Eurasian Prehistory, 7 (2): 287–308.
SHELL ORNAMENTS FROM THE UPPER PALEOLITHIC
THROUGH MESOLITHIC LAYERS OF KLISSOURA CAVE 1
BY PROSYMNA (PELOPONESE, GREECE)
Mary C. Stiner
School of Anthropology, P.O. Box 210030, University of Arizona, Tucson, Arizona 85721-0030, USA;
[email protected]
Abstract
More than 1500 shell ornaments were recovered during the excavations of the early Upper Paleolithic through Mesolithic layers of Klissoura Cave 1. The ornament assemblages from the middle and lower Aurignacian and the earliest Upper Paleolithic (Uluzzian) layers associate with well preserved hearths and other intact cultural features. The older ornament assemblages are exceptionally rich in mollusk species, whose shells humans collected from marine shorelines, freshwater habitats and Pliocene fossil sources. A particularly dense concentration of ornamental shells occurs within the area
of a small structure in the lower Aurignacian (layer IV). Taxonomic diversity in the ornament assemblages declines precipitously after the formation of layer IIIe–g (upper Aurignacian), and this condition of low diversity persists through the
end of the cultural sequence. The changes in ornament diversity seem to reflect natural changes in coastline habitat structures of the region. All of the Upper Paleolithic ornament assemblages are “high-graded” or selectively winnowed for harmony in color, form and quality. There are few if any hints of manufacturing errors and debris typical of shell ornament assemblages in coastal sites. Rather, the ornaments display high frequencies of use wear (polish), usually in a preferred orientation, indicating that most of them arrived on site while affixed to human bodies or organic artifacts. There are no remains of edible marine mollusks in Klissoura Cave 1, consistent with its inland location. The taxonomic composition of
the early Upper Paleolithic shell assemblages is similar to those documented in Italy, whereas the very limited taxonomic
composition of the later ornament assemblages is most consistent with those found at Franchthi Cave on the southern
Argolid.
Key words: Paleolithic shell ornaments, Aurignacian, Uluzzian, taphonomy, site formation processes, site structure.
INTRODUCTION
Comparatively little is known about Paleolithic ornaments from Greece. The number of excavated Paleolithic sites has climbed slowly over
the last five decades, and only a few of these sites
– Franchthi Cave and now Klissoura Cave 1 –
provide much information about early decorative
traditions and the contexts of their use. Paleolithic
art, including beads, is considered an important
criterion for the emergence of modern human behavior during the Late Pleistocene (e.g., Klein,
1989; Mellars, 1989; White, 2003).
Decorative traditions exist in virtually every
recent human culture, so much so that early Paleolithic beads and other ornaments might seem or-
dinary and without interest. The behavioral importance of the emergence of bead-making traditions in the early Upper Paleolithic is underscored, however, by the dearth of them in archaeological sites prior to this period in much of
Eurasia (but see d’Errico et al., 1998; Bar-Yosef
Mayer et al., 2009) in contrast to some African
Middle Stone Age sites (Bouzouggar et al., 2007;
Henshilwood et al., 2004).
Ornaments made from mollusk shell, ostrich
eggshell, mammal teeth and ivory, and soft stone
are considered to be a unique evolutionary development on account of their visually striking qualities, transferability among persons (sensu durable
items of trade, gifts or burial paraphernalia) (Kuhn
and Stiner, 2007), and the stylistic formalizations
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M. C. Stiner
apparent from consistencies in size and appearance (Hahn, 1972; Bar-Yosef, 1989; White, 1989,
1993; Taborin, 1993; Stiner, 1999, 2003; d’Errico
et al., 2001; Kuhn et al., 2001; Bouzouggar et al.,
2007). Mineral oxides, on the other hand, were
widely used as early as the Middle Paleolithic period in Eurasia, perhaps to color hides, human
skin, or tools. While decoration is only one of
several possible explanations for the presence of
pigments in Middle Paleolithic toolkits, the possibility of such practices in the Middle Paleolithic
cannot be refuted with current evidence. Even if
ochre and other minerals were used as pigments in
the earlier periods, archaeologists’ efforts to link
their occurrence in sites to prehistoric decorative
traditions remains controversial (but see Henshilwood et al., 2009). The information that painted
body designs may carry is transient and nontransferrable (Kuhn and Stiner, 2007), in contrast
to the durable properties of some ornaments. Evidence for design repetition is necessary to any argument for shared symbolic significance in prehistoric decorative objects (Kuhn and Stiner,
2007). It is likely that ornament production and
use was greatest where interaction among social
groups was high, possibly in connection with regionally high human population densities (Kuhn
et al., 2001). The sudden appearance of durable
ornamental objects in human cultures of the Late
Plei- stocene may even suggest the emergence of
a basic visual grammar in social interactions (e.g.,
Gamble, 1986; d’Errico et al., 2003; Kuhn and
Stiner, 2007).
The timing of the first appearance and subsequent proliferation of ornament traditions varies
across world regions. An important question for
this study is the chronology and evolution of ornament traditions in southern Greece. Few paleoanthropologists would be surprised to learn that
ornaments appear suddenly in the archaeological
record of Greece with beginning of the Upper
Paleolithic culture period. Most early Upper
Paleolithic industries in Greece are attributed to
the Aurignacian (Runnels, 1996) and have been
identified 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 recently at Lakonis
(Panagopoulou et al., 2002–2004). Early Upper
Paleolithic industries that resemble the Uluzzian
of Italy (Palma di Cesnola, 1966, 1993; Benini et
al., 1997; Gambassini, 1997) have been found in
Klissoura Cave 1 (Koumouzelis et al., 2001;
Kaczanowska et al., this issue). Several Upper
Paleolithic sites in Greece contain ornaments
made from shells (e.g., Shackleton, 1988; Koumouzelis et al., 2001). Recent excavations at
Klissoura Cave 1 have yielded exceptionally rich
ornament traditions in the early Upper Paleolithic
of Klissoura Cave 1, including the Uluzzian layer.
