6/7 in fact represent themixed filling of what may be an anthropogenicditch and will not be discussed further.

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The artifactual record in Klissoura Cave 1 af-fords important insights on the process of culturalevolution and differentiation in the region. Abovethe Uluzzian layer V is a series of Aurig na cianlayers (IV, IIIa-g) overlain by layer III that couldrepresent the final phase of the Uluzzian. Higherin the stratigraphic column, layer III contains aMediterranean backed bladelet/blade industry,which, after sedimentological/erosional hiatus, isoverlain by the Epigravettian (layer II). These ob-servations from the lithic assemblages (Kacza-nowska et al., this issue) suggest that the variousculture units identified in Klissoura 1 correspondnot so much to the adaptation of the same foragersto specific raw materials and ecological condi-

tions, but rather represent different cultural tradi-tions corresponding to distinct groups who peri-odically coexisted within the larger region.

Culturally diagnostic artifact forms in Klis-soura 1 include the following: arched backed

blades and convex truncations for the Uluzzian;carenate cores/endscrapers and micro-retouched

bladelets for the Aurignacian; backed blades andbladelets for the Med iterra nean Early Gravet-toid; component various backed blades (alsowith ventral retouch), para-geometric forms and

shouldered points for the Epigravettian; and geo-metric microliths for the Mesolithic.

Technological diversity is quite narrow fromthe Upper Paleolithic through the Mesolithic as-semblages and contrasts with the cultural taxon-omy based on the indicative ar tifact classes. Thelimited technological variability in these assem-

blages is probably due to the fact that a fairly ho-mogenous group of primarily local raw materials(radiolarites and flints) were exploited throughoutthe sequence (Koumouzelis et al., 1996). The pro-

portion of radiolarites remains fairly sta ble(6070%) through time, whereas the proportionof flint is more variable; the lowest frequency offlint is found in the Aurignacian layers (2029%),and the highest in layer III (42%) and the Meso-lithic (3337%). Extra-local raw materials arerare throughout the layers, with some of the redradiolarites in layer V being of the highest quality.Unquestionably exotic materials from the latercultural layers include Melian obsidian from

Mesolithic layer 5a. The rel atively uni form com-po sition of the raw material across assemblages,and the dominance of local raw materials overall,

suggest a fairly limited range of mobility of all ofthe groups while inhabiting this cave. Local rawmaterials were determined from field survey ofraw material outcrops over a radius of 34 kmaround the site. The nearest known sources ofhigher quality stone are Mesozoic outcrops lo-cated in the northeastern Peloponnese. Severalsites on the Argolid, particularly those of theEarly Neolithic, contain these high quality rawmaterials, but Klissoura Cave 1 does not.

The structure of the major technological cate-gories also is similar across most of the lithic as-semblages. The large quantities of lithic shat ter/chunks in all lev els are clearly attributable to thelow quality of the lo cal raw materials. Tab ular

fragments of both radiolarite and flint from thestudy area con tain many in ternal frac tures andflaws as a rule. The high frequency of chips andsmall flakes is also partly explained by the use of

poor quality raw materials, but it simultaneouslyattests to the inten sive use of these materials, in-cluding the frequent rejuvenation of retouchedtools.

Shatter/chunks, chips and small flakes consti-tute more than half of the lithic ar tifacts. This

property of the stone tool assemblages reduces the

indices of other technological categories cores,blades, and retouched tools. Indices for the lattercategories nonetheless are consistent with valuesfor other sites at which the full cycle of blank andtool production and diverse activities took place.The high indices of backed pieces used as insertsin the Uluzzian and Gravettoid industries indicatethe use of hafts, apparently made from perishablematerials. In the Aurignacian levels, on the otherhand, the bladelet index is low, and carenatecores/endscrapers from which the bladelets weredetached, occur in large quantities. This patternsuggests that some bladelets were taken awayfrom the site during the Aurignacian occupations,hafted as exchangeable parts of weapons, tools or

both of these. Macro scopic impact fractures andmicroscopic marginal scars indicate that bothAurignacian micro-retouched (or unretouched)

bladelets and Gravettoid backed bladelets wereused for projectile hunting weapons and as insertsfor cutting or scraping tools.

The contrast be tween the Aurignacian andother industries is most evident in the tools/corestypological group. In the Aurignacian levels, end-

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scrap ers/cores (mainly carenate forms) are verycommon, with an index of about 60 for the indus-tries of layers IV and IIIe-g. By contrast, thescraper/core index is between 16 and 36 in theUluzzian, and about 30 in levels III and II. Suchhigh tool to core indices in the Aurignacian aredue to the dual roles of carenate pieces, whichfunctioned both as cores for bladelets and asendscrapers. In other lev els, bladelets were de-tached from other, spe cialized core types, whileendscrapers functioned primarily as tools for ac-tivities such as hide processing.

The indices of blades and bladelets do notfluctuate very much through the Upper Paleolithicsequence, oscillating between the values of 4 and

6. It is only in the Late Mesolithic (layer 3) thatthe blade index is significantly higher (9.5). Moresignificant differ ences can be seen among the cul-tural layers in the frequency of flake tools (i.e.denticulated, notched and retouched flakes). Anespecially high index of these tools (20) is foundin layer III and in some of the Aurignacianlayers.

DATING

The dates for the GravettoidAurignacian se-quence (layers III, IIIe-g and IV) are relativelywell constrained. They show a generally mono-tonic trend of increasing age, from ca. 2729 14Ckyrs BP to ca. 3233 14C kyrs BP. These ages areconsistent with other classic or late Aurigna ciansites in southern Europe (Kuhn et al., this issue).

The Uluzzian indus try in layer V clearly pre-dates the Aurigna cian stratigraphically, but the ra-diocarbon results for layer V are ambiguous.Based on one date on a sample reported to befrom layer V (source lo cation is unclear, see Kuhnet al., this issue), and two dates from layer VI(which is stratgraphically mixed), it is possible tosuggest an age of >40,000 years for layer V.Other radiocarbon data are significantly youngerand may or may not truly originate from layer V,the limits of which are locally difficult to distin-guish from layer IV.

