Eur asian Pre his tory, 7 (2): 287–308.

SHELL OR NA MENTS FROM THE UP PER PALEOLITHICTHROUGH MESOLITHIC LAY ERS OF KLISSOURA CAVE 1

BY PROSYMNA (PELOPONESE, GREECE)

Mary C. Stiner

School of An thro pol ogy, P.O. Box 210030, Uni ver sity of Ar i zona, Tuc son, Ar i zona 85721-0030, USA;mstiner@email.ar i zona.edu

Ab stractMore than 1500 shell or na ments were re cov ered dur ing the ex ca va tions of the early Up per Paleolithic through Meso-

lithic lay ers of Klissoura Cave 1. The or na ment as sem blages from the mid dle and lower Aurig na cian and the ear li est Up -per Paleolithic (Uluzzian) lay ers as so ci ate with well pre served hearths and other in tact cul tural fea tures. The older or na -ment as sem blages are ex cep tion ally rich in mol lusk spe cies, whose shells hu mans col lected from ma rine shore lines, fresh -wa ter hab i tats and Plio cene fos sil sources. A par tic u larly dense con cen tra tion of or na men tal shells oc curs within the areaof a small struc ture in the lower Aurig na cian (layer IV). Tax o nomic di ver sity in the or na ment as sem blages de clines pre -cip i tously af ter the for ma tion of layer IIIe–g (up per Aurig na cian), and this con di tion of low di ver sity per sists through theend of the cul tural se quence. The changes in or na ment di ver sity seem to re flect nat u ral changes in coast line hab i tat struc -tures of the re gion. All of the Up per Paleolithic or na ment as sem blages are “high-graded” or se lec tively win nowed for har -mony in color, form and qual ity. There are few if any hints of man u fac tur ing er rors and de bris typ i cal of shell or na ment as -sem blages in coastal sites. Rather, the or na ments dis play high fre quen cies of use wear (pol ish), usu ally in a pre ferred ori -en ta tion, in di cat ing that most of them ar rived on site while af fixed to hu man bod ies or or ganic ar ti facts. There are no re -mains of ed ible ma rine mol lusks in Klissoura Cave 1, con sis tent with its in land lo ca tion. The tax o nomic com po si tion ofthe early Up per Paleolithic shell as sem blages is sim i lar to those doc u mented in It aly, whereas the very lim ited tax o nomiccom po si tion of the later or na ment as sem blages is most con sis tent with those found at Franchthi Cave on the south ernArgolid.

Key words: Paleolithic shell or na ments, Aurig na cian, Uluzzian, taphonomy, site for ma tion pro cesses, site struc ture.

IN TRO DUC TION

Com par a tively lit tle is known about Paleo-lithic or na ments from Greece. The num ber of ex -ca vated Paleolithic sites has climbed slowly overthe last five de cades, and only a few of these sites– Franchthi Cave and now Klissoura Cave 1 –pro vide much in for ma tion about early dec o ra tivetra di tions and the con texts of their use. Paleolithic art, in clud ing beads, is con sid ered an im por tantcri te rion for the emer gence of mod ern hu man be -hav ior dur ing the Late Pleis to cene (e.g., Klein,1989; Mellars, 1989; White, 2003).

Dec o ra tive tra di tions ex ist in vir tu ally ev eryre cent hu man cul ture, so much so that early Pale-olithic beads and other or na ments might seem or -

di nary and with out in ter est. The be hav ioral im -por tance of the emer gence of bead-mak ing tra di -tions in the early Up per Paleolithic is un der-scored, how ever, by the dearth of them in ar chae -o log i cal sites prior to this pe riod in much ofEurasia (but see d’Errico et al., 1998; Bar-YosefMayer et al., 2009) in con trast to some Af ri canMid dle Stone Age sites (Bouzouggar et al., 2007;Henshilwood et al., 2004).

Or na ments made from mol lusk shell, os trichegg shell, mam mal teeth and ivory, and soft stoneare con sid ered to be a unique evo lu tion ary de vel -op ment on ac count of their vi su ally strik ing qual i -ties, trans fer abil ity among per sons (sensu du ra bleitems of trade, gifts or burial par a pher na lia) (Kuhnand Stiner, 2007), and the stylistic formalizations

ap par ent from con sis ten cies in size and ap pear -ance (Hahn, 1972; Bar-Yosef, 1989; White, 1989, 1993; Tab or in, 1993; Stiner, 1999, 2003; d’Errico et al., 2001; Kuhn et al., 2001; Bouzouggar et al.,2007). Min eral ox ides, on the other hand, werewidely used as early as the Mid dle Paleolithic pe -riod in Eur asia, per haps to color hides, hu manskin, or tools. While dec o ra tion is only one ofsev eral pos si ble ex pla na tions for the pres ence ofpig ments in Mid dle Paleolithic toolkits, the pos si -bil ity of such prac tices in the Mid dle Paleolithiccan not be re futed with cur rent ev i dence. Even ifochre and other min er als were used as pig ments in the ear lier pe ri ods, ar chae ol o gists’ ef forts to linktheir oc cur rence in sites to pre his toric dec o ra tivetra di tions re mains con tro ver sial (but see Henshil-wood et al., 2009). The in for ma tion that paintedbody de signs may carry is tran sient and non-trans fer ra ble (Kuhn and Stiner, 2007), in con trastto the du ra ble prop er ties of some or na ments. Ev i -dence for de sign rep e ti tion is nec es sary to any ar -gu ment for shared sym bolic sig nif i cance in pre -his toric dec o ra tive ob jects (Kuhn and Stiner,2007). It is likely that or na ment pro duc tion anduse was great est where in ter ac tion among so cialgroups was high, pos si bly in con nec tion with re -gion ally high hu man pop u la tion den si ties (Kuhnet al., 2001). The sud den ap pear ance of du ra bleor na men tal ob jects in hu man cul tures of the LatePlei- stocene may even sug gest the emer gence ofa ba sic vi sual gram mar in so cial in ter ac tions (e.g., Gam ble, 1986; d’Errico et al., 2003; Kuhn andStiner, 2007).

The tim ing of the first ap pear ance and sub se -quent pro lif er a tion of or na ment tra di tions var iesacross world re gions. An im por tant ques tion forthis study is the chro nol ogy and evo lu tion of or -na ment tra di tions in south ern Greece. Few paleo-an thro pol o gists would be sur prised to learn thator na ments ap pear sud denly in the ar chae o log i calre cord of Greece with be gin ning of the Up perPaleolithic cul ture pe riod. Most early Up perPaleolithic in dus tries in Greece are at trib uted tothe Aurig na cian (Run nels, 1996) and have beeniden ti fied in cave sites such as Franchthi (PerlÀs,1987), Kephalari, Klissoura 1 by Prosymna(Koumouzelis et al., 1996, 2001), Asprochaliko(Bailey et al., 1983), and most re cently at Lakonis (Panagopoulou et al., 2002–2004). Early Up perPaleolithic in dus tries that re sem ble the Uluzzian

of It aly (Palma di Cesnola, 1966, 1993; Benini etal., 1997; Gambassini, 1997) have been found inKlissoura Cave 1 (Koumouzelis et al., 2001;Kaczanowska et al., this is sue). Sev eral Up perPaleolithic sites in Greece con tain or na mentsmade from shells (e.g., Shackle ton, 1988; Kou-mouzelis et al., 2001). Re cent ex ca va tions atKlissoura Cave 1 have yielded ex cep tion ally richor na ment tra di tions in the early Up per Paleolithicof Klissoura Cave 1, in clud ing the Uluzzian layer.

BACK GROUND TO KLISSOURACAVE 1

More than 1500 shell or na ments were re cov -ered dur ing the ex ca va tions of the early Up perPaleolithic through Mesolithic lay ers of Klissoura Cave 1. The or na ments from the early Up perPaleolithic lay ers (V–IIIc) oc cur within andamong a dense ar ray of well pre served sed i men -tary fea tures, mainly hearths. Shell or na ments aremost abun dant in the mid dle to lower Aurig na cian lay ers, par tic u larly layer IV.

Sev eral gen eral ob ser va tions about KlissouraCave 1 are im por tant to this pre sen ta tion of theshell or na ments. This shal low cave is sit u ated onthe Berbatias River and com mands a wide view of the Argos plain, roughly 12 km in land of theArgolikos Gulf (Fig. 1). Most of the or na men talshells none the less were col lected from ma rineshores. The great va ri ety of fea tures, ar ti facts andver te brate fau nal re mains in the Up per Paleolithic lay ers in di cates that the site served mainly at ares i den tial base, par tic u larly in the early phases of oc cu pa tion. An area of 13–15 square me ters wasex ca vated (Karkanas et al., 2004; Koumouzelis et al., 2001), and the units cut through the heart ofsev eral oc cu pa tional lay ers in this small cave.