BACKGROUND TO KLISSOURA
CAVE 1
More than 1500 shell ornaments were recovered during the excavations of the early Upper
Paleolithic through Mesolithic layers of Klissoura
Cave 1. The ornaments from the early Upper
Paleolithic layers (V–IIIc) occur within and
among a dense array of well preserved sedimentary features, mainly hearths. Shell ornaments are
most abundant in the middle to lower Aurignacian
layers, particularly layer IV.
Several general observations about Klissoura
Cave 1 are important to this presentation of the
shell ornaments. This shallow cave is situated on
the Berbatias River and commands a wide view of
the Argos plain, roughly 12 km inland of the
Argolikos Gulf (Fig. 1). Most of the ornamental
shells nonetheless were collected from marine
shores. The great variety of features, artifacts and
vertebrate faunal remains in the Upper Paleolithic
layers indicates that the site served mainly at a
residential base, particularly in the early phases of
occupation. An area of 13–15 square meters was
excavated (Karkanas et al., 2004; Koumouzelis et
al., 2001), and the units cut through the heart of
several occupational layers in this small cave.
The cultural stratigraphy in Klissoura Cave 1
is more than 5 m deep and includes a long Middle
Paleolithic sequence (Sitlivy et al., 2007) capped
by Upper Paleolithic and Mesolithic horizons
(Karkanas, this issue). Apart from locally disturbed zones at the interface of the Middle and
Upper Paleolithic (VII/VI), ornaments are confined to the Upper Paleolithic (V, IV, III–III” &
6a), Epipaleolithic (II) and Mesolithic layers
(5–3a) (Table 1). In the early Upper Paleolithic
layers, hearth features occur in more than onethird of all excavation units. The Aurignacian lay-
Shell ornaments from the Upper Paleolithic
289
Fig. 1. (Left) Location of Klissoura Cave 1 by Prosymna on the Argolid of Peloponnese, Greece; other sites
shown are Franchthi Cave (2) and Kephalari Cave (3). (Right) Comparison of modern shorelines to the distribution
of paleolakes (black fill) and shorelines at the Last Glacial Maximum in southern Greece (adapted from Petit-Maire
et al., 2005). White fill represents exposed land surfaces at the time of the LGM, which are now inundated
ers are unique not only for their high concentrations of wood ash and intact hearths, but the fact
that many were lined with clay (Karkanas et al.,
2004; Pawlikowski et al., 2000). Burning damage
is common on the bones, ornaments, and lithics
from all of the layers (Koumouzelis et al., 2001;
Tomek and Bocheñski, 2002; Karkanas et al.,
2004; Starkovich and Stiner, this issue).
Variation in ornament abundance in the Upper Paleolithic through Mesolithic layers is not
explained by differences in the thickness of the
excavated deposits. Following Karkanas (this issue), the Mesolithic and probable Epigravettian
(IIa–d) horizons are 10–20 cm in thickness, except where large pits invade the older layers. The
Upper Paleolithic sequence (III–V) is much
Table 1
Layer terms, cultural attributions, and geological sequences for the ornament samples from Klissoura
Cave 1
Culture attribution
Layer terms
Geological sequence
Ornament NISP
3, 5a
A
5
IIa, IIb, IId
B
9
6, 6a, 6/7
B
8
III-III'
D
38
Upper Paleolithic (non-Aurig.)
III"
D
28
Upper Aurignacian
IIIc
D
23
IIIe-g
D
138
IV
E
1218
Mesolithic
Epigravettian
Disturbed zone
Medit. Backed-bladelet industry
Mid-Aurignacian
Lower Aurignacian
Earliest UP (Uluzzian)
V
F
32
Disturbed, mostly MP
VI
(F/)G
28
VII
G
52
Middle Paleolithic
The geological sequences are layer groups or facies with similar sedimentological characteristics that form coherent layer sets,
separated by discrete contacts that indicate minor or major depositional hiatuses (following Karkanas, this issue). The Mediterranean backed-bladelet industries of layers III-III' and III'' differ from one another in some aspects but have in common the high frequency of the implements.
290
M. C. Stiner
thicker – about 190–210 cm overall. A disturbed,
mixed deposit was encountered in layers 6–7 (see
Table 1) and will not be discussed further.
In the layer III series, clay-lined hearths are
common only in layer III’. A large space devoid
of hearths and large rocks was found in squares
AA-CC1 at the depths of 75–85 cm below datum,
and a short arc of clustered rocks was encountered
between 85–105 cm. Sediments containing a nonAurignacian backed-bladelet industry of typical
Mediterranean type (layers III–III’) and another
layer (III”) containing a non-Aurignacian industry occur above the Aurignacian layers (IIIb–g,
IV) (Kaczanowska et al., this issue).
The earliest Aurignacian layer IV varies between 25–50 cm in thickness and is exceptionally
rich in bones and artifacts. In addition to many
clay-lined and unlined hearths, this layer preserves a wide assortment of lenses, pits, and the
outline of what was probably a small man-made
shelter. Rare antler points and significant amounts
of what appears to be osseous manufacturing debris also occur in layer IV (Christidou, personal
communication, 2010; Starkovich and Stiner, this
issue).
Layer V is a thin, undulating deposit that occurs only in the western part of the excavation and
is characterized by many concave hearth features.
A small ornament assemblage accompanies the
distinctive lithic industry (Table 1), whose arched
backed bladelets and comparatively high proportion of microblades resemble Uluzzian industries
in Italy (cf. Palma di Cesnola, 1993; Benini et al.,
1997; Gambassini, 1997; Kuhn and Stiner, 1998),
warranting a comparison of the Greek and Italian
ornament assemblages.