Fortunately, microtephra analysis conductedas part of the RESET project has identified one

major peak concentration of tephra shards at theinterface of layers IV and V, tailing upwardly tolayer III, along with a minor peak at the interface

of layers VI and VII. Attempts currently are un-derway to correlate the tephra(s) with specificeruptions of known age (Dustin White, personalcommunication, 2010). In nearby Franchthi Cave,a wind-blown Y-5 tephra (Campanian Ignimbrite)was found in stratum Q. Originating from the Na-

ples area of Italy, this tephra is dated to 39.280.11 kyrs by 40Ar/39Ar (De Vivo et al., 2001).There is a strong pos sibility that Klissoura 1 mayalso contain this tephra.

Impor tantly, the tephra seals layer V in Klis-soura 1. The radiocarbon results suggest that theindustry of layer V is about 6000 years older thanmost currently reported ages for similar lithic as-semblages from Europe.

Karkanas (this issue) concludes that the tem-poral gap between layers VII and V is consider-ably greater than the gap between layers V andIV. Published ages from Uluzzian sites in Italytend to be significantly younger than 39,000 yearsBP (reviewed by Kuhn et al., this is sue). How-ever, almost all published dates for the Uluzzianinsouthern Europe should be considered minimumes timates, and current ef forts at re-dat ing the sitesare likely to push age estimates backward in time.

The ABOX pre-treatment tech nique was used

successfully used to obtain finite dates of >60,000years on wood charcoal from the Middle Paleo-lithic layers. Although these should be also takenas minimum ages, they illustrate the techniques

potential for pushing back the maximum limits ofradiocarbon dating. ABOX proved less usefulwhen applied to more recent Upper Paleolithicsamples for reasons that may include greater wa-ter circulation or other sources of contamination.

The two important depositional/occupationalhi atuses in the later part of the Klissoura 1 se-quence correspond to major paleoclimatic events.One of these occurs within se quence B betweenthe Med iterranean Backed Bladelet layer III andthe Epigravettian (layer II). Be cause layer III isdated at about 3028 Kyr and the Epigravettiancannot be older than 1615 Kyrs BP, this hiatusshould include the Last Gla cial Maximum (LGM)within Marine Oxygen Isotope Stage (MIS) 2.Sites dated to the LGM are absent in the gorge aswell as in other parts of Argolid. The only cultural

en tities that are chro nologically close to LGM arelithic phase II in Franchthi Cave (22.321.4 kyrsBP; see Perls, 1987) and probably layer D3 in



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Kephalari Cave, which unfortunately lacks radio-carbon dates. It is possible that the inland areaswere depopulated during the LGM.

The last major hiatus in Klissoura 1 occursbetween the geological sequences B and A, or be-tween the Epigravettian (layer II) and the lowerMesolithic (layer 5a). Given that the Epigra-vettian occu pation almost all cer tainly spans thetime interval from 16 to 14 kyrs BP, and theMesolithic is synchronous with the Early Holo-cene, the second hiatus must cover the end of theLate Glacial, spanning at least 12 to 10 kyrs BP.This most recent hiatus in Klissoura Cave 1 may

be unique to the site, since there are numerousLate/Fi nal Paleolithic sites in the area, including

the dated layers from Klissoura Caves 4 and 7(Koumouzelis et al., 2004), sites in the Voido-matis Gorge, and in other parts of Argolid such asat Kephalari Cave (layer C) and Franchthi Cave(Perls, 1987).

PALEOENVIRONMENTS

AND PALEOCOMMUNITIES

OF THE ARGOLID

The results from the botanical, faunal and geo-

logical analyses suggest a gradual trend toward cli-matic cooling through the Upper Paleolithic se-quence in Klissoura Cave 1. Warmer, wet terconditions returned only well after MIS 2, or dur-ing the Mesolithic. These general conclusions can

be qualified by indications about the broader envi-ronment and the sedimentary environment insidethe cave.

Grass phytoliths are the most impor tant el e-ments in the phytolith assemblages from Klisso-ura 1 (Albert, this is sue). Most of the iden tifiedspecimens correspond to the C3 festucoid sub-family, which is very common in the Mediterra-nean basin. The grass phytolith assemblages oflayers IIIe-g and IV indicate only a moderatelyhumid environment. Pytoliths representing C4grasses and probably also reeds (Arundo donax)are present in the Epipaleolithic (II) layers, andreed phytoliths occur in the uppermost portion ofthe III sequence (III-III). Although reeds requirevery wet conditions, their presence may simply

indi cate small pockets of wet land somewhere inthe area and possibly localized changes in wa tertables caused by sea level changes, tec tonic

events or other non-climatic factors. The pres enceof C4 grasses, on the other hand, suggests a sig-nificantly drier and more open environmentduring the Epipaleolithic.

Wood charcoal remains from the Upper Pa-leolithic layers re flect a mosaic of peren nial vege-tation types (Ntinou, this issue). It is likely thatdry parkland vegetation covered the rocky hills,giving way to open woodland with mesophilousand thermophilous trees in the foothills and valleyfloors. Burned wood remnants of oak (Quercussp., deciduous type) and elm (Ulmus), genera that

prefer somewhat moister conditions such as mightoccur in gullies and small canyons, are most com-mon in layers V though IIIe-g. Elm all but disap-

pears from the charcoal assemblages thereaf ter.The wood-charcoal record of the early part of theUpper Paleolithic sequence indicates interstadialconditions during mid-MIS 3 (4030 kyrs BP)and gradual cooling and drying towards the end ofthe MIS 3 (after 30 kyrs BP).