The cul tural stra tig ra phy in Klissoura Cave 1is more than 5 m deep and in cludes a long Mid dlePaleolithic se quence (Sitlivy et al., 2007) cappedby Up per Paleolithic and Mesolithic ho ri zons(Karkanas, this is sue). Apart from lo cally dis -turbed zones at the in ter face of the Mid dle andUp per Paleolithic (VII/VI), or na ments are con -fined to the Up per Paleolithic (V, IV, III–III” &6a), Epipaleolithic (II) and Mesolithic lay ers(5–3a) (Ta ble 1). In the early Up per Paleolithiclay ers, hearth fea tures oc cur in more than one-third of all ex ca va tion units. The Aurig na cian lay -

288 M. C. Stiner

ers are unique not only for their high con cen tra -tions of wood ash and in tact hearths, but the factthat many were lined with clay (Karkanas et al.,2004; Pawlikowski et al., 2000). Burn ing dam age is com mon on the bones, or na ments, and lithicsfrom all of the lay ers (Koumouzelis et al., 2001;Tomek and Bocheñski, 2002; Karkanas et al.,2004; Starkovich and Stiner, this is sue).

Vari a tion in or na ment abun dance in the Up -per Paleolithic through Mesolithic lay ers is notex plained by dif fer ences in the thick ness of theex ca vated de pos its. Fol low ing Karkanas (this is -sue), the Mesolithic and prob a ble Epigravettian(IIa–d) ho ri zons are 10–20 cm in thick ness, ex -cept where large pits in vade the older lay ers. TheUp per Paleolithic se quence (III–V) is much

Shell or na ments from the Upper Paleolithic 289

Fig. 1. (Left) Lo ca tion of Klissoura Cave 1 by Prosymna on the Argolid of Peloponnese, Greece; other sitesshown are Franchthi Cave (2) and Kephalari Cave (3). (Right) Com par i son of mod ern shore lines to the dis tri bu tionof paleolakes (black fill) and shore lines at the Last Gla cial Max i mum in south ern Greece (adapted from Pe tit-Maire et al., 2005). White fill rep re sents ex posed land sur faces at the time of the LGM, which are now in un dated

Ta ble 1Layer terms, cul tural at tri bu tions, and geo log i cal se quences for the or na ment sam ples from Klissoura

Cave 1

Cul ture at tri bu tion Layer terms Geo log i cal se quence Or na ment NISP

Mesolithic 3, 5a A 5

Epigravettian IIa, IIb, IId B 9

Dis turbed zone 6, 6a, 6/7 B 8

Medit. Backed-bladelet in dus try III-III' D 38

Up per Paleolithic (non-Aurig.) III" D 28

Up per Aurig na cian IIIc D 23

Mid-Aurig na cian IIIe-g D 138

Lower Aurig na cian IV E 1218

Ear li est UP (Uluzzian) V F 32

Dis turbed, mostly MP VI (F/)G 28

Mid dle Paleolithic VII G 52

The geo log i cal se quences are layer groups or fa cies with sim i lar sedimentological char ac ter is tics that form co her ent layer sets,sep a rated by dis crete con tacts that in di cate mi nor or ma jor depositional hi a tuses (fol low ing Karkanas, this is sue). The Med i ter ra -nean backed-bladelet in dus tries of lay ers III-III' and III'' dif fer from one an other in some as pects but have in com mon the high fre -quency of the im ple ments.

thicker – about 190–210 cm over all. A dis turbed,mixed de posit was en coun tered in lay ers 6–7 (seeTa ble 1) and will not be dis cussed fur ther.

In the layer III se ries, clay-lined hearths arecom mon only in layer III’. A large space de voidof hearths and large rocks was found in squaresAA-CC1 at the depths of 75–85 cm be low da tum,and a short arc of clus tered rocks was en coun tered be tween 85–105 cm. Sed i ments con tain ing a non- Aurig na cian backed-bladelet in dus try of typ i calMed i ter ra nean type (lay ers III–III’) and an otherlayer (III”) con tain ing a non-Aurig na cian in dus -try oc cur above the Aurig na cian lay ers (IIIb–g,IV) (Kaczanowska et al., this is sue).

The ear li est Aurig na cian layer IV var ies be -tween 25–50 cm in thick ness and is ex cep tion allyrich in bones and ar ti facts. In ad di tion to manyclay-lined and un lined hearths, this layer pre -serves a wide as sort ment of lenses, pits, and theout line of what was prob a bly a small man-madeshel ter. Rare ant ler points and sig nif i cant amounts of what ap pears to be os se ous man u fac tur ing de -bris also oc cur in layer IV (Christidou, per sonalcom mu ni ca tion, 2010; Starkovich and Stiner, thisis sue).

Layer V is a thin, un du lat ing de posit that oc -curs only in the west ern part of the ex ca va tion and is char ac ter ized by many con cave hearth fea tures.A small or na ment as sem blage ac com pa nies thedis tinc tive lithic in dus try (Ta ble 1), whose archedbacked bladelets and com par a tively high pro por -tion of microblades resemble Uluzzian in dus triesin It aly (cf. Palma di Cesnola, 1993; Benini et al.,1997; Gambassini, 1997; Kuhn and Stiner, 1998), war rant ing a com par i son of the Greek and Ital ianor na ment as sem blages.

The first ra dio car bon de ter mi na tions attem-pted for the Up per Paleolithic lay ers sug gestedcom par a tively young ages for the Aurig na cian atKlissoura 1 (Koumouzelis et al., 2001; com pareKoz³owski, 1982, 1992, 1999). Pre lim i narythermo-lu mi nes cence re sults on burnt flint ar ti -facts sug gested that layer V is sig nif i cantly older.New AMS ra dio car bon re sults us ing the ABOXpre treat ment method (Bird et al., 1999) on woodchar coal sam ples from the late Mid dle Paleolithicthrough Up per Paleolithic lay ers per mit some re -vi sions to the Up per Paleolithic chro nol ogy(Pigati et al., 2007; Kuhn et al., this is sue). Still,the dates for the ear li est Aurig na cian layer (IV) –

32,690±110 and 33,150±120 uncali- brated ra dio -car bon years be fore pres ent (BP) – are not nearlyas old as those for many other Aurig na cian casesin Eu rope (e.g., Conard et al., 2003), but they aresome what older than Aurig na cian sites in theLev ant. The Uluzzian-like in dus try in layer V issealed from the layer above by a fine tephra andmay be older than 39,000 cal i brated BP.

GOALS AND METH ODS

This study ad dresses sev eral is sues sur round -ing or na ment use and dis card at Klissoura Cave 1: (a) Spe cies com po si tion as it re lates to ma rine and ter res trial en vi ron men tal con di tions, di ver sity inraw ma te rial sources, and ev i dence for se lec tiv ityby hu mans; (b) dam age pat terns on the shells thatmay re flect raw ma te rial sources, or na ment man -u fac ture, use and dis card prac tices; (c) the spa tialdis tri bu tion of or na ments and dam age phe nom ena in the de pos its, in clud ing as so ci a tions with hearthsand other fea tures; (d) trends in or na ment as sem -blage com po si tion from the early Upper Paleo-lithic through Mesolithic; and, (e) com par i sons ofthe Klissoura 1 or na ment se ries to those fromearly Up per Paleolithic Ital ian sites and to thelater or na ment as sem blages from Franchthi Caveon the south ern Argolid (see Fig. 1). The quan ti ta -tive units used in the anal y ses are the num ber ofiden ti fied spec i mens (NISP) and the min i mumnum ber of in di vid ual an i mals (MNI) rep re sentedby whole and frag men tary re mains. NISP is im -por tant for many of the taphonomic anal y ses,whereas MNI better rep re sents or na ment quan ti -ties from a func tional point of view.

To ad dress ques tions about the con texts of or -na ment use and their pos si ble sig nif i cance indaily life, this study con sid ers both the gen eralproces ses of as sem blage for ma tion and how theor na ments were made, used and ex hausted asitems of tech nol ogy. Spe cies-spe cific ecol o gies of the mol lusks are used to in fer the range of hab i tatsources, and ob ser va tions about shell con di tionand spa tial dis tri bu tions help to de ter mine thecon texts of or na ment use on site. Re con struc tionsof Late Pleis to cene shore lines and other wa terbod ies at the Last Gla cial Max i mum (LGM, ca.20,000 years BP; Lambeck, 1996; Pe tit-Maire etal., 2005) and re cent Ho lo cene pe ri ods de fine two ex tremes in the Pleis to cene shore line and hab i tat

290 M. C. Stiner

con fig u ra tions of south ern Greece (Fig. 1). En vi -ron men tal con di tions on the Argolid dur ing theearly Up per Paleolithic would have fallen some -where within these ex tremes, though closer toLGM con di tions with re spect to land sur face ex -po sure, pen in sula-is land con fig u ra tions, and thedi ver sity of aquatic hab i tats within a 50 km ra dius of the site. The later oc cu pa tions, par tic u larly dur -ing the Mesolithic pe riod, would have experien-ced con di tions more like those of the re cent Ho lo -cene.