The first radiocarbon determinations attempted for the Upper Paleolithic layers suggested
comparatively young ages for the Aurignacian at
Klissoura 1 (Koumouzelis et al., 2001; compare
Koz³owski, 1982, 1992, 1999). Preliminary
thermo-luminescence results on burnt flint artifacts suggested that layer V is significantly older.
New AMS radiocarbon results using the ABOX
pretreatment method (Bird et al., 1999) on wood
charcoal samples from the late Middle Paleolithic
through Upper Paleolithic layers permit some revisions to the Upper Paleolithic chronology
(Pigati et al., 2007; Kuhn et al., this issue). Still,
the dates for the earliest Aurignacian layer (IV) –
32,690±110 and 33,150±120 uncali- brated radiocarbon years before present (BP) – are not nearly
as old as those for many other Aurignacian cases
in Europe (e.g., Conard et al., 2003), but they are
somewhat older than Aurignacian sites in the
Levant. The Uluzzian-like industry in layer V is
sealed from the layer above by a fine tephra and
may be older than 39,000 calibrated BP.
GOALS AND METHODS
This study addresses several issues surrounding ornament use and discard at Klissoura Cave 1:
(a) Species composition as it relates to marine and
terrestrial environmental conditions, diversity in
raw material sources, and evidence for selectivity
by humans; (b) damage patterns on the shells that
may reflect raw material sources, ornament manufacture, use and discard practices; (c) the spatial
distribution of ornaments and damage phenomena
in the deposits, including associations with hearths
and other features; (d) trends in ornament assemblage composition from the early Upper Paleolithic through Mesolithic; and, (e) comparisons of
the Klissoura 1 ornament series to those from
early Upper Paleolithic Italian sites and to the
later ornament assemblages from Franchthi Cave
on the southern Argolid (see Fig. 1). The quantitative units used in the analyses are the number of
identified specimens (NISP) and the minimum
number of individual animals (MNI) represented
by whole and fragmentary remains. NISP is important for many of the taphonomic analyses,
whereas MNI better represents ornament quantities from a functional point of view.
To address questions about the contexts of ornament use and their possible significance in
daily life, this study considers both the general
processes of assemblage formation and how the
ornaments were made, used and exhausted as
items of technology. Species-specific ecologies of
the mollusks are used to infer the range of habitat
sources, and observations about shell condition
and spatial distributions help to determine the
contexts of ornament use on site. Reconstructions
of Late Pleistocene shorelines and other water
bodies at the Last Glacial Maximum (LGM, ca.
20,000 years BP; Lambeck, 1996; Petit-Maire et
al., 2005) and recent Holocene periods define two
extremes in the Pleistocene shoreline and habitat
Shell ornaments from the Upper Paleolithic
291
Table 2
Indexed comparison of species richness for early Upper Paleolithic through Mesolithic shell ornament
assemblages by intact layer from Klissoura Cave 1
Layer group
Number of species
Ornament MNI
Species richness
Mesolithic (3, 5a)
5
5
7.15
Epipaleolithic (IIa-d)
Med. Backed-bladelet Industry
(III-III')
Upper Paleolithic (non-Aurig.) (III")
3
9
3.14
6
38
3.80
2
23
1.47
Upper Aurignacian (IIIc)
4
23
2.94
Mid-Aurignacian (IIIe-g)
18
138
8.41
Lower Aurignacian (IV)
45
1218
14.58
Early UP/Uluzzian (V)
14
32
9.30
14
Undetermined (VI-VII)
Species richness is calculated as N-species/logMNI.
53
8.12
configurations of southern Greece (Fig. 1). Environmental conditions on the Argolid during the
early Upper Paleolithic would have fallen somewhere within these extremes, though closer to
LGM conditions with respect to land surface exposure, peninsula-island configurations, and the
diversity of aquatic habitats within a 50 km radius
of the site. The later occupations, particularly during the Mesolithic period, would have experienced conditions more like those of the recent Holocene.
RESULTS ON THE PALEOLITHIC
ORNAMENT ASSEMBLAGES
All of the ornaments from Klissoura Cave 1
were made from aquatic mollusk shells, mainly
small marine gastropod species along with some
fresh or brackish water and fossil types. A few red
deer (Cervus elaphus) canines occur among the
faunal remains (Koumouzelis et al., 2001; Starkovich and Stiner, this issue) but none of these
was altered for suspension. A few perforated teeth
were noted in the early years of the excavation
project but these could not be located or confirmed
Mollusk species abundance and diversity
The diversity of mollusk species varies greatly
among the Upper Paleolithic and Mesolithic layers
in Klissoura Cave 1 (Table 2 and Appendix 1), but
species distributions do not vary significantly
among features or excavation units within each
layer. Species richness, here calculated as N-species/log MNI to correct for sample size effects, is
greatest in layers IIIe–g and VI, especially in IV
where a minimum of 44 taxa were identified. The
ornament assemblages from the later UP and Epipaleolithic layers are much poorer in species,
even after correcting for the smaller sample sizes,
followed by a mild increase in diversity in the
Mesolithic. Taxonomic diversity is uniformly low
in the large Epipaleolithic and Mesolithic ornament assemblages from Franchthi Cave on the
southeastern margin of the Argolikos Gulf (compare Shackleton, 1988; PerlÀs and Vanhaeren,
2010).
Steep rocky or heterogeneous Mediterranean
coasts with high nutrient turnover tend to support
many mollusk species. Nearby sites containing
Upper Paleolithic shell ornament assemblages
also tend to be species-rich (e.g., Riparo Fumane
and Riparo Mochi in northern Italy; Leonardi,
1935; Bartolomei et al., 1994; Fiocchi, 1996-97;
Stiner, 1999, 2003). Today Klissoura Cave 1 lies
as close as it ever has to the Argolikos Gulf. The
site would have been a few more kilometers distant from shore at the time of the Aurignacian occupations, but without radical alterations in shoreline shape or topography (Fig. 1).