The scope of variation in moisture that wouldhave been available to plants near Klissoura Cave1 warrants some discussion. Even the earliest Auri-gnacian phases, in which some moisture-depend-ent tree spe cies are rep resented, coin cided with

generally dry conditions. A marginal balance ofmoisture availability and water uptake by plantswas enough during this interval to support the de-velopment of some mesophilous and thermophi-lous vegetation. While this situation helped to sup-

press erosion in the area during the formation oflayer IV (and possibly layer V), conditions werestill suffi ciently dry to pre vent ash in the site from

becoming cemented by dripping water. Duringthe later occupations, the environment becamevery dry and, based on higher rates of clastic sedi-ment accumulation, more prone to erosion duringinfrequent storms. The main explanation for in-creased erosion in the area is a decline in pe ren-nial vegetation. Precipitation seems to have in-creased again only with the Mesolithic, but ero-sion was considerable until forest developmentcaught up with water availabil ity.

The composition of the mammal and avianfaunas in Klissoura 1 suggests correspondingchanges in animal community structure during the

Upper Paleolithic through Mesolithic (Bocheskiand Tomek, this issue; Starkovich and Stiner, thisissue). The ungulate assemblages from layers



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IIIe-g, IV and V are relatively diverse for theirsizes. Hares (and tortoises in V) dominate thesmall game fractions. The assemblages from themiddle and upper Aurignacian layers (IIId-a) areless rich in ungulates, and they are dominated byone ungulate species in particular, European fal-low deer (Dama dama). Hunting of large and me-dium-sized ground birds (bustard and partridges,respectively) also be came important in the in ter-val represented by layers IIId-a. The dominanceof fallow deer together with the high incidence of

partridges and bustards in the later Aurignacianlayers suggests an expansion of open grassy ar-eas. Ungulate diversity expands again in the Me-solithic (and possibly in the Epipaleolithic, but

this is a small sample), and hares once againdominate the small game fraction.It is unlikely that the vari a tions in ungulate

species diversity stem mainly from human hunt-ing pref erences. Changes in temper ature and sealevel forced qualitative shifts in the struc ture ofterrestrial animal communities on the Argolid. Amore heterogeneous environment would supporta broader range and more even proportions of un-gu late species, because greater macro-struc turalvaria tion in available hab itats makes it more dif fi -

cult for one species to outcompete others. Rela-tively heterogeneous habitats are indicated by thefaunal and botanical results for the earliest UpperPaleolithic. Subsequently, drier or cooler condi-tions prevailed and vegetation became more uni-form, allowingDama populations to dominate lo-cally.

The small animal component of the dietaryspectrum poses the main contra dic tion strictly cli-mate-driven patterns in prey choice. The overallcontri bution of small animals to the meat diet in-creased dramatically in layer III and above. Thisis clearly appar ent within the vertebrate assem-

blages, but also from the rising economic impor-tance of large land snails. Expansions in dietary

breadth are generally thought to represent eithertemporary or long-term responses by consumersto the decline in the most profitable resources(Stephens and Krebs, 1986) large game in thecase of the Paleolithic. Such trends may repre sentcultural preferences only in the sense of their be-

coming permanent solutions with time, supportedby significant technological investments made inspite of other important demands on foragers

time. It is likely, therefore, that the relent less ex-pansion in dietary breadth evidenced in KlissouraCave 1 reflects a growing human ecological foot-

print in the region and probably also mild in-creases in human population densities. That thetrend is evidenced principally within the smallgame fraction of the faunas is not surprising, asthese re sources were the main means for fillinggaps in the availability of large game animals(Kelly, 1995; Kuhn and Stiner, 2006).

The cultural sequence of Franchthi Cave onthe southern Argolid partly overlaps with themost recent part of the Klissoura 1 chronology(Perls, 1987; Farrand, 2000). The Aurignacianfauna from Franchthi is small and as yet un-

der-documented, but the large Gravettoid assem-blage dated to ca. 2122 kyrs appears to follow intime the III layer series of Klissoura 1. Althoughthe southern Argolid experienced sea changesmuch more directly than the Klissoura Gorgearea, the Franchthi data are of comparative inter-est.1 Five of the ungulate species found in theKlissoura 1 faunas also occur in the upper Aurig-nacian, Gravettoid, Epipaleolithic, and Meso-lithic layers of Franchthi Cave (Stiner and Munro,n.d.; Payne, 1975, 1982). Both red deer and Euro-

pean wild ass were important prey in the earlypart of the Franchthi sequence, but red deer wasthe only significant large prey item in the later

part. Other ungulates are represented in low fre-quencies, namely aurochs, wild pig and ibex. In-terestingly, no fallow deer remains were found inFranchthi,2 despite the singular importance of thisspecies in Klissoura 1. The contrast in dominantdeer species in Klissoura 1 and Franchthi are oneof several lines of evidence that cli mate-drivenenvironmental conditions were strongly influ-enced by local factors on the Peloponnese.

Another contrast in ungulate representationbetween the two sites concerns the European wildass. In Franchthi, wild ass remains dominateduring the Gravettoid occupations (Stiner andMunro, n.d.). Deer dominate the entire Upper Pa-leolithicMesolithic se quence in Klissoura 1 wherewild asses were always rare. The importance ofwild ass at Franchthi Cave during the Gravettoidoccupations must relate to more open conditions

on the southern end of the peninsula going intothe LGM. European equids of the Late Pleisto-cene are thought to have preferred open and



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steppic conditions, particularly E. hydruntinus.Modern fallow deer feed mainly in open grassyareas but must have some tree cover for protectionfrom the elements and predators. The optimalhabitat for fallow deer therefore is deciduous andmixed woodlands on gently rolling terrain. Reddeer, the dominant deer species in Franchthi, aregraz ers by prefer ence but can also feed on dwarfshrubs such as heather and other low quality

browse, provided that conditions are relativelymoist. They are also more tolerant of wet windsand cool, exposed conditions than are fallow deer.