RE SULTS ON THE PALEOLITHICOR NA MENT AS SEM BLAGES

All of the or na ments from Klissoura Cave 1were made from aquatic mol lusk shells, mainlysmall ma rine gas tro pod spe cies along with somefresh or brack ish wa ter and fos sil types. A few red deer (Cervus elaphus) ca nines oc cur among thefau nal re mains (Koumouzelis et al., 2001; Star-kovich and Stiner, this is sue) but none of thesewas al tered for sus pen sion. A few per fo rated teeth were noted in the early years of the excavationpro ject but these could not be lo cated or con-firmed

Mol lusk spe cies abun dance and di ver sityThe di ver sity of mol lusk spe cies var ies greatly

among the Up per Paleolithic and Mesolithic lay ersin Klissoura Cave 1 (Ta ble 2 and Ap pen dix 1), but spe cies dis tri bu tions do not vary sig nif i cantlyamong fea tures or ex ca va tion units within each

layer. Spe cies rich ness, here cal cu lated as N-spe -cies/log MNI to cor rect for sam ple size ef fects, isgreat est in lay ers IIIe–g and VI, es pe cially in IVwhere a min i mum of 44 taxa were iden ti fied. Theor na ment as sem blages from the later UP and Epi-paleolithic lay ers are much poorer in spe cies,even af ter cor rect ing for the smaller sam ple sizes,fol lowed by a mild in crease in di ver sity in theMesolithic. Tax o nomic di ver sity is uni formly low in the large Epipaleolithic and Mesolithic or na -ment as sem blages from Franchthi Cave on thesouth east ern mar gin of the Argolikos Gulf (com -pare Shackle ton, 1988; PerlÀs and Vanhaeren,2010).

Steep rocky or het er o ge neous Med i ter ra neancoasts with high nu tri ent turn over tend to sup portmany mol lusk spe cies. Nearby sites con tain ingUp per Paleolithic shell or na ment as sem blagesalso tend to be spe cies-rich (e.g., Riparo Fumaneand Riparo Mochi in north ern It aly; Leonardi,1935; Bartolomei et al., 1994; Fiocchi, 1996-97;Stiner, 1999, 2003). To day Klissoura Cave 1 liesas close as it ever has to the Argolikos Gulf. Thesite would have been a few more ki lo me ters dis -tant from shore at the time of the Aurig na cian oc -cu pa tions, but with out rad i cal al ter ations in shore -line shape or topography (Fig. 1).

The cen tral geo graphic po si tion of the Kli-ssoura Gorge to brack ish la goons, rivers, lakesand ma rine shore lines of the Peloponnese andmain land Greece dur ing gla cial pe ri ods con trib -uted to the tax o nomic va ri ety in the or na ment as -sem blages in lay ers IIIe–g through V. Shells were

Shell or na ments from the Upper Paleolithic 291

Ta ble 2In dexed com par i son of spe cies rich ness for early Up per Paleolithic through Mesolithic shell or na ment

as sem blages by in tact layer from Klissoura Cave 1

Layer group Num ber of spe cies Or na ment MNI Spe cies rich ness

Mesolithic (3, 5a) 5 5 7.15

Epipaleolithic (IIa-d) 3 9 3.14

Med. Backed-bladelet Industry(III-III')

6 38 3.80

Up per Paleolithic (non-Aurig.) (III") 2 23 1.47

Up per Aurig na cian (IIIc) 4 23 2.94

Mid-Aurig na cian (IIIe-g) 18 138 8.41

Lower Aurig na cian (IV) 45 1218 14.58

Early UP/Uluzzian (V) 14 32 9.30

Un de ter mined (VI-VII) 14 53 8.12

Spe cies rich ness is cal cu lated as N-spe cies/logMNI.

col lected from the wide range of ma rine and in -land hab i tats dur ing these oc cu pa tion phases (Ta -ble 3 and Ap pen dix 2). At least 8% of the or na -men tal shells from layer IV are fresh wa ter spe cies (Theodoxus), and an other 4% are fos sils fromPlio cene lake bed de pos its (Fig. 2; Magrograssi,1928; G. Manganelli and A. C. Colonese, per -sonal com mu ni ca tion, 2007). More than half of

the shells orig i nate from Late Pleis to cene brack -ish la goons or es tu ar ies (Cyclope), and the re -main ing 35% are from the ma rine lit to ral. Otherfac tors that may have con trib uted to spe cies rich -ness in the older or na ment as sem blages were thegreater in ten sity or du ra tion of oc cu pa tion and the greater diversity of ac tiv i ties on site (see be low).

292 M. C. Stiner

Ta ble 3Rep re sen ta tion of hab i tat sources and trophic cat e go ries in shell or na ments from layer IV,

the most di verse as sem blage

By hab i tat type % By trophic group %

Fos sil source 4 Car ni vore 10

Ma rine mixed shore 35 Car ni vore-scav en ger 42

Estuarine-lagunal 52 Om ni vore 6

Fresh-brack ish 8 Her bi vore-detritivore 40

Fig. 2. Fos sil taxa from Plio cene lake de pos its of the gen era Corymbina (prob a bly C. rhodiensis and C. aegae) andMelanopsis (e.g. M. gorceixi), fol low ing Magrograssi (1928). Im age com piled from pho to graphs by G. Hartrman

Ta ble 4Sum mary of rel a tive abun dances of com mon gen era (% of to tal MNI) in the or na ment as sem blages

from Klissoura Cave 1

Layer(s): 3, 5a IIa-d 6-6/7 III-III' III" IIIc IIIe-g IV V VI-VII

Taxon % of to tal MNI

Dentalium spp. 20 44 0 2 0 0 1 1 29 4

Gibbula albida 0 0 0 0 0 0 0 4 0 0

Gibbula spp. 0 0 0 0 0 0 8 13 3 15

Clanculus spp. 0 0 0 0 0 0 0 6 0 11

Monodonta spp. 0 0 0 0 0 0 2 2 0 2

Homalopoma sp. 0 0 0 0 0 0 3 9 3 6

Columbella rustica 0 0 0 13 30 22 25 7 9 6

Mitrella scripta 0 0 0 0 0 0 0 < 0 0

Cyclope spp. 20 45 87 81 70 61 46 35 34 30

Hinia reticulata 0 0 0 0 0 0 2 7 3 11

Theodoxus spp. 0 0 13 0 0 17 6 8 4 9

Fos sil lake spp. 0 0 0 0 0 0 1 3 3 2

Bi valve spp. 60 11 0 4 0 0 2 1 9 4

All other taxa 0 0 0 0 0 0 4 3 3 0

Sam ple size (MNI) 5** 9** 8** 38 23 23 138 1218 32 53

(**) Sam ple size is very small, mak ing per cent age cal cu la tions sus pect.

The or na ment as sem blage from layer V isalso spe cies-rich for its size, but the prom i nenceof Dentalium (tusk) shells sets it apart (Ta ble 4).The layer V as sem blage is small, how ever, and itshould be kept in mind that one dis tinc tive beadstrand would be enough to al ter the char ac ter ofthis small as sem blage. In ter est ingly, Dentaliumshells are also com mon in the Uluzzian ho ri zonsof Grotta del Ca val lo in It aly (Palma di Cesnola,1966).

Small quan ti ties of or na ments also oc cur inlay ers VI and VII of Klissoura 1. These must becon sid ered sep a rately, since a mix ture of Up perPaleolithic and (mostly) Mous terian ar ti facts oc -

curs in these lay ers, along with few fau nal re -mains (Starkovich and Stiner, this is sue). The spe -cies con tent of the or na ment as sem blage fromVI–VII best re sem bles that of layer IV (Fig. 3),which lies in di rect con tact with layer VI hor i zon -tal units in the area where most of the or na mentswere found (squares AA1-2 at 175–185 cm be low da tum). This and other ev i dence (K. Douka, per -sonal com mu ni ca tion, Feb ru ary 2010) in di catesthat the few or na ments that lo cally co-oc cur inMid dle Paleolithic ar ti facts in layers VI and VIIoriginated from layer IV.