The central geographic position of the Klissoura Gorge to brackish lagoons, rivers, lakes
and marine shorelines of the Peloponnese and
mainland Greece during glacial periods contributed to the taxonomic variety in the ornament assemblages in layers IIIe–g through V. Shells were
M. C. Stiner
292
Table 3
Representation of habitat sources and trophic categories in shell ornaments from layer IV,
the most diverse assemblage
By habitat type
%
By trophic group
%
Fossil source
4
Carnivore
10
Marine mixed shore
35
Carnivore-scavenger
42
Estuarine-lagunal
52
Omnivore
6
Fresh-brackish
8
Herbivore-detritivore
40
Fig. 2. Fossil taxa from Pliocene lake deposits of the genera Corymbina (probably C. rhodiensis and C. aegae) and
Melanopsis (e.g. M. gorceixi), following Magrograssi (1928). Image compiled from photographs by G. Hartrman
Table 4
Summary of relative abundances of common genera (% of total MNI) in the ornament assemblages
from Klissoura Cave 1
Layer(s):
3, 5a
IIa-d
6-6/7
III-III'
III"
IIIc
IIIe-g
IV
V
VI-VII
4
% of total MNI
Taxon
Dentalium spp.
20
44
0
2
0
0
1
1
29
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
Fossil lake spp.
0
0
0
0
0
0
1
3
3
2
Bivalve spp.
60
11
0
4
0
0
2
1
9
4
All other taxa
0
0
0
0
0
0
4
3
3
0
5**
9**
8**
38
23
Sample size (MNI)
(**) Sample size is very small, making percentage calculations suspect.
23
138
1218
32
53
collected from the wide range of marine and inland habitats during these occupation phases (Table 3 and Appendix 2). At least 8% of the ornamental shells from layer IV are freshwater species
(Theodoxus), and another 4% are fossils from
Pliocene lake bed deposits (Fig. 2; Magrograssi,
1928; G. Manganelli and A. C. Colonese, personal communication, 2007). More than half of
the shells originate from Late Pleistocene brackish lagoons or estuaries (Cyclope), and the remaining 35% are from the marine littoral. Other
factors that may have contributed to species richness in the older ornament assemblages were the
greater intensity or duration of occupation and the
greater diversity of activities on site (see below).
Shell ornaments from the Upper Paleolithic
Fig. 3.
293
Relative frequencies of the major ornament genera in the Upper Paleolithic layers of Klissoura Cave 1
The ornament assemblage from layer V is
also species-rich for its size, but the prominence
of Dentalium (tusk) shells sets it apart (Table 4).
The layer V assemblage is small, however, and it
should be kept in mind that one distinctive bead
strand would be enough to alter the character of
this small assemblage. Interestingly, Dentalium
shells are also common in the Uluzzian horizons
of Grotta del Cavallo in Italy (Palma di Cesnola,
1966).
Small quantities of ornaments also occur in
layers VI and VII of Klissoura 1. These must be
considered separately, since a mixture of Upper
Paleolithic and (mostly) Mousterian artifacts oc-
curs in these layers, along with few faunal remains (Starkovich and Stiner, this issue). The species content of the ornament assemblage from
VI–VII best resembles that of layer IV (Fig. 3),
which lies in direct contact with layer VI horizontal units in the area where most of the ornaments
were found (squares AA1-2 at 175–185 cm below
datum). This and other evidence (K. Douka, personal communication, February 2010) indicates
that the few ornaments that locally co-occur in
Middle Paleolithic artifacts in layers VI and VII
originated from layer IV.
The structure of the Peloponnese landscape
during the Late Pleistocene and early Holocene is
294
M. C. Stiner
important to understanding ornament raw material sources as well as the potential foraging opportunities for humans of the period. The Gulf of
Korinthos, Argolid Peninsula, and Cyclade Islands have been the focus of several shoreline reconstructions (van Andel and Sutton, 1987;
Shackleton, 1988; Lambeck, 1996; van Andel and
Tzedakis, 1996). These reconstructions are invaluable for estimating the distance between site
and coast at the time of occupation, the complexity of coastal and inland aquatic habitats, and the
heterogeneity of the environment in successively
larger catchments around the site. The most recent
reconstruction by Lambeck (1996) integrates
models of eustacy and glacio-hydro isostasy with
topographic data for tectonically stable areas to
estimate net changes in land exposure. Groundtruthing was accomplished in this and prior studies partly by reference to dated archaeological and
paleontological deposits. Layers III and below in
Klissoura Cave 1 would have formed during intermediate glacial conditions, and layer II and the
Mesolithic layers would have formed around the
time of the Pleistocene–Holocene temporal
boundary. The older ornament assemblages predate the Last Glacial Maximum by many thousands of years, but according to Lambeck (1996:
596), [during] “…much of the time between about
70,000 years b.p. and the Glacial Maximum, ice
volumes significantly exceeded those of today
such that the global sea-levels did not rise above
40–50 m below present level during this interval.”
Landscapes to the north and east of the Klissoura Gorge had greater land mass exposure during the early Upper Paleolithic, interspersed locally with marshes and estuaries and at least two
large freshwater lakes to the northeast. The landscape and ecology of this period therefore was
more complex than the modern one, and the terrestrial ecosystems would have been more productive. The central location of the gorge and its
small caves to diverse habitats was a major attraction to Pleistocene foragers in the eastern Peloponnese.
The rapid decline in species richness in the
shell ornaments after layer IIIe–g but within the
Aurignacian period may be explained by the
growth and eventual dominance of lagunal habitats on the western shores of the Argolid (see
Shackleton, 1988). While changes in culturally
bound aesthetics may also have contributed to the
narrowed range of mollusk species used for ornament-making in layers IIIc–d and after, changes
in natural biotic diversity were the primary constraint on human choices and form the baseline
against which variety must be interpreted.