The findings on the shell ornaments from Kli-ssoura Cave 1 also speak to questions about thedegree of environmental heterogeneity in south-

ern Greece dur ing the Late Pleistocene (Stiner,this issue). Although Klissoura 1 was never situ-ated on the Aegaen shore during the Paleolithic orMesolithic, the in habitants visited the sea andother aquatic habitats, and they brought manysmall or namental shells back to the site. Theseshells fall within a nar row range of sizes andshapes. However, the species collected during theearliest Upper Paleolithic phases are quite varied,whereas few species were uti lized for or na mental

purposes in the later occupations (above layers

IIIg-e). The great variety of ornamental molluskspe cies in the assemblages from layers IV and Vreflect a mo saic of aquatic hab itats, more complexthan exists in the Peloponnese today. The taxo-nomic diversity of the early Upper Paleolithic or -nament assemblages from Klissoura 1 also greatlyexceeds that of every post-Aurignacian ornamentassemblage from Franchthi Cave (Shackle ton,1988; C. Perls, personal communication, 2010).The more re cent shell as semblages from Klis-soura 1 and Franchthi invariably are dominated

by just a few brackish water and lagunal molluskspecies (Columbella rustica and Cyclope spp.).Reduction in taxonomic diversity in the marineshell types was almost certainly linked to changesin sea level.

Based on the dating and environmental data,the early Upper Paleolithic part of the Klissoura 1cultural se quence corre lates with an interstadialand the last minor high sea stand (ca. 35 kyrs cali-

brated BP), before the big drop in sea level that

began about 30 kyrs (calibrated BP) and culmi-nated in the LGM (Chappel, 2002; Wright et al.,2009). The high sea stand in the early Upper

Paleolithic helps to ex plain the great diversity ofcoastal and inland habitats, since el evated wa tertables would result in the formation of marshes,lagoons and sand bars. According to van Andel etal. (1990), for example, the northern part of theArgolis Gulf is an extended flat shallow area lessthan 50 m deep. During the last high sea stand ofMIS 3a this area would have been only partly sub-merged and thus would have supported a widerrange of aquatic hab itats. The de cline in sea lev elsthereafter probably also explains the abrupt de-cline in orna ment shell diversity in the layersabove IIIe-g in Klissoura 1.

HUNTING PATTERNSKlissoura Cave 1 lies at the inter face of rug-ged hills and a large plain, thereby providing ac-cess to a varied food supply. The site is locatedstrategically where the gorge opens onto the up-

per Argos plain. There is no indication of masshunting of ungulates at this site. Rather the datasuggest opportunistic hunting of a variety of spe-cies, one or a few at a time (Starkovich and Stiner,this issue). As the local animal community chan-ged with climate and vegetation, hunters respon-

ded opportunistically and pursued whatever un-gulate species were avail able. There is the ques-tion of where the Upper Paleolithic inhabitants ofKlissoura 1 obtained the ibex and chamois (layerIV only), since the area does not include true al-

pine habitats. In fact ibex may inhabit a muchwider variety of elevations, provided that the ter-rain is rugged (Phoca-Cosmetatou, 2004). Thelow but persistent presence of ibex in the ungulatefaunas may simply re flect the ex tent to whichhunters chose to search craggy uplands nearby.Today chamois tend to occupy rocky or alpine ar-eas, but they along with wild goats may descendto much lower, forested pastures in winter (For-syth, 2000; Baumann et al., 2005).

Small game ex ploitation at Klissoura 1 ex-hibit a pattern of increasing dietary breadth. Gen-er ally similar trends have been documented inother regions of the Mediterranean Basin (Tcher-nov, 1992; Hockett and Bicho, 2000; Stineret al.,2000; Stiner, 2001; Munro, 2004; Manne and

Bicho, 2009). Specifically, there is a decreasethrough time in the proportion of small, slow-moving game species and an increasing reliance



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of more productive quick small animals such ashares and/or birds. At Klissoura 1, this trend be-gins around the time of the Middle to UpperPaleolithic transition (Starkovich, 2010), thoughoccasional hare exploitation occurred before thistime, in the late Mid dle Paleolithic. The rel ativecontri bution of small animals to the meat diet in-creased further in the most recent layers (3-5a, II),where the NISP counts for quick small game actu-ally surpass those for medium-sized ungulates(Bocheski and Tomek, this issue; Starkovich andStiner, this issue).

The shells of the large edible land snail,Helixfigulina, also are prevalent in the late UpperPaleolithic through Mesolithic strata of Klissoura

1. Most of these were modified by humans ratherthan small predators, although none is burned byfire (Koumouzelis et al., 2001; Starkovich andStiner, this is sue). The relative quantity of landsnails in the archaeofaunas increases ex po nen -tially with time, and shell sizes become larger andmore uniform. Spe cifically, land snails are rare inlayers IV or V and show no clear evidence of hu-man modification, whereas human-modified snailshells are moderately abundant in layer IIIc andincrease greatly through layers III and III, and

snail abundance peaks in Mesolithic layers 3-5a.Epigravettian layer IIa-b rep resents a strik ing ex -ception in that snails are uncommon and a widerange of tiny to large species are rep re sented, sim-ilar to the natural snail assemblages that litter theground in the site vicinity today. Snails are notdifficult to find or collect after heavy rains, butcooking and extraction is relatively labor inten-sive.

Other findings on Upper Paleolithic subsis-tence at Klissoura 1 relate to large game hunt ing,spe cifically the patterns of prey age se lection andfood transport. Minor biases were found for body

part repre sentation in this site. These biases arenot explained by in situ attri tion and thereforemust reflect human transport decisions. The partsof ungulate skeletons are fairly evenly repre-sented, except for the scarcity of ax ial elements

below the neck. Nearly all meaty parts of car-casses were brought to the site for processing, andaxial parts were often left at or near the kill sites.