The struc ture of the Peloponnese land scapedur ing the Late Pleis to cene and early Ho lo cene is

Shell or na ments from the Upper Paleolithic 293

Fig. 3. Rel a tive fre quen cies of the ma jor or na ment gen era in the Up per Paleolithic lay ers of Klissoura Cave 1

im por tant to un der stand ing or na ment raw ma te -rial sources as well as the po ten tial for ag ing op -por tu ni ties for hu mans of the pe riod. The Gulf ofKorinthos, Argolid Pen in sula, and Cyclade Is -lands have been the fo cus of sev eral shore line re -con struc tions (van Andel and Sutton, 1987;Shackle ton, 1988; Lambeck, 1996; van Andel and Tzedakis, 1996). These re con struc tions are in -valu able for es ti mat ing the dis tance be tween siteand coast at the time of oc cu pa tion, the com plex -ity of coastal and in land aquatic hab i tats, and thehet er o ge ne ity of the en vi ron ment in suc ces sivelylarger catch ments around the site. The most re cent re con struc tion by Lambeck (1996) in te gratesmod els of eustacy and glacio-hy dro isos tasy withtop o graphic data for tec toni cally sta ble ar eas toes ti mate net changes in land ex po sure. Ground-truthing was ac com plished in this and prior stud -ies partly by ref er ence to dated ar chae o log i cal and paleontological de pos its. Lay ers III and be low inKlissoura Cave 1 would have formed dur ing in -ter me di ate gla cial con di tions, and layer II and theMesolithic lay ers would have formed around thetime of the Pleis to cene–Ho lo cene tem po ralbound ary. The older or na ment as sem blages pre-date the Last Gla cial Max i mum by many thou -sands of years, but ac cord ing to Lambeck (1996:596), [dur ing] “…much of the time be tween about 70,000 years b.p. and the Gla cial Max i mum, icevol umes sig nif i cantly exceeded those of todaysuch that the global sea-levels did not rise above40–50 m below present level during this interval.”

Land scapes to the north and east of the Kli-ssoura Gorge had greater land mass ex po sure dur -ing the early Up per Paleolithic, in ter spersed lo -cally with marshes and es tu ar ies and at least twolarge fresh wa ter lakes to the north east. The land -scape and ecol ogy of this pe riod there fore wasmore com plex than the mod ern one, and the ter -res trial eco sys tems would have been more pro -duc tive. The cen tral lo ca tion of the gorge and itssmall caves to di verse hab i tats was a ma jor at trac -tion to Pleis to cene for ag ers in the eastern Pelo-ponnese.

The rapid de cline in spe cies rich ness in theshell or na ments af ter layer IIIe–g but within theAurig na cian pe riod may be ex plained by thegrowth and even tual dom i nance of lagunal hab i -tats on the west ern shores of the Argolid (seeShackle ton, 1988). While changes in cul tur ally

bound aes thet ics may also have con trib uted to thenar rowed range of mol lusk spe cies used for or na -ment-mak ing in lay ers IIIc–d and af ter, changesin nat u ral bi otic di ver sity were the pri mary con -straint on hu man choices and form the base lineagainst which variety must be interpreted.

Raw ma te rial sources and ev i dence for hu man se lec tiv ity

The types of or na men tal shells in Klissoura 1are not the most com mon spe cies on Med i ter ra -nean shores to day or in the past. “Un nat u ral” bi -ases in tax o nomic com po si tion in clude com par a -tively high fre quen cies of car niv o rous spe cies,and ar ti fi cially nar row shell color, shape and sizedis tri bu tions. The early Up per Paleolithic oc cu -pants clearly pre ferred shells of the gen era, Gib-bula, Clanculus, Homalopoma, Columbella, Cy-clope, Hinia, and Theodoxus (Ta ble 4). The pro -por tions of Columbella and Cyclope in crease with time and these two gen era ul ti mately dom i nate the youn ger Up per Paleolithic as sem blages (Fig. 3).Gibbula albida was im por tant only dur ing the for -ma tion of layer IV, and most Clanculus shellsalso oc cur there. Dentalium shells are com mon inlayer V, as noted pre vi ously by Koumouzelis etal. (2001), and they re gain im por tance only dur -ing the Epipaleolithic and Mesolithic.

Car niv o rous spe cies (pred a tors and scav en -gers) in the or na ment as sem blages are rep re sented well in ex cess of the ex pected en coun ter rate inliv ing mol lusk com mu ni ties. Global cen suses ofmol lusk com mu ni ties (e.g., Sabelli, 1980) andcol lect ing ex per i ments by the au thor on east ernMed i ter ra nean shores (Stiner, 1999) sug gest av er -age en coun ter rates for car ni vores at roughly 15% of all gas tro pod in di vid u als in beach-cast ma te -rial. In Aurig na cian layer IV, spe cial ized car ni -vores oc cur at a rate of 10% but to gether with car -niv o rous scav en gers con sti tute 52% of shell MNI(Ta ble 4). The pat tern of trophic rep re sen ta tion inthe or na ments is the op po site of that for nat u ralmol lusk com mu ni ties and must re flect strong se -lec tiv ity on the part of Paleolithic hu mans. Shellcol lec tors prob a bly were re spond ing to vi su allyat trac tive properties of the shells coupled with asense of their ecological rarity (Stiner, 2003).

Dam age pat terns on the shells in di cate thatmost or all of the spec i mens were col lected asempty shells from ma rine and es tu ary beaches,

294 M. C. Stiner

river banks and mouths, and fos sil beds. Wave-in -duced abra sion is wide spread on the ma rine shells and on some of the fos sil types (Ta ble 4b). Abra -sion dam age is least com mon on fresh and brack -ish wa ter spe cies and prob a bly re flects the loweren ergy lev els of the aquatic hab i tats where theseshells were ob tained. Abra sion dam age and bore-holes from molluskan pred a tors oc cur on some ofthe shells from Klissoura 1 (0–8% of shells), alsocon sis tent with col lec tion from wave-cast beachsources. There are no large, ed ible ma rine mol -lusk re mains in any of the layers of KlissouraCave 1.

Hu man se lec tiv ity is ap par ent from the nar -row range of shell shapes and sizes in the Klisso-ura or na ments. This pat tern holds true for all ofthe shell as sem blages. Small spe cies were stronglypre ferred (Fig. 4, Ta ble 5b), with the mean lengthfor all mea sur able (whole) shells in the Up perPaleolithic lay ers be tween 1.3–1.4 cm, and be -tween 1.4–1.7 cm in the youn ger lay ers. The full

size range is 0.6 to 5.5 cm, but nearly all of theshells are un der 2 cm in length. The con sis tency in or na ment size within this range is ex plained bythe hu mans’ nar row pref er ence for cer tain spe -cies. Round, oval or bas ket-like forms were thenorm in the Klissoura as sem blages (Fig. 5), alongwith Dentalium tubes. Sim i lar mean sizes andranges are re ported for other Up per Paleolithicsites in Eu rope (e.g., White,1989; Stiner, 2003),in clud ing or na ments from the Périgord re gion ofFrance that were la bo ri ously carved from ivoryand stone (White, 1989: 382).

Red and pink shells were sought out pref er en -tially in ad di tion to those with bright opal es centhues or con trast ing stripes. Ochre stained shellsoc cur through out the Up per Paleolithic lay ers(Ta ble 6a), but ochre ap pli ca tion cen tered on twogen era: Clanculus, a group with nat u rally pink orred shells and rough, knobby sur faces; and Theo-doxus, a group with bright white or viv idly striped shells and smooth sur faces (Ta ble 6). Ochre traces

Shell or na ments from the Upper Paleolithic 295

Fig. 4. Size dis tri bu tion (cm) of mea sured or na ments, ex em pli fied by the shells from lay ers IV and V of Klis-soura Cave 1

Fig. 5. Typ i cal shell forms and per fo ra tions made by hu mans.The holes were made by a sim ple punch ing tech -nique, ex cept for tusk shells (Dentalium) which were snapped to cre ate tube beads. Shell taxa from left to right areHomalopoma sanguineus, Nucella lapillus, Cyclope neritea, Dentalium sp. Theodoxus spp (2 spec i mens, one withstripes), Monodonta sp., Columbella rustica, and Clanculus corallinus. Im age com piled from pho to graphs by G.Hartman

oc cur most of ten on Clanculus shells, at about 3times the rate ob served for Theodoxus. Ochretraces were found on sev eral other shell types but

at much lower fre quen cies. Though a rough sur -face might gen er ally in crease the chances ofochre traces be ing pre served, ochre is also fre -quently pre served on smooth shells in the Klisso-ura 1 as sem blages, so the bias is not from dif fer -en tial pres er va tion of ap plied pig ment. Red dishhues were val ued, and peo ple went to some ef fortto make naturally reddish (but faded) shellsredder.