Raw material sources and evidence for human
selectivity
The types of ornamental shells in Klissoura 1
are not the most common species on Mediterranean shores today or in the past. “Unnatural” biases in taxonomic composition include comparatively high frequencies of carnivorous species,
and artificially narrow shell color, shape and size
distributions. The early Upper Paleolithic occupants clearly preferred shells of the genera, Gibbula, Clanculus, Homalopoma, Columbella, Cyclope, Hinia, and Theodoxus (Table 4). The proportions of Columbella and Cyclope increase with
time and these two genera ultimately dominate the
younger Upper Paleolithic assemblages (Fig. 3).
Gibbula albida was important only during the formation of layer IV, and most Clanculus shells
also occur there. Dentalium shells are common in
layer V, as noted previously by Koumouzelis et
al. (2001), and they regain importance only during the Epipaleolithic and Mesolithic.
Carnivorous species (predators and scavengers) in the ornament assemblages are represented
well in excess of the expected encounter rate in
living mollusk communities. Global censuses of
mollusk communities (e.g., Sabelli, 1980) and
collecting experiments by the author on eastern
Mediterranean shores (Stiner, 1999) suggest average encounter rates for carnivores at roughly 15%
of all gastropod individuals in beach-cast material. In Aurignacian layer IV, specialized carnivores occur at a rate of 10% but together with carnivorous scavengers constitute 52% of shell MNI
(Table 4). The pattern of trophic representation in
the ornaments is the opposite of that for natural
mollusk communities and must reflect strong selectivity on the part of Paleolithic humans. Shell
collectors probably were responding to visually
attractive properties of the shells coupled with a
sense of their ecological rarity (Stiner, 2003).
Damage patterns on the shells indicate that
most or all of the specimens were collected as
empty shells from marine and estuary beaches,
Shell ornaments from the Upper Paleolithic
295
Fig. 4. Size distribution (cm) of measured ornaments, exemplified by the shells from layers IV and V of Klissoura Cave 1
Fig. 5. Typical shell forms and perforations made by humans.The holes were made by a simple punching technique, except for tusk shells (Dentalium) which were snapped to create tube beads. Shell taxa from left to right are
Homalopoma sanguineus, Nucella lapillus, Cyclope neritea, Dentalium sp. Theodoxus spp (2 specimens, one with
stripes), Monodonta sp., Columbella rustica, and Clanculus corallinus. Image compiled from photographs by G.
Hartman
river banks and mouths, and fossil beds. Wave-induced abrasion is widespread on the marine shells
and on some of the fossil types (Table 4b). Abrasion damage is least common on fresh and brackish water species and probably reflects the lower
energy levels of the aquatic habitats where these
shells were obtained. Abrasion damage and boreholes from molluskan predators occur on some of
the shells from Klissoura 1 (0–8% of shells), also
consistent with collection from wave-cast beach
sources. There are no large, edible marine mollusk remains in any of the layers of Klissoura
Cave 1.
Human selectivity is apparent from the narrow range of shell shapes and sizes in the Klissoura ornaments. This pattern holds true for all of
the shell assemblages. Small species were strongly
preferred (Fig. 4, Table 5b), with the mean length
for all measurable (whole) shells in the Upper
Paleolithic layers between 1.3–1.4 cm, and between 1.4–1.7 cm in the younger layers. The full
size range is 0.6 to 5.5 cm, but nearly all of the
shells are under 2 cm in length. The consistency in
ornament size within this range is explained by
the humans’ narrow preference for certain species. Round, oval or basket-like forms were the
norm in the Klissoura assemblages (Fig. 5), along
with Dentalium tubes. Similar mean sizes and
ranges are reported for other Upper Paleolithic
sites in Europe (e.g., White,1989; Stiner, 2003),
including ornaments from the Périgord region of
France that were laboriously carved from ivory
and stone (White, 1989: 382).
Red and pink shells were sought out preferentially in addition to those with bright opalescent
hues or contrasting stripes. Ochre stained shells
occur throughout the Upper Paleolithic layers
(Table 6a), but ochre application centered on two
genera: Clanculus, a group with naturally pink or
red shells and rough, knobby surfaces; and Theodoxus, a group with bright white or vividly striped
shells and smooth surfaces (Table 6). Ochre traces
M. C. Stiner
296
Table 5
Damage and shell size statistics for the larger assemblages of Upper Paleolithic ornaments by layer
from Klissoura Cave 1
a. burning damage, perforation rates, breaks through perforation point, cord wear and the incidence
of ochre traces
Layer group
Burned
%
Perforated
%
Hole broken
%
With cord-wear
%
With ochre
%
Non-Aurig. UP industries (III-III")
51
97
14
69
3
Upper Aurignacian (IIIc)
58
92
8
50
8
Mid-Aurignacian (IIIe-g)
55
92
26
38
3
Lower Aurignacian (VI)
18
90
21
18
6
Early UP/Uluzzian (V)
48
96
17
7
0
Undetermined (VI-VII)
4
99
32
47
4
b. completeness, size statistics, and non-cultural types of damage
Layer group
Complete-ness
With natural color
%
Mean length
(cm)
Sd
(cm)
Wave-worn Predated
%
%
Non-Aurig. UP Industries (III-III")
0.97
11
1.43
0.26
23
6
Upper Aurignacian (IIIc)
1
17
1.47
0.3
17
8
Mid-Aurignacian (IIIe-g)
0.91
8
1.45
0.6
52
2
Lower Aurignacian (VI)
0.94
10
1.28
0.44
35
4
Early UP/Uluzzian (V)
0.93
3
1.28
0.55
31
3
0.94
13
1.27
0.29
49
0
Undetermined (VI-VII)
Notes: Only the larger samples from the III series are below are considered. Perforation refers to holes made by humans; hole broken refers to specimens that were broken through the perforation point on the shell, either during manufacture or (more commonly) from long-term use; cord wear refers to polish on some or all edges of the perforation but may also include polish on the
outer surface near the perforation. Shell completeness is calculated as MNI/NISP. Length measurements are only for those shells
whose natural dimension could be determined. Predated shells have holes drilled in them by a mollusk predator. Natural color refers to the retention of natural pigment within the shell.