In fact the dressed weight for most of the prey ani-mals would not have exceeded what a few hunterscould carry back to camp within a day. The mor-

tality pat terns of the ungulates in dicate fairly evenrepresentation of young, prime adult and old adultin dividuals in all layers ex cept the early Aurig na-cian (IV), where old adults are less well repre-sented. The lack of strong age biases in the ungu-late faunas suggests a consistently emphasis onencounter hunting, without focusing on sex-sorted herds.

Small quan tities of fe tal or neo nate re mainsare present in the ungulate assemblages through-out the sequence, and most of these represent un-

born animals. Hunting of pregnant female fallowdeer must have taken place before or during thespring birthing season (late MayJune). Otherseasons of occupation are not necessarily ex-

cluded by these findings, however, since the phy-tolith evidence from layers IIIg, and to a lesser de-gree in IIIe, indi cates the pres ence of grass flow-ers. In modern Greece, grasses may begin bloom-ing in March, but most inflorescenses developfrom April to June, and at much lower frequenciesfrom July to September (Al bert, this issue).Hearths and large volumes of wood ash are also amajor component of the Klissoura 1 record, andsome of the occupations must have included coolmonths of the year.

Deer antler is proportionately common in theMesolithic (3-5a), though this is a small sample,and in layers III, IIIe-g, IV, and V (also a smallsample). Worked antler and (rarely) bone artifactsoccur in all of the layers, but most of these arefrom Aurignacian layers IIIe-g and IV. The layersthat contain the most antler fragments of any sortgenerally also contain the most worked antler arti-facts, with the excep tion of the Mesolithic (Star-kovich and Stiner, this issue). Microscopic analy-sis of the antler fragments revealed few unequivo-cal examples of trimming debris from antlerworking, but the lack of such evidence in this sitemay be explained by the extensive breakage of theantler by the humans and post-depositional micro-surface alterations (Christidou, personal commu-nication, 2010).

Male fallow deer pos sess antlers fromroughly July to April (Chapman and Chapman,1975), and the antlers harden in time for the au-tumn rut. Some of the male fallow deer in the Up-

per Paleolithic Klissoura as semblages thereforemust have died in the colder months of the year(autumn through early spring). It is possible, how-



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ever, that antler was col lected and curated overlong periods in anticipation of tool-making. Thusthe presence of ant ler in direct connection withosseous tool-making may not provide a reliableindication of the season of occupation.

PALEOLITHIC ORNAMENTS

Shell ornaments occur throughout the UpperPaleolithic, Epipaleolithic and Mesolithic lay ers(Stiner, this issue). As is gener ally true of the Up -

per Paleolithic in Europe, the ornament assem-blages from Klissoura Cave 1 are well developedin character. The earliest Upper Paleolithic orna-ments oc cur in layer V in asso ci a tion with an

Uluzzian industry. The largest assemblage of or-naments comes from the earliest Aurignacian orlayer IV. There are a few orna ments in layersVIVII, but most of these were found immedi-ately below the area of the man-made shelter inlayer IV. Layer V does not extend to this area ofthe excavation, and taphonomic evidence and di-rect dating of some of the shells (K. Douka, per-sonal communication, February 2010) indicatethat the shells in VIVII represent very localizeddownward intrusions from layer IV.

The Klissoura 1 ornament assemblages differfrom those typical of coastal sites in that the Klis-soura 1 assemblages consist almost exclusively offinished products. There is considerable evidenceof high-grading or human selection of the assem-

blages for harmony in shell color, form and quality,and there are few, if any, ex amples of man ufactur -ing errors. The prevalence of cord-wear suggeststhat many of the or na ments arrived al ready at-tached to organic materials or human bodies. What

breakage occurred to the shells resulted primarilyfrom long-term use. Faded or worn shells of spe-cies that would have originally been red in color(Clanculus spp.) were renewed with red ochre.

The ornament assemblages from the earliestUp per Paleolithic layers are particularly rich inspecies. The high frequency ofDentalium (tusk)shells none theless sets the small Uluzzian orna-ment assemblage apart from the Aurignacian andlater bead as semblages in the site. Dentaliumshells are also prominent in the Uluzzian horizons

of Grotta del Caval lo in south ern It aly (Palma diCesnola, 1966), though sample size variation maybe a complicating factor.

The ornament is considerable variation inabundance through the Upper Paleolithic andMesolithic layers in Klissoura 1 is not explained

by dif fer ences in the thickness of the ex cavatedlayers. As noted above, the assemblage from theearliest Aurignacian layer IV is exceptionallylarge, and most of the ornaments from this layeroccur within the inferred perimeter of the man-made shelter, ringed by large stones. This featureis surrounded by hearths but none occurs withinthe inferred area of the shelter where the or na-ments are concentrated.

SITE FUNCTION AND OCCUPATION

INTENSITYThe variety of features, artifacts and faunalre mains in the early Up per Paleolithic layers ofKlissoura 1 in dicate that the site served as somekind of residential base during most or all of theseoccupations. The intensity or duration of the oc-cupations probably varied greatly, however, withthe most intense use of the site occurring duringthe formation of Aurig nacian layers IV and IIIe-g.In addition to many clay-lined and unlinedhearths, these layers contain a diverse as sort ment

of lenses, pits and other features. Antler pointsand probable manufacturing debris, mainly onantler, are particularly abun dant in layer IV. Un-like the situation in the layers above, where osse-ous artifacts are widely scattered among hor izon-tal units, worked ant ler speciments are spatiallyconcentrated within and around the immediatearea of the shelter fea ture in layer IV (Christidou,

personal communication, 2010).The indications of human activities are only

somewhat less varied in the later part of the UpperPaleolithic-Epipaleolithic sequence. Unfortu-nately, the upper lay ers suffered from greateramounts of erosion or disturbance, possibly re-ducing the diversity of visible activity areas. Forexample, the material in layers 6, 6a and 6/7,which includes a mixture of Aurignacian andother artifacts, represents material that wasdumped episodically into a large pit. Its sedimentis homogenous, generally loose and porous, andthe matrix is almost pure ash and contains more

than the usual proportion of fragmented lithicdebitage. Normally such concentrated refusewould associate with an intensive occupation, but



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we find no horizontal layer associated with thepit. A significant erosional hiatus is apparent inthis part of the stratigraphic column (Karkanas,this issue), however, and some sediment mayhave been lost during the LGM.