Con texts of or na ment use and dis cardShell frag men ta tion is mea sured in this study

by an in dex of “com plete ness." The in dex is cal -cu lated by di vid ing the num ber of in di vid u als bythe to tal num ber of iden ti fied frag ments in eachtax o nomic group and layer (MNI/NISP, Ta ble5b). Nearly all of the gas tro pod shells from Kli-ssoura 1 are com plete or nearly com plete (89–

296 M. C. Stiner

Ta ble 5Dam age and shell size sta tis tics for the larger as sem blages of Up per Paleolithic or na ments by layer

from Klissoura Cave 1

a. burn ing dam age, per fo ra tion rates, breaks through per fo ra tion point, cord wear and the in ci denceof ochre traces

Layer groupBurned

%Per fo rated

%Hole bro ken

%With cord-wear

%With ochre

%

Non-Aurig. UP in dus tries (III-III") 51 97 14 69 3

Up per Aurig na cian (IIIc) 58 92 8 50 8

Mid-Aurig na cian (IIIe-g) 55 92 26 38 3

Lower Aurig na cian (VI) 18 90 21 18 6

Early UP/Uluzzian (V) 48 96 17 7 0

Un de ter mined (VI-VII) 4 99 32 47 4

b. com plete ness, size sta tis tics, and non-cul tural types of dam age

Layer group Com plete-nessWith nat u ral color

%Mean length

(cm)Sd

(cm)Wave-worn

%Pre dated

%

Non-Aurig. UP In dus tries (III-III") 0.97 11 1.43 0.26 23 6

Up per Aurig na cian (IIIc) 1 17 1.47 0.3 17 8

Mid-Aurig na cian (IIIe-g) 0.91 8 1.45 0.6 52 2

Lower Aurig na cian (VI) 0.94 10 1.28 0.44 35 4

Early UP/Uluzzian (V) 0.93 3 1.28 0.55 31 3

Un de ter mined (VI-VII) 0.94 13 1.27 0.29 49 0

Notes: Only the larger sam ples from the III se ries are be low are con sid ered. Per fo ra tion re fers to holes made by hu mans; hole bro -ken re fers to spec i mens that were bro ken through the per fo ra tion point on the shell, ei ther dur ing man u fac ture or (more com -monly) from long-term use; cord wear re fers to pol ish on some or all edges of the per fo ra tion but may also in clude pol ish on theouter sur face near the per fo ra tion. Shell com plete ness is cal cu lated as MNI/NISP. Length mea sure ments are only for those shellswhose nat u ral di men sion could be de ter mined. Pre dated shells have holes drilled in them by a mol lusk pred a tor. Nat u ral color re -fers to the re ten tion of nat u ral pig ment within the shell.

Ta ble 6Dis tri bu tion of ochre traces by mol lusk ge nusin all of the Up per Paleolithic lay ers com bined

Ge nus % with red ochre

Clanculus 29

Columbella 1

Cyclope 5

Gibbula 2

Homalopoma 4

Hinia 7

Theodoxus 11

All other gen era 3

100%, de pend ing on the spe cies), ex cept for frac -tures at the per fo ra tion point. Dentalium shellswere usu ally sec tioned to make tube beads. Rarebi valves (Pecten and Chlamys) were nearly al -ways bro ken, prob a bly be cause they are large,frag ile and prone to tram pling.

A sig nif i cant num ber of the or na men tal shells are burned (Ta ble 5a). Most or all of this dam ageap pears to have been ac ci den tal. There is muchev i dence of hearth re build ing, re newal, and su -per po si tion in the Up per Paleolithic lay ers (Kou-mouzelis et al., 2001; Karkanas et al., 2004),caus ing older de bris to be dam aged by the heatfrom su per im posed hearths (see Stiner et al.,1995). Burn ing dam age is least com mon on theor na ment shells from layer IV, VI and VII (Ta ble5a). Most of the or na ments in layer IV were found within the per im e ter of the man-made structureand immediately below it.

The great ma jor ity of the shells in each as -sem blage have holes in their flanges or whorls(90–95%, Ta ble 5a), and there is lit tle if any waste ma te rial on site. The holes in gas tro pod shellsusu ally were made by hu mans us ing a punch ingtech nique that pro duced small round open ingswith rough edges. Some of the ma rine shells haveholes caused by wave ac tion or drill ing by mollus- kan pred a tors. Hu mans some times took ad van -tage of these “nat u ral” holes if lo cated in the shellflange or dor sally (op po site) to the flange. Wave-worn and pred a tor-drilled holes tend to be dis trib -uted more ran domly on shell sur faces, and thosedrilled by molluskan pred a tors are al most per -fectly round and bev eled in con trast to man-madeholes (Figs 7 and 8). Holes caused by wave ac tion are as abraded and thinned along with the rest ofthe shell, in con trast to lo cal ized pol ishes re sult -ing from wear against organic fibers or hide.

Gibbula albida shells were treated dif fer ently from other types of shells in that fewer than halfof the spec i mens were per fo rated. This is a com -par a tively large top shell with an el e gant py ram i -dal form and pro nounced con cen tric ridges. G.albida shells are com mon only in layer IV (MNI = 71). They con cen trated be tween the depths of150–170 cm be low da tum, mainly in squaresAA1-AA2 (a few also oc cur in BB1-BB2) andthere fore co in cide with the in te rior of the man-made shelter (see below).

The gen er ally high per fo ra tion rate for the

shell or na ments from Klissoura 1 over all lies atone ex treme in the con tin uum of vari a tion ob -served among Up per Paleolithic cases that con -tain or na men tal shells. At coastal sites, such asRiparo Mochi (Liguria, It aly; Stiner, 1999) andÜçaÈÏzlÏ I Cave (Hatay, Tur key; Stiner, 2003;Kuhn et al., 2009), per fo ra tion rates are lower and man u fac tur ing er rors more ap par ent in shell or na -ment as sem blages, along with ev i dence forstock-pil ing or raw ma te rial and ac cu mu la tion ofre jected pieces.

Cord wear was iden ti fied on the shell spec i -mens from Klissoura 1 with the aid of a hand lens, and thus the per cent ages in Ta ble 5a rep re sentmin i mum es ti mates of oc cur rence. Cord-wear in -side the per fo ra tions tends to be asym met ri cal,sug gest ing that the shells were fas tened or strungin one po si tion for long pe ri ods. Sur face pol ish on the outer sur face near the per fo ra tions ac com pa -nies the ev i dence of cord-wear on some of theshells (Figs 6, 7). The in ci dence of cord wear at

Shell or na ments from the Upper Paleolithic 297

Fig. 6. Ex am ple of cord wear in hu man-made holeand outer sur face of Theodoxus shell. Photo by G.Hartman

Fig. 7. Com par i son of holes made by hu mans (1),and an other by molluskan pred a tors (2) of theNaticidae or Muricidae fam i lies

Klissoura is high in com par i son to the coastal siteof Riparo Mochi in It aly.

About one fifth of the spec i mens from Klisso- ura 1 Cave were bro ken through the per fo ra tionpoint (Ta ble 5a, hole bro ken 8–32%). The fre -quen cies of breaks at the per fo ra tion is pos i tivelycor re lated with the in ci dence of cord pol ish(.05>>p>>.02), prob a bly be cause both of thesephe nom ena re late to or na ments be com ing wornout from ex tended use. The prev a lence of cordwear, the high per fo ra tion rates, and the near ab -sence of man u fac tur ing waste ar gue against the or -na ments hav ing been man u fac tured on site. Thereis, as well, an ap pre cia ble win now ing or “highgrad ing” ef fect ap par ent in the or na ment as sem -blages. All told, the or na ments are gen er ally com -plete, at trac tive spec i mens with a nar row size dis -tri bu tion and con sid er able ev i dence long-term use.

Spa tial dis tri bu tions of the or na mentsOr na ments are abun dant in layer IV and es pe -

cially within the do main of the hy poth e sizedman-made struc ture. This fea ture spans squaresAA1-2 of the ex ca va tion (Ta ble 7), where sim pli -fied site plan draw ings (Fig. 8) re veal the mu tu -ally ex clu sive dis tri bu tions of large lime stonerocks and hearths at 150–175 cm be low da tum.The struc ture is de fined by the cir cu lar jum ble oflarge rocks, min i mally 2 m in di am e ter, and a com -plete ab sence of hearths. Doz ens of hearth fea turesen cir cle this area. The rocks may orig i nally haveweighted the edges an or ganic cov er ing, and rolled in wardly upon re moval or de com po si tion of thecover. Dark stained sed i ments were ob served at155–165 cm be low da tum and may be the rem -nants of a floor cov er ing. Given that the or na ments clus ter within the shel ter area to a large de gree,and many dis play cord wear from pro longed use,it is pos si ble that they were once at tached to an or -ganic ob ject such as a large hide or matt. The con -cen tra tion of or na ments within the struc ture alsoex plains the lower in ci dence of burned spec i mens in layer IV in com par i son to the or na ment fromthe other lay ers (Ta ble 5a). The only other shellas sem blage that dis plays a low burn ing rate isfrom lay ers VII–VI, im me di ately be low the shel -ter fea ture, and where layer V does not ex tendhorizontally. Small num bers of or na ments ex tenddown ward into lay ers VII–VI at one edge of theshel ter fea ture area of layer IV (Ta ble 7).