occur most often on Clanculus shells, at about 3
times the rate observed for Theodoxus. Ochre
traces were found on several other shell types but
Table 6
Distribution of ochre traces by mollusk genus
in all of the Upper Paleolithic layers combined
Genus
% with red ochre
Clanculus
29
Columbella
1
Cyclope
5
Gibbula
2
Homalopoma
4
Hinia
7
Theodoxus
11
All other genera
3
at much lower frequencies. Though a rough surface might generally increase the chances of
ochre traces being preserved, ochre is also frequently preserved on smooth shells in the Klissoura 1 assemblages, so the bias is not from differential preservation of applied pigment. Reddish
hues were valued, and people went to some effort
to make naturally reddish (but faded) shells
redder.
Contexts of ornament use and discard
Shell fragmentation is measured in this study
by an index of “completeness." The index is calculated by dividing the number of individuals by
the total number of identified fragments in each
taxonomic group and layer (MNI/NISP, Table
5b). Nearly all of the gastropod shells from Klissoura 1 are complete or nearly complete (89–
Shell ornaments from the Upper Paleolithic
100%, depending on the species), except for fractures at the perforation point. Dentalium shells
were usually sectioned to make tube beads. Rare
bivalves (Pecten and Chlamys) were nearly always broken, probably because they are large,
fragile and prone to trampling.
A significant number of the ornamental shells
are burned (Table 5a). Most or all of this damage
appears to have been accidental. There is much
evidence of hearth rebuilding, renewal, and superposition in the Upper Paleolithic layers (Koumouzelis et al., 2001; Karkanas et al., 2004),
causing older debris to be damaged by the heat
from superimposed hearths (see Stiner et al.,
1995). Burning damage is least common on the
ornament shells from layer IV, VI and VII (Table
5a). Most of the ornaments in layer IV were found
within the perimeter of the man-made structure
and immediately below it.
The great majority of the shells in each assemblage have holes in their flanges or whorls
(90–95%, Table 5a), and there is little if any waste
material on site. The holes in gastropod shells
usually were made by humans using a punching
technique that produced small round openings
with rough edges. Some of the marine shells have
holes caused by wave action or drilling by molluskan predators. Humans sometimes took advantage of these “natural” holes if located in the shell
flange or dorsally (opposite) to the flange. Waveworn and predator-drilled holes tend to be distributed more randomly on shell surfaces, and those
drilled by molluskan predators are almost perfectly round and beveled in contrast to man-made
holes (Figs 7 and 8). Holes caused by wave action
are as abraded and thinned along with the rest of
the shell, in contrast to localized polishes resulting from wear against organic fibers or hide.
Gibbula albida shells were treated differently
from other types of shells in that fewer than half
of the specimens were perforated. This is a comparatively large top shell with an elegant pyramidal form and pronounced concentric ridges. G.
albida shells are common only in layer IV (MNI =
71). They concentrated between the depths of
150–170 cm below datum, mainly in squares
AA1-AA2 (a few also occur in BB1-BB2) and
therefore coincide with the interior of the manmade shelter (see below).
The generally high perforation rate for the
297
Fig. 6. Example of cord wear in human-made hole
and outer surface of Theodoxus shell. Photo by G.
Hartman
Fig. 7. Comparison of holes made by humans (1),
and another by molluskan predators (2) of the
Naticidae or Muricidae families
shell ornaments from Klissoura 1 overall lies at
one extreme in the continuum of variation observed among Upper Paleolithic cases that contain ornamental shells. At coastal sites, such as
Riparo Mochi (Liguria, Italy; Stiner, 1999) and
ÜçaÈÏzlÏ I Cave (Hatay, Turkey; Stiner, 2003;
Kuhn et al., 2009), perforation rates are lower and
manufacturing errors more apparent in shell ornament assemblages, along with evidence for
stock-piling or raw material and accumulation of
rejected pieces.
Cord wear was identified on the shell specimens from Klissoura 1 with the aid of a hand lens,
and thus the percentages in Table 5a represent
minimum estimates of occurrence. Cord-wear inside the perforations tends to be asymmetrical,
suggesting that the shells were fastened or strung
in one position for long periods. Surface polish on
the outer surface near the perforations accompanies the evidence of cord-wear on some of the
shells (Figs 6, 7). The incidence of cord wear at
298
M. C. Stiner
Klissoura is high in comparison to the coastal site
of Riparo Mochi in Italy.
About one fifth of the specimens from Klissoura 1 Cave were broken through the perforation
point (Table 5a, hole broken 8–32%). The frequencies of breaks at the perforation is positively
correlated with the incidence of cord polish
(.05>>p>>.02), probably because both of these
phenomena relate to ornaments becoming worn
out from extended use. The prevalence of cord
wear, the high perforation rates, and the near absence of manufacturing waste argue against the ornaments having been manufactured on site. There
is, as well, an appreciable winnowing or “high
grading” effect apparent in the ornament assemblages. All told, the ornaments are generally complete, attractive specimens with a narrow size distribution and considerable evidence long-term use.
Spatial distributions of the ornaments
Ornaments are abundant in layer IV and especially within the domain of the hypothesized
man-made structure. This feature spans squares
AA1-2 of the excavation (Table 7), where simplified site plan drawings (Fig. 8) reveal the mutually exclusive distributions of large limestone
rocks and hearths at 150–175 cm below datum.