Plant phytoliths are abundant in most of theUpper Paleolithic sediment samples, but they gen-erally are not found in the hearths. The contrast in

phytolith distributions between hearths and openareas testifies to the horizontal integrity of featuresin the Upper Paleolithic lay ers, consistent with thegeological observations. The input of plant matterinto fireplaces was selective mainly dicotyledo-nous wood and bark-producing plants. It is thegrasses that show the great est spatial separa tion

from the hearths. The abundant grass inflorescen-ces in Layer IIIe-g may point to their use as food,or the harvesting of mature stems for fiber work-ing or bedding. Small amounts of sedge phytolithsin layer IV, and reed phytoliths in layers II andIII-III could also relate to fiber working on site.There is considerable evidence of osseous tech-nology use and production on site (Christidou,

personal communication, 2010), particularly inlayers III IIIe-g and IV. Plant fiber working andhunting may rank among the many potential uses

of these tools. Clearly, a wide range of ac tivitiestook place in the cave, a situation typical of basecamps.

The record of fire use in the Upper Paleolithiclayers of Klissoura 1 is unusually complex (Kar-kanas et al., 2004; Karkanas, this issue). Hearthsare often superimposed in Klissoura 1, with re-

peated building and maintenance of fires in cer-tain areas of the site. The function of the clay-lined hearths in the Aurignacian layers is difficultto interpret from the sparse wood charcoal re-mains alone. However, the scar city of charcoal inthese layers might be the result of intentional pro-duction of hot embers for use in the clay hearths.Embers the incandes cent stage of a fire allowone to exploit the properties of conduction andcon vection to heat a small, semi-en closed space.Complete combustion from continuous stoking ofembers would destroy most of the wood charcoal

produced in these fires. Embers may also be usedto cook food indirectly with radiant heat, for dry-

ing and curing of food and other materials, andpossibly heating sweat lodges (saunas). Embertransfer and multiple uses of hearths likely oc-

curred during the formation of the Upper Paleo-lithic layers, particularly in layers where clay-lined hearths co-occur with unlined hearths.

CONCLUSION

The Paleolithic deposits in Klissoura Cave 1preserve a well-ordered stratigraphy that covers alarge portion of the Late Pleistocene MiddlePaleolithic and Up per Paleolithic, along with Epi-

paleolithic and Mesolithic components. The Up-per Paleolithic record is especially well preservedand complex due to a dense assortment of intactfeatures and artifacts. These cultural components

provide a rich and only partly overlapping com-

plement to the Paleolithic record of FranchthiCave on the southern Argolid.Among the important findings of this re-

search on the Upper Paleolithic through Meso-lithic in Klissoura Cave 1 are the identifica tion ofan early Uluzzian occupation in layer V that may

be more than 39,000 years old, the first UpperPaleolithic oc cu pa tion in the stratigraphic se ries,followed by a long and relatively well-dated Auri-gnacian sequence. Ornament assemblages appearsuddenly with the onset of the early Upper Paleo-

lithic (Uluzzian) in this site, and all intact (un-mixed) Middle Paleolithic layers that underlie theUpper Paleolithic lack or naments en tirely. Someinter-stratification of Aurignacian, later Uluzzian-like (III) and Gravettoid (III) components issuggested on the basis of formal artifact types.The lithic in dus tries in some lay ers of Klissoura 1show demonstrable, periodic links to Paleolithic

populations in ltaly, particularly the Uluzzian oflayer V.

Zooarchaeological findings indicate early in-creases in dietary breath dur ing the Upper Paleo-lithic, consistent with trends observed in otherMediterranean areas where this phenomenon has

been studied intensively. Diets expanded furtherdur ing the later Paleolithic and Mesolithic oc cu-

pations. The small game trends in Klissoura 1 andother Mediterranean faunal sequences are not ex-

plained by climate driven environmental changes,since they persist through MIS 3 and generally in-tensify in MIS 2 (Stiner, 2001).3

Paleoenvironmental data from the studies ofintact charcoal fragments, plant phytoliths, sedi-mentary characteristics, and animal species repre-



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sentation indicate moderate changes in moistureavail abil ity over the Upper Paleolithic throughMesolithic sequence. Overall, dry conditions pre-vailed in the area throughout the Upper Paleo-lithic, but moisture availabil ity was somewhatgreater during the formation of layers V and IV.Local environments became drier and cooler dur-ing the formation of the middle and upper Aurig-nacian layers and more so through the Epipaleo-lithic. Moisture availability increased again onlyin the Mesolithic.

Important hiatuses marked by erosion eventssep arate the Mid dle and Up per Paleolithic sedi-mentary groups. There is no seamless transition

between the two cultural en tities in this site.

Depositional hi atuses also separate the Uluzzianfrom the earliest Aurigancian, and the UpperPaleolithic layer se ries from the Epipaleolithic.There is no record of human occupation at Klis-soura 1 during the Last Glacial Maximum nor, ap-

par ently, at the nearby site of Franchthi Cave(Farrand, 2000).