Layer V is com par a tively thin, un even andrid dled with hearth de pres sions. About 42% ofthe layer V or na ments oc cur within hearths, andmany of these or na ments are burned (Ta ble 5a).

The fre quent pres ence of orig i nal (bi o log i cal) pig ment in the shells is note wor thy (nat u ral colorvari able in Ta ble 5b) and sug gests a fa vor ablepres er va tion en vi ron ment. Ver ti cally, the fre -quency of this phe nom e non is high est at 85–115and 140–160 cm be low da tum. The re ten tion ofnat u ral col or ation in shells is strongly and pos i -tively cor re lated to the pres ence of cord wearthrough out the cul tural lay ers (p = 0.0001). Itseems that the lon ger an or na ment re mainedstrung or at tached to an other ob ject, the better thechance that its more tran sient nat u ral prop er tieswould also be pre served. This re la tion sug geststhe ex is tence of one or more sta ble microenviron-ments within the de pos its that en hanced shellpres er va tion. Nei ther nat u ral color re ten tion norcord wear is con cen trated in any par tic u lar square, but the large hor i zon tal (1 × 1 m) units of the ex -ca va tion do not allow us to distinguish randomfrom finely structured distributions in the sedi-ments.

Fi nally, or na ments are rare items in Klissoura Cave 1 if con sid ered against the to tal num ber ofchipped stone ar ti facts in each layer (Ta ble 8). Or -na ments are for mal ar ti facts, how ever, and thus itseems ap pro pri ate to con sider their num bers in re -la tion to chipped stone tools in particular. Or na -ments are com par a tively few even in re la tion tothe num bers of tools, but their pro por tions are no -tice ably greater in lay ers III’ and be low, and es pe -cially in layer IV. These and other ob ser va tionssug gest that the oc cu pa tions were more intenseduring the Aurignacian.

CON CLU SIONS

Pre his toric shell or na ments were mere par ti -cles in larger dec o ra tive for mu lae, and in Paleo-lithic sites they usu ally are found mixed withother cul tural de bris. Lost to ar chae ol o gists inmost in stances are the rules of com bi na tion ofthese ob jects on strings, sur faces or in de lib er ately as sem bled caches. The de gree to which or na -ments were re cruited for com plex sym bolic rep re -sen ta tions is un clear and no doubt var ied greatly.Their ba sic func tion for con vey ing sim ple vi sual

298 M. C. Stiner

mes sages about per sonal state (Kuhn and Stiner,2007) or af fil i a tion (Vanhaeren and d’Errico,2006) may have been nearly uni ver sal, how ever,and likely ex plains why small or na ments are the

most wide spread and com mon art form of thelater Paleolithic in Eurasia and Africa.

Even in a dis as sem bled or scat tered state,Paleolithic or na ments can pro vide in for ma tion

Shell or na ments from the Upper Paleolithic 299

Ta ble 7Num ber of shell or na ments by square at 5-199 cm be low da tum in Klissoura Cave 1

Spit A1 A2 A3 AA1 AA2 AA3 AA4 BB1 BB2 BB3 BB4 CC1 CC2 CC3 Layer(s)

5-20 - - - - - 1 - - - - - - - 1

25 - - - - - 1 - - - 2 - - - -

30 - - - - - - 2 - - 1 - - - -

35 - - - - - - - - - 3 - - - -

40 - - - - 1 1 1 - 1 1 - - - -

45 - - - - - - 3 - - 1 - - 4 1 III'

50 - - - - 1 1 - - 1 - - - 1 1 III'

55 - - - 1 1 - - - 1 - - - 2 - III'

60 - - - - - 1 3 - - - 1 - - - III'

65 - - - - - 1 1 - - - 2 3 - - III'

70 - - - 2 - - - - 2 - 1 - 1 - III"

75 - - - - 1 1 2 2 - 1 - - 1 - III"

80 - - - - - 1 1 - - - - - - - III"/IIIc

85 - - - - - 3 - - - 1 - - 4 2 III"/IIIc

90 - - - 3 2 3 2 - 1 - 1 3 - 2 IIIe-g

95 - - - 3 4 2 2 - 2 - - 1 - 1 IIIe-g

100 - - - 3 3 - 7 - 1 - - 1 1 - IIIe-g

105 - - - - 2 - - - - - 1 2 - - IIIe-g

110 - - - - 2 - - - 11 2 2 2 4 - IIIe-g

115 1 - - 1 2 - 7 - - - 4 - 14 - IIIe-g

120 3 - - 3 1 - 4 - - - - - - - IIIe-g

125 1 1 - - 3 - 1 - - - - - - - IIIe-g

130 - - - 1 - - 1 - - - - - - - IIIe-g

135 - 2 7 2 6 7 4 2 7 5 - 3 2 3 IIIe-g

140 1 4 1 8 6 8 7 8 12 7 1 6 7 5 IIIe-g/IV

145 - 4 2 9 29 - 8 11 6 1 6 5 3 11 IIIe-g/IV

150 4 4 4 10 36 25 19 24 23 14 4 5 21 25 IV

155 7 6 - 21 17 14 16 21 12 15 7 10 36 22 IV

160 3 10 1 20 38 6 3 17 23 14 14 - 15 13 IV

165 4 - 6 55 62 17 5 12 16 16 1 - 12 5 IV

170 12 6 1 53 33 2 1 21 36 3 - 24 22 3 IV

175 - 2 - 27 37 3 3 32 21 1 - 21 9 7 IV/V

180 - - - 7 15 - 1 6 4 - - - 3 2 IV/VI

185 - - - - - - 2 - 1 - 1 - - - VI

190 - - - - - - 2 - - - - - - - VI/VII

195 - - - - - - 2 - - - - - - - VII

Value in first col umn re fers to the top of each ~5 cm spit. Gray shad ing in di cates the pres ence of one or more hearth fea tures.Layer des ig na tions are gen er al ized, since some are dis con tin u ous (e.g. Layer V) or are not nec es sar ily lie per fectly hor i zon talacross units. Be cause the se quence of ex ca va tion squares in the ta ble can not cor re spond to the orig i nal grid lay out, squares arelisted alphanumerically.

300 M. C. Stiner

Fig. 8. The dis tri bu tions of lime stone rocks (white) and hearth fea tures (black and dark gray) in 5 cm cuts at 145through 180 cm be low da tum. The in ferred shel ter fea ture is ap par ent in cuts 150–175, in the area where rocks arecom mon but hearths con sis tently ab sent. Light gray back ground rep re sents sed i men tary ma trix

about the con texts of dec o ra tive be hav ior, pat -terns of hu man se lec tiv ity and ge og ra phy of thesepref er ences, and how the na tive biota were wo ven into lo cal or gen er al ized ex pres sions of hu maniden tity (Stiner, 2003). Lo cal bi otic com mu ni tiesex erted pow er ful back ground ef fects on the com -po si tion of or na ments made from an i mal skel e talma te ri als. Hu mans fil tered through this nat u ralback ground with their own strong pref er ences forshell shape, color, size, and nat u ral rar ity. Thesepat terns of se lec tiv ity are sur pris ingly con sis tentthrough the Up per Paleolithic across Med i ter ra -nean and in land re gions of Eur asia, while spe ciesor types of raw ma te ri als used for bead-makingfrequently were substituted (White, 1989; Tabo-rin, 1993; Stiner, 2003).

The early Up per Paleolithic or na ment as sem -blages from Klissoura Cave 1 are most sim i lar tothose from Adri atic sites with re spect to fa voredmol lusk spe cies. The nar row size dis tri bu tion ofthe Klissoura 1 shells rep re sents an other point ofsim i lar ity to or na ment as sem blages from coastalIt aly but also to Levantine Tur key, and to carvednon-shell or na ments from the in te rior of Franceand Germany.