The structure is defined by the circular jumble of
large rocks, minimally 2 m in diameter, and a complete absence of hearths. Dozens of hearth features
encircle this area. The rocks may originally have
weighted the edges an organic covering, and rolled
inwardly upon removal or decomposition of the
cover. Dark stained sediments were observed at
155–165 cm below datum and may be the remnants of a floor covering. Given that the ornaments
cluster within the shelter area to a large degree,
and many display cord wear from prolonged use,
it is possible that they were once attached to an organic object such as a large hide or matt. The concentration of ornaments within the structure also
explains the lower incidence of burned specimens
in layer IV in comparison to the ornament from
the other layers (Table 5a). The only other shell
assemblage that displays a low burning rate is
from layers VII–VI, immediately below the shelter feature, and where layer V does not extend
horizontally. Small numbers of ornaments extend
downward into layers VII–VI at one edge of the
shelter feature area of layer IV (Table 7).
Layer V is comparatively thin, uneven and
riddled with hearth depressions. About 42% of
the layer V ornaments occur within hearths, and
many of these ornaments are burned (Table 5a).
The frequent presence of original (biological)
pigment in the shells is noteworthy (natural color
variable in Table 5b) and suggests a favorable
preservation environment. Vertically, the frequency of this phenomenon is highest at 85–115
and 140–160 cm below datum. The retention of
natural coloration in shells is strongly and positively correlated to the presence of cord wear
throughout the cultural layers (p = 0.0001). It
seems that the longer an ornament remained
strung or attached to another object, the better the
chance that its more transient natural properties
would also be preserved. This relation suggests
the existence of one or more stable microenvironments within the deposits that enhanced shell
preservation. Neither natural color retention nor
cord wear is concentrated in any particular square,
but the large horizontal (1 × 1 m) units of the excavation do not allow us to distinguish random
from finely structured distributions in the sediments.
Finally, ornaments are rare items in Klissoura
Cave 1 if considered against the total number of
chipped stone artifacts in each layer (Table 8). Ornaments are formal artifacts, however, and thus it
seems appropriate to consider their numbers in relation to chipped stone tools in particular. Ornaments are comparatively few even in relation to
the numbers of tools, but their proportions are noticeably greater in layers III’ and below, and especially in layer IV. These and other observations
suggest that the occupations were more intense
during the Aurignacian.
CONCLUSIONS
Prehistoric shell ornaments were mere particles in larger decorative formulae, and in Paleolithic sites they usually are found mixed with
other cultural debris. Lost to archaeologists in
most instances are the rules of combination of
these objects on strings, surfaces or in deliberately
assembled caches. The degree to which ornaments were recruited for complex symbolic representations is unclear and no doubt varied greatly.
Their basic function for conveying simple visual
Shell ornaments from the Upper Paleolithic
299
Table 7
Number of shell ornaments by square at 5-199 cm below datum in Klissoura Cave 1
Spit
A1
A2
A3
AA1
AA2
AA3
AA4
BB1
BB2
BB3
BB4
CC1
CC2
CC3
5-20
-
-
-
-
-
1
-
-
-
-
-
-
-
1
Layer(s)
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 column refers to the top of each ~5 cm spit. Gray shading indicates the presence of one or more hearth features.
Layer designations are generalized, since some are discontinuous (e.g. Layer V) or are not necessarily lie perfectly horizontal
across units. Because the sequence of excavation squares in the table cannot correspond to the original grid layout, squares are
listed alphanumerically.
messages about personal state (Kuhn and Stiner,
2007) or affiliation (Vanhaeren and d’Errico,
2006) may have been nearly universal, however,
and likely explains why small ornaments are the
most widespread and common art form of the
later Paleolithic in Eurasia and Africa.
Even in a disassembled or scattered state,
Paleolithic ornaments can provide information
300
M. C. Stiner
Fig. 8. The distributions of limestone rocks (white) and hearth features (black and dark gray) in 5 cm cuts at 145
through 180 cm below datum. The inferred shelter feature is apparent in cuts 150–175, in the area where rocks are
common but hearths consistently absent. Light gray background represents sedimentary matrix
Shell ornaments from the Upper Paleolithic
301
Table 8
Density of ornaments in relation to all chipped stone artifacts and to chipped stone tools by layer or
layer group
Layer(s)
N chipped stone N chipped stone
artifacts
tools
Percent
groundstone
of all lithics
MNI ornaments
Ornaments/
all lithics
Ornaments/
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
Percent groundstone artifacts is calculated based on all lithic artifacts in layer. Lithic data provided by Kaczanowska et al. (this issue).
about the contexts of decorative behavior, patterns of human selectivity and geography of these
preferences, and how the native biota were woven
into local or generalized expressions of human
identity (Stiner, 2003). Local biotic communities
exerted powerful background effects on the composition of ornaments made from animal skeletal
materials. Humans filtered through this natural
background with their own strong preferences for
shell shape, color, size, and natural rarity. These
patterns of selectivity are surprisingly consistent
through the Upper Paleolithic across Mediterranean and inland regions of Eurasia, while species
or types of raw materials used for bead-making
frequently were substituted (White, 1989; Taborin, 1993; Stiner, 2003).
The early Upper Paleolithic ornament assemblages from Klissoura Cave 1 are most similar to
those from Adriatic sites with respect to favored
mollusk species. The narrow size distribution of
the Klissoura 1 shells represents another point of
similarity to ornament assemblages from coastal
Italy but also to Levantine Turkey, and to carved
non-shell ornaments from the interior of France
and Germany.
The shell ornaments from Klissoura 1 are distinguished by their refined contents if compared
to assemblages from coastal sites. Though never
far from the sea, Klissoura 1 was always an inland
site. Marine shells suitable for ornament making
were not readily at hand, and few if any of the ornaments were manufactured on site. There is also
considerable evidence of “high-grading” or selec-
tive winnowing of the ornamental shells from
Klissoura 1 for harmony in color, form and quality. One finds few if any hints of child’s play from
the raw materials used, or the miscellaneous junk
so typical of coastal assemblages. Instead, nicely
finished objects were the rule. The prevalence of
cord-wear suggests that many of the ornaments
arrived already attached to organic materials or
human bodies. What breakage occurred to the ornamental shells reflects damage from long-term
use rather than errors in manufacture.