Klissoura Cave 1 provides considerable detailabout the daily existence of Upper Paleolithic for-ag ers at one site. The sed imen tary fea tures, arti-facts and an imal remains tes tify a wide range of

on-site activities that is typical of residential bases.Anthropogenic processes greatly shaped the char-acter of the Upper Paleolithic through Mesolithicsediments, par ticularly the many cycles of hearth

building, cleaning, renovation and trampling. Hu-mans were also the main sources of disturbance inthe cave de pos its, as the re sult of creating and clea-ning hearth basins, clear ing activity surfaces, anddigging small pits. The remarkable clay hearths ofKlissoura 1 Aurignacian are unique as of this writ-ing, and may have been used as sat ellite fire in stal-lations to which hot coals were moved for the pur-

poses of cooking or heating activity areas. Theintensity of the occupations seems to have variedover time, with those of layer IV being particularlyintense. Greater post-depositional disturbances tothe later lay ers may have undermined the vis ibilityof features, and thus the relative intensity of thelater occupations is more difficult to judge, but siteuse does seem to have been lighter during theEpipaleolithic and Mesolithic.

The cultural sequences identified at KlissouraCave 1 and at other important sites in southernGreece (see Perls, 1987, 1999 on Franchthi

Cave; Panagopoulou et al., 20022004 on Lako-nis) testify to significant regional differences theclassic chronologies of western and central Eu-rope beyond the Balkans (e.g., Kozowski, 1999).As might be expected for a part of the world de-fined by distinctive ecosystems and a uniquely

broken and diverse topography, there is much evi-dence for regional or en demic patterns of cul-tural evolution. While processes that promote cul-tural divergence are to be expected for peninsularconditions, as the Peloponnese certainly repre-sents, there were also significant intervals of in-creased con tact westwardly across the upperAdriatic seabed, such as during the Uluzzian.

REFERENCES

BAUMANN M., BABOTAL C., SCHIBLER J. 2005.Native or naturalized? Validating alpine chamoishab itat models with archaeozoological data. Eco-logical Applications 15(3), 10961110.

BERKE H. 1984. The distri butions of bones from largemammals at Petersfels. In: H. Berke, J. Hahn, C.-J.Kind (eds) Upper Palaeolithic Settlement Patternsin Eu rope. Verlag Archaeologica Venatoria, Institutfr Urgeschichte der Universitat Tbingen, Tbin-gen, 103108.

CHAPMAN D.I., CHAP MAN N.G. 1975. FallowDeer, their history, distribution and biology. Te-rence Dal ton Ltd., Lavenham, Suf folk.

CHAPPELL J. 2002. Sea level changes forced icebreakouts in the Last Glacial cy cle: new results fromcoral terraces. Quaternary Science Reviews 21,12291240.

DE VIVO B., ROLANDI G., GANS P.B., CALVERTA., BOHRSON W.A., SPERA F. J., BELKIN H.E.2001. New con straints on the pyroclastic eruptivehistory of the Campanian volca nic Plain (Italy).

Mineralogy and Petrology 73, 4765.FARRAND W.R. (with JACOBSON T.W.). 2000.

Depositional history of Franchthi Cave: stratig ra-

phy, sedimentology, and chro nology.Excavations atFranchthi Cave, Greece, fasc. 7. Indiana UniversityPress, Bloomington, IN.

FORSYTH D.M. 2000. Habitat se lection and coexis-tence of the alpine chamois (Rupicapra rupicapra)and Himalayan tahr (Hemitragus jemlahicus) in theeastern Southern Alps, New Zealand.Journal of Zo-ology, London 252, 215225.

HOCKETT B., BICHO N. 2000. The rabbits of Pica-

reiro Cave: Small mammal hunting during the lateUpper Palaeo lithic in the Por tuguese Estremadura.Journal of Archaeological Science 27, 715723.



12/13

JOCHIM M. 1998. A Hunter-Gatherer Landscape:Southwest Germany in the Late Paleolithic and

Mesolithic. Plenum Press, New York.KARKANAS P., KOUMOUZELIS M., KOZOWSKI

J.K., SITLIVY V., SOBCZYK K., BERNA F.,

WEINER S. 2004. The earliest evidence for clayhearths: Aurignacian features in Klissoura Cave 1,southern Greece.Antiquity 78, 513525.

KELLY R. 1995. The Foraging Spectrum: Diversity inHunter-gatherer Lifeways. Smithsonian InstitutionPress, Washington D.C.

KOUMOUZELIS M., KOZOWSKI J.K., NOWAKM., SOBCZYK K., KACZANOWSKI M., PAWLI-KOWSKI M., PAZDUR A. 1996. Prehistoric settle-ment in the Klissoura Gorge, Argolid, Greece (exca-vations 1993, 1994). Prhistoire Europenne 8,143173.

KOUMOUZELIS M., GINTER B., KOZOWSKIJ.K., PAWLIKOWSKI M., BAR-YOSEF O., AL -BERT R.M., LITYSKA-ZAJC M., STWORZE-WICZ E., WOJTAL P., LIPECKI G., TOMEK T.,BOCHESKI Z.M., PAZDUR A. 2001. The earlyupper Palaeolithic in Greece: The excavations inKlissoura Cave.Journal of Archaeological Science28, 515539.

KOUMOUZELIS M., KOZOWSKI J.K., KACZA-NOWSKA M. 2004. End of the Palaeolithic in theArgolid (Greece): Exca va tions in cave 3 and Cave 7in the Klissoura gorge. Eurasian Prehistory 2, 3356.

KOZOWSKI J.K. 1999. The Evolution of the Bal kanAurignacian. In: S. Davies, R. Charles (eds) Doro-thy Garrod and the Prog ress of the Palaeolithic.Oxbow Books, Oxford, 97117.

KUHN S.L., STINER M.C. 2006. Whats a mother todo? A hypothe sis about the division of labor and mo-dern human origins. Current Anthropology 47(6),953980.

MANNE T., BICHO N. 2009. Vale Boi: Renderingnew understandings of resource intensification and

diversification in southwestern Iberia.Before Farm-ing2009/2, ar ticle 1, 121.MUNRO N.D. 2004. Zooarchaeological mea sures of

human hunting pressure and site occupation inten-sity in the Natufian of the south ern Lev ant and theimplications for agricultural origins. Current An-thropology 45, S5S33.