The shell or na ments from Klissoura 1 are dis -tin guished by their re fined con tents if com paredto as sem blages from coastal sites. Though neverfar from the sea, Klissoura 1 was al ways an in land site. Ma rine shells suit able for or na ment mak ingwere not readily at hand, and few if any of the or -na ments were man u fac tured on site. There is alsocon sid er able ev i dence of “high-grad ing” or se lec -

tive win now ing of the or na men tal shells fromKlissoura 1 for har mony in color, form and qual -ity. One finds few if any hints of child’s play from the raw ma te ri als used, or the mis cel la neous junkso typ i cal of coastal as sem blages. In stead, nicelyfin ished ob jects were the rule. The prev a lence ofcord-wear sug gests that many of the or na mentsar rived al ready at tached to or ganic ma te ri als orhu man bod ies. What break age oc curred to the or -na men tal shells re flects damage from long-termuse rather than errors in manufacture.

The or na ment as sem blage from the ear li estAurig na cian (layer IV) is the rich est and the mostin struc tive about site func tion of the pe riod. Thisas sem blage is quite large, and the di ver sity of itscon tents is ex cep tion ally high, even af ter cor rec -tions for sam ple size dif fer ences among lay ers.Most of the or na ments in layer IV oc cur withinwhat ap pears to have been a man-made shel ter.This hearth-free fea ture is de fined by a jum bledring of large stones around a thin or ganic stain,and is sur rounded by many hearths. The or na -ments from in side the shel ter fea ture may havebeen at tached to one or more leather or tex tile ob -jects that once lined the floor of the structure.

As is gen er ally true of Up per Paleolithic or -na ments in Eu rope (Koz³owski and Otte, 2000),those from Klissoura Cave 1 are well de vel opedin char ac ter and ap pear sud denly in the strati -graphic se ries. Dis tinct, well-strat i fied Mous -terian in dus tries oc cur in all lay ers be low VI.There are no or na ments in the Mid dle Paleolithiclay ers ex cept in VI an VII im me di ately be low the

Shell or na ments from the Upper Paleolithic 301

Ta ble 8Den sity of or na ments in re la tion to all chipped stone ar ti facts and to chipped stone tools by layer or

layer group

Layer(s)N chipped stone

ar ti factsN chipped stone

tools

Per centgroundstoneof all lithics

MNI or na mentsOr na ments/all lithics

Or na ments/tools

5a 3955 134 0.1 5 0.0014 0.037

IIa-d 6281 251 3 9 0.0014 0.036

III-III' 5096 158 3 38 0.0075 0.24

III" 2935 97 4 23 0.0078 0.24

IIIa-g 31631 822 0.01 161 0.0051 0.20

IV 63922 2237 0.06 1218 0.0190 0.54

V 4153 137 0.07 32 0.0077 0.23

Per cent groundstone ar ti facts is cal cu lated based on all lithic ar ti facts in layer. Lithic data pro vided by Kaczanowska et al. (this is -sue).

shel ter fea ture of layer IV. Layer V has an in ter -mit tent dis tri bu tion and does not ex tend un der thearea in layer IV that con tains the shel ter fea ture.In stead, layer IV meets VI in this area of the ex ca -va tion. The spa tial dis tri bu tion of the shell or na -ments in lay ers VI–VII and young dates on someof the shell spec i mens are al most certainly theresult of localized intrusions and mix ing fromabove.

The ear li est Up per Paleolithic in dus try in theKlissoura 1 strati graphic se ries co mes from layerV and as so ci ates with an un am big u ous shell or na -ment as sem blage. Dated to greater than ca. 39,000 years BP, the lithic in dus try is dis tin guished by ahigh in ci dence of lunates, re sem bling Uluzzianindustries in It aly, and it is clearly an Up per Pale-olithic tech nol ogy. The or na ment as sem blagesfrom all of the early Up per Paleolithic lay ers ofKlissoura 1 show gen eral tax o nomic af fin i tieswith those from some older and co eval Auri-gnacian sites in It aly. The small or na ment as sem -blage from layer V re sem bles that from Grotta delCa val lo in It aly, due sim ply to the high in ci denceof Dentalium shells.

Vari a tion in the tax o nomic con tent of theshell or na ment as sem blages from Klissoura Cave1 also speaks to cli mate-driven changes in en vi -ron ment het er o ge ne ity in south ern Greece dur ingthe Late Pleis to cene. The great num ber of spe ciesrep re sented in the early Up per Paleolithic as sem -

blages re flects a mo saic of hab i tats that was morecom plex than ex ists in the Peloponnese to day.Lon ger oc cu pa tions, as is sug gested for the lowerAurig na cian oc cu pa tion, might tend to am plifytax o nomic di ver sity quasi-in de pend ently of en vi -ron men tal change, but this is not suf fi cient to ac -count for the dif fer ences in spe cies rich ness. Theearly Up per Paleolithic or na ments from Klisso-ura 1 greatly ex ceed the tax o nomic di ver sity ofor na men tal shells both from the youn ger lay ers ofthis site and those from all of the late Paleolithicand Mesolithic lay ers in Franchthi Cave (Shackle -ton, 1988). The re duc tion in tax o nomic di ver sityaf ter the LGM was al most cer tainly linked to theglobal rise in sea level, which drowned the inlandlakes, raised water tables and swamped manyshorelines.

Ac knowl edg ments

I am very grate ful to André Carlo Colonese (Uni-versit´ degli Studi di Firenze) and Giuseppe Manga-nelli (Università di Si enna) for their as sis tance in clar i -fy ing the paleontological con text of the Plio cene fos silshells from Klissoura Cave 1. I also thank the mem bersof the Klissoura 1 ex ca va tion team for ac cess of geo -log i cal data, site maps, and artifactual data, and the vol -un teers from the com mu nity of Berbati for their gen er -os ity and lo gis ti cal as sis tance dur ing the study. Gid eonHartman (Har vard Uni ver sity) generously made manyof the pho to graphs used in this manu script.

302 M. C. Stiner

Ap pen dix 1: Spe cies abun dance (NISP and MNI) in the in tact Up per Paleolithic through Mesolithiclay ers of Klissoura Cave 1, and for the in ter face be tween MP and UP de pos its (VII-VI)

a. Lay ers 3-5 to IIIc 3-5a 3, 5a IIa-d IIa-d III-III' III-III' III'' III'' IIIc IIIc

Taxon NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI

Dentalium, ridged types 1 1 4 4 0 0 0 0 0 0

Dentalium, smooth types 0 0 0 0 1 1 0 0 0 0

Columbella rustica 0 0 0 0 5 5 7 7 5 5

Cylope neritea 1 1 4 4 25 25 14 14 12 12

Cylope pelucida 0 0 0 0 1 1 0 0 2 2

Cylope sp. 0 0 0 0 5 5 2 2 0 0

Theodoxus spp. 0 0 0 0 0 0 0 0 4 4

Glycymeris spp. 1 1 0 0 0 0 0 0 0 0

Pecten maximus 2 1 1 1 1 1 0 0 0 0

Acanthocardia tuberc. 1 1 0 0 1 1 0 0 0 0

TO TAL 6 5 9 9 38 38 23 23 23 23

Shell or na ments from the Upper Paleolithic 303

b. Lay ers IIIe-g to VII IIIe-g IIIe-g IV IV V V VI-VII VI-VII

Taxon NISP MNI NISP MNI NISP MNI NISP MNI

Scaphopoda

Dentalium, indet. Type 0 0 6 3 0 0 0 0

Dentalium, smooth types 1 1 9 7 9 7 2 2

Dentalium, ridged types 0 0 4 3 2 2 0 0

Gastropoda

Haliotis lamellose 0 0 5 2 1 1 0 0

Calliostoma sp. 0 0 2 2 0 0 0 0

Gibbula adansoni 8 8 123 118 0 0 8 7

Gibbula albida 0 0 61 53 0 0 0 0

Gibbula richardi 3 3 19 19 1 1 0 0

Gibbula cf. um bi li cus 0 0 3 3 0 0 0 0

Gibbula spp. (other) 0 0 16 16 0 0 2 1

Clanculus corallinus 0 0 73 70 0 0 6 6

Clanculus cruciatus (?) 0 0 2 2 0 0 0 0

Monodonta ar tic u late 1 1 6 6 0 0 1 1

Monodonta mutabilis 1 1 10 10 0 0 0 0

Monodonta turbinate 0 0 6 6 0 0 0 0

Homalopoma sanguineum 4 4 111 110 1 1 3 3

Littorina neritoides 0 0 2 2 0 0 0 0

Turritella communis 0 0 2 2 0 0 0 0

Vermetus sp. 0 0 2 2 0 0 0 0

Cerithium vulgaris 0 0 1 1 0 0 0 0

Cerithium sp. 1 1 2 1 0 0 0 0

Naticarius sp. 0 0 4 4 0 0 0 0

Neverita/Naticarius sp. 1 1 1 1 0 0 0 0

Phalium sp. 0 0 3 3 0 0 0 0

Columbella rustica 42 34 88 85 3 3 3 3

Mitrella/Pyrene scripta 0 0 1 1 0 0 0 0

Pisania maculosa 1 1 6 6 0 0 0 0

Cancellaria cancellata 0 0 3 3 0 0 0 0

Cyclope neritea 56 54 410 395 7 7 16 15

Cyclope pelucida 7 7 34 34 3 3 1 1

Cyclope spp. 6 6 0 0 1 1 0 0

Hexaplex trunculus 0 0 1 1 0 0 0 0

Thais haemastoma 0 0 1 1 0 0 0 0

Hinia re tic u late 3 3 94 90 1 1 6 6

Sphaeronassa mutabilis 0 0 2 2 0 0 0 0

Colus jeffreysianus 0 0 1 1 0 0 0 0

Co nus mediterraneus 1 1 4 4 0 0 0 0

Indet. ma rine gas tro pod 0 0 3 3 0 0 0 0

Theodoxus spp. 8 8 97 95 1 1 5 5

"A"* 1 1 8 8 0 0 0 0

"B1" Corymbina rhodiensis* 0 0 15 15 1 1 1 1

"B2"* 0 0 8 8 0 0 0 0

"D,G" Melanopsis gorceixi* 0 0 12 12 0 0 0 0

304 M. C. Stiner

b. Lay ers IIIe-g to VII IIIe-g IIIe-g IV IV V V VI-VII VI-VII

Taxon NISP MNI NISP MNI NISP MNI NISP MNI

Bivalvia

Glycymeris sp. 0 0 5 3 1 1 0 0

Pecten maximus 1 1 17 1 0 0 1 1

Chlamys varia 0 0 3 1 0 0 0 0

Acanthocardia tuberculatum 2 2 14 3 1 1 1 1

Cerastoderma edule 0 0 0 0 1 1 0 0

TO TAL 148 138 1300 1218 34 32 56 53

(*) fos sil spe cies of Plio cene age, orig i nat ing from lake bed or lakeshore de pos its, source lo cal ity un de ter mined

Ap pen dix 2: Eco log i cal sum mary of Late Pleis to cene molluskan taxa used as or na mentsat Klissoura Cave 1, Greece

Fam ily Ge nus spe cies Name source Com mon name Diet Substratesize (mm)

Adult

SCAPHOPODA (Class)

Dentaliidae Dentalium dentale L. tusk C s 35-50

Dentalium vulgare DaCosta tusk C m,s 35-50

GASTROPODA (Class)

ARCHAEOGASTROPODA (Or der)

Haliotidae Haliotis lamellosa Lamarck ab a lone, ormer HA r 60-75

Trochidae

Gibbula adansoni Payr. top HD r,s,w 8-15

Gibbula albida Gmelin top HD – 10-24

Gibbula richardi Payr. top HD r,s,w 8-15

Clanculus cruciatus L. top shell HD r,g 10-13

Clanculus corallinus Gmelin top shell HD r,g 9-15

Monodonta =Gibbula articulata Lamarck top shell HD r 10-25

Monodonta =Gibbula mutabilis Philippe top shell HD r 10-15

Monodonta turbinata Born check ered top HD r 20-35

Turbinidae Homalopoma sanguineum L. red tur ban HA r,w 3-7

MESOGASTROPODA (Or der)

Littorinidae Littorina neritoides L. per i win kle H r 5-7

Turritellidae Turritella communis Lamarck turrit shell HD g,m,s 20-45

CerithiidaeCerithium rupestre Risso horn shell HD m,w,s 20-35

Cerithium vulgatum Brug. horn shell HD m,w,s 20-65

NaticidaeNaticarius= Neverita josephina Risso moon snail C s,g,m 25-40

Naticarius millepunctata + oth ers Lamarck moon snail C s 30-45

Cassididae Phalium = Cassis undulatum Born hel met shell Curch s 60-70

NEOGASTROPODA (Or der)

Pyrenidae =Columbellidae

Columbella rustica L. dove shell O s,r,w 15-20

Pyrene = Mitrella scripta L. dove shell O r,s,c 15-18

Buccinidae Pisania maculosa = striata Lamarck spot ted pisania C,SC r 15-32

Muricidae Hexaplex = Murex trunculus – – C – –

Nassariidae

Cyclope neritea1 L. mud snail C-SC s,m 8-17

Nassarius = Hinia costulata Renieri mud snail C-SC r,s 8-15

Sphaeronassa mutabilis L. bas ket whelk O s,m 18-28

Conidae Co nus mediterraneus Brug. cone shell C r,w 60-65

Shell or na ments from the Upper Paleolithic 305

Ap pen dix 2 continued

Fam ily Ge nus spe cies Name source Com mon name Diet Substratesize (mm)

Adult

MESOGASTROPODA (Or der)

Littorinidae Littorina neritoides L. per i win kle H r 5-7

Turritellidae Turritella communis Lamarck turrit shell HD g,m,s 20-45

CerithiidaeCerithium rupestre Risso horn shell HD m,w,s 20-35

Cerithium vulgatum Brug. horn shell HD m,w,s 20-65

NaticidaeNaticarius= Neverita josephina Risso moon snail C s,g,m 25-40

Naticarius millepunctata + oth ers Lamarck moon snail C s 30-45

Cassididae Phalium = Cassis undulatum Born hel met shell Curch s 60-70

NEOGASTROPODA (Or der)

Pyrenidae =Columbellidae

Columbella rustica L. dove shell O s,r,w 15-20

Pyrene = Mitrella scripta L. dove shell O r,s,c 15-18

Buccinidae Pisania maculosa = striata Lamarck spot ted pisania C,SC r 15-32

Muricidae Hexaplex = Murex trunculus – – C – –

Nassariidae

Cyclope neritea1 L. mud snail C-SC s,m 8-17

Nassarius = Hinia costulata Renieri mud snail C-SC r,s 8-15

Sphaeronassa mutabilis L. bas ket whelk O s,m 18-28

Conidae Co nus mediterraneus Brug. cone shell C r,w 60-65

FRESH- AND BRACK ISH WA TER MOL LUSKS (lakes, ponds, slow mov ing rivers)

GASTROPODA (Class)

NeritidaeTheodoxus cf. jordani Sowerby river nerite C s,m 7-9

Theodoxus cf. fluviatalis – river nerite C s,m 7-10

BIVALVIA or PELECYPODA (Class)

FILIBRANCHIA (Or der)

Glycymeridae Glycymeris = Pectunculus sp. – bit ter sweet F s,m,g 35-65

Pectinidae Pecten maximus L. gi ant scal lop F r,s 100-150

EULAMELIBRANCHIA (Or der)

CardiidaeAcanthocardia tuberculatum L. cockle shell F s,m,g 25-90

Cerastoderma edule 1 L. ed ible cockle F s,m,g 30-50

Notes: Plio cene fos sil taxa are not in cluded in this ta ble. (¹) tol er ates or pre fers brack ish wa ter. Sub strate codes: (r) rock and otherfirm sur faces, (m) mud, (f) float ing mat ter and bub bles, (s) sand, (v) var ied, (w) weeds, (c) cor als, (g) gravel or coarse sand, (sp)sponges, (r/s) ad heres to hard sur faces as ju ve nile but free swim ming as adult. No men cla ture: (L.) Linnaeus; (Blainv.) Blainville;(Monter.) Monterosato; (Payr.) Payraudeau; (Brug.) BruguiÀre. Molluskan diet codes: (H) her bi vore, (O) om ni vore, (D) detriti-vore, (SC) scav en ger, (F) fil ter or sus pen sion feeder, (C) car ni vore.

Ap pen dix 3: Rel a tive fre quen cies of nat u ral and hu man-caused dam age to a sam ple of mol lusk shellsfound within well-de fined hearths in Layer IV

Ge nus(if > 5 MNI)

NISPex am ined

MNIex am ined

Shell com -pleteness

wave-worn%

pre-dated%

burned%

per fo rated%

holebroken%

with cordwear

%

withochre

%

Homalopoma 15 14 0.93 93 20 40 100 0 0 0

Cyclope 40 37 0.92 27 2 32 100 27 13 0

Theodoxus 10 10 1 0 0 20 100 30 20 10

Other gen era 59 30 0.93 37 0 25 100 34 0 4

All taxa 124 116 0.93 38 3 29 100 28 6 3

List ings are by gen era for com mon types, ex cept G. albida, which is sig nif i cantly larger than all other spe cies of Gibbula in the or -na ment as sem blages

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