PALMA DI CESNOLA A. 1966. Il Paleolitico supe-riore arcaico (facies uluzziana) della Grotta del Ca-vallo, Lecce (continuazione).Rivista di Scienze Pre-istoriche 21, 359.

PANAGOPOULOU E., KARKANAS P., TSARTSI-

DOU G., KOTJABOPOULOU E., HARVATI K.,NTINOU M. 20022004. Late Pleistocene ar chaeo-logical and fossil human evidence from Lakonis

Cave, southern Greece.Journal of Field Archaeol -ogy 29, 323349.

PAWLIKOWSKI M., KOUMOUZELIS M., GINTERB., KOZOWSKI J.K. 2000. Emerg ing ceramictechnology in structured Aurignacian hearths at

Klissoura Cave 1 in Greece.Archaeology, Ethnolo-gy and An thropology of Eur asia 4, 1929.

PAYNE S. 1975. Faunal Change at Franchthi Cavefrom 20,000 B.C. 3000 B.C. In: A.T. Clason (ed.)

Archaeozoological Studies. North Holland Publish-ing, Amsterdam, 120131.

PAYNE S. 1982. Faunal Ev idence for En vironmen-tal/Climatic Change at Franchthi Cave (South ernArgolid, Greece), 25,000 B.P. 5000 B.P. Prelimi -nary Results. In: J.L. Bintliff, W. van Zeist (eds)

Palaeoclimates, Palaeoenvironments and Human

Communities in the Eastern Mediterranean Region

in Later Prehistory. BAR Internationl Se ries 133,Oxford, 133137.

PERLS C. 1987. Les industries lithiques tailles deFranchthi (Argolide, Grce). Tome I: Prsentation

gnrale et industries palolithiques, Excavationsat Franchthi Cave, fasc. 3. Indiana University Press,Bloomington, IN.

PERLS C. 1999. Long-term perspec tives on the occu -pation of the Franchthi Cave: continuity and discon -tinuity. In: G.N. Bailey, E. Adam, E. Panagopoulou,C. Perls, K. Zachos (eds) The Palaeolithic Archae-ology of Greece and Adjacent Areas, Proceedings ofthe ICOPAG Conference, Ioannina, Sep tember1994, British School at Athens Studies no. 3, Ath -ens, 311318.

PHOCA-COSMETATOU N. 2004. A Zooarchaeolo-gical Reassessment of the Habitat and Ecology ofthe Ibex (Capra ibex). In: R.C.G.M. Lauwerier, I.Plug (eds) The Future from the Past: Archaeozool -ogy in Wildlife Conservation and Heritage Manage -

ment. Oxbow Books, Oxford, 6478.RUNNELS C. 1996. The Palaeo lithic and Mesolithic

remains. In: B. Wells, C. Runnels (eds) The Berbati-

Limnes Archaeological Survey 19881990. SwedishInstitute in Athens, Stockholm, 2325.SHACKLE TON J.C. (with DEITH M. R, SHACKLE -

TON N. J.). 1988.Marine molluscan remains fromFranchthi Cave. Indiana University Press, Bloom-ington, IN.

SITLIVY V., SOBCZYK K., KARKANAS P., KOU-MOUZELIS M. 2007. Middle Paleolithic lithic as-semblages of the Klissoura Cave, Peloponnesus,Greece: A comparative analysis.Archaeology, Eth -nology and Anthropology of Eurasia 3(31), 215.

STARKOVICH B. 2011. Subsistence Change during

the Middle to Upper Paleolithic at Klissoura Cave 1(Peloponnese, Greece). Ph.D. Dis serta tion, School ofAnthropology, University of Arizona, Tucson, AZ.



13/13

STEPHENS D.W., KREBS J.R. 1986. Foraging The-ory. Princeton University Press, Princeton, NJ.

STINER M.C. 2001. Thirty years on the Broad Spec -trum Revolution and Palaeolithic demography.

Proceed ings of the Na tional Acad emy of Sci ences

98(13), 69936996.STINER M.C., MUNRO N.D. (n.d.). On the Evolution

of Diet and Landscape dur ing the Upper Paleolithicthrough Mesolithic at Franchthi Cave (Peloponnese,Greece).Journal of Human Evolution (in press).

STINER M.C., MUNRO N.D., SUROVELL T.A.2000. The tortoise and the hare: Small game use, theBroad Spectrum Revolution, and Palaeolithic de-mography. Current Anthropology 41, 3973.

TCHERNOV E. 1992. Evo lution of complex ities,exploitatation of the biosphere and zooarchaeology.

Archaeozoologica V(1), 942.

TOMEK T., BOCHESKI Z.M. 2002. Bird scrapsfrom a Greek table: the case of Klissoura Cave.Acta

Zoologica Cracoviensia 45, 133138.VAN ANDEL T.H., ZANGGER E., PERISSORATIS

C. 1990. Quaternary transgressive/regressive cyclesin the Gulf of Argos, Greece. Quaternary Research34, 317329.

WRIGHT J.D., SHERIDAN R.E., MILLER K.G.,UPTEGROVE J., CRAMER B.S., BROWNINGJ.V. 2009. Late Pleistocene Sea level on the NewJersey Margin: Implications to eustasy and deep-seatemperature. Global and Plan etary Change 66, 93

99.

Notes

1. The ages of the cultural layers differ; the early UP inKlissoura 1 is older, and there is a major gap in cul-tural ma te rial between 2716 kyrs BP at Klissoura1; Franchthi is particularly rich in deposits contain-ing Gravettoid, Final Paleo- lithic and Mesolithicindustries (Perls, 1987; Farrand, 2000).

2. Payne mentions hav ing found a few fal low deerbones in Franchthi Cave, but this diag nosis remainsunclear.

3. Data on a variety of higher latitude archaeo- faunasin con tinental Eu rope (e.g., Berke, 1984; Jochim,1988) indicate related but significantly later shifts indietary breadth.


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