The ornament assemblage from the earliest
Aurignacian (layer IV) is the richest and the most
instructive about site function of the period. This
assemblage is quite large, and the diversity of its
contents is exceptionally high, even after corrections for sample size differences among layers.
Most of the ornaments in layer IV occur within
what appears to have been a man-made shelter.
This hearth-free feature is defined by a jumbled
ring of large stones around a thin organic stain,
and is surrounded by many hearths. The ornaments from inside the shelter feature may have
been attached to one or more leather or textile objects that once lined the floor of the structure.
As is generally true of Upper Paleolithic ornaments in Europe (Koz³owski and Otte, 2000),
those from Klissoura Cave 1 are well developed
in character and appear suddenly in the stratigraphic series. Distinct, well-stratified Mousterian industries occur in all layers below VI.
There are no ornaments in the Middle Paleolithic
layers except in VI an VII immediately below the
M. C. Stiner
302
shelter feature of layer IV. Layer V has an intermittent distribution and does not extend under the
area in layer IV that contains the shelter feature.
Instead, layer IV meets VI in this area of the excavation. The spatial distribution of the shell ornaments in layers VI–VII and young dates on some
of the shell specimens are almost certainly the
result of localized intrusions and mixing from
above.
The earliest Upper Paleolithic industry in the
Klissoura 1 stratigraphic series comes from layer
V and associates with an unambiguous shell ornament assemblage. Dated to greater than ca. 39,000
years BP, the lithic industry is distinguished by a
high incidence of lunates, resembling Uluzzian
industries in Italy, and it is clearly an Upper Paleolithic technology. The ornament assemblages
from all of the early Upper Paleolithic layers of
Klissoura 1 show general taxonomic affinities
with those from some older and coeval Aurignacian sites in Italy. The small ornament assemblage from layer V resembles that from Grotta del
Cavallo in Italy, due simply to the high incidence
of Dentalium shells.
Variation in the taxonomic content of the
shell ornament assemblages from Klissoura Cave
1 also speaks to climate-driven changes in environment heterogeneity in southern Greece during
the Late Pleistocene. The great number of species
represented in the early Upper Paleolithic assem-
blages reflects a mosaic of habitats that was more
complex than exists in the Peloponnese today.
Longer occupations, as is suggested for the lower
Aurignacian occupation, might tend to amplify
taxonomic diversity quasi-independently of environmental change, but this is not sufficient to account for the differences in species richness. The
early Upper Paleolithic ornaments from Klissoura 1 greatly exceed the taxonomic diversity of
ornamental shells both from the younger layers of
this site and those from all of the late Paleolithic
and Mesolithic layers in Franchthi Cave (Shackleton, 1988). The reduction in taxonomic diversity
after the LGM was almost certainly linked to the
global rise in sea level, which drowned the inland
lakes, raised water tables and swamped many
shorelines.
Acknowledgments
I am very grateful to André Carlo Colonese (Universit´ degli Studi di Firenze) and Giuseppe Manganelli (Università di Sienna) for their assistance in clarifying the paleontological context of the Pliocene fossil
shells from Klissoura Cave 1. I also thank the members
of the Klissoura 1 excavation team for access of geological data, site maps, and artifactual data, and the volunteers from the community of Berbati for their generosity and logistical assistance during the study. Gideon
Hartman (Harvard University) generously made many
of the photographs used in this manuscript.
Appendix 1: Species abundance (NISP and MNI) in the intact Upper Paleolithic through Mesolithic
layers of Klissoura Cave 1, and for the interface between MP and UP deposits (VII-VI)
3-5a
3, 5a
IIa-d
IIa-d
III-III'
III-III'
III''
III''
IIIc
IIIc
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
TOTAL
6
5
9
9
38
38
23
23
23
23
a. Layers 3-5 to IIIc
Taxon
Shell ornaments from the Upper Paleolithic
b. Layers IIIe-g to VII
Taxon
303
IIIe-g
IIIe-g
IV
IV
V
V
VI-VII
VI-VII
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. umbilicus
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 articulate
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 reticulate
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
Conus mediterraneus
1
1
4
4
0
0
0
0
Indet. marine gastropod
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
M. C. Stiner
304
b. Layers IIIe-g to VII
IIIe-g
IIIe-g
IV
IV
V
V
VI-VII
VI-VII
NISP
MNI
NISP
MNI
NISP
MNI
NISP
MNI
Taxon
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
148
138
1300
1218
34
32
56
TOTAL
(*) fossil species of Pliocene age, originating from lake bed or lakeshore deposits, source locality undetermined
53
Appendix 2: Ecological summary of Late Pleistocene molluskan taxa used as ornaments
at Klissoura Cave 1, Greece
Family
Genus species
Name source
Common name
Diet
Substrate
size (mm)
Adult
SCAPHOPODA (Class)
Dentaliidae
Dentalium dentale
L.
tusk
C
s
35-50
Dentalium vulgare
DaCosta
tusk
C
m,s
35-50
60-75
GASTROPODA (Class)
ARCHAEOGASTROPODA (Order)
Haliotidae
Haliotis lamellosa
Lamarck
abalone, ormer
HA
r
Gibbula adansoni
Payr.
top
HD
r,s,w
8-15
Gmelin
top
HD
–
10-24
Payr.
top
HD
r,s,w
8-15
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
Born
checkered top
HD
r
20-35
L.
red turban
HA
r,w
3-7
Gibbula albida
Gibbula richardi
Trochidae
Clanculus cruciatus
Monodonta turbinata
Turbinidae
Homalopoma sanguineum
MESOGASTROPODA (Order)
Littorinidae
Littorina neritoides
L.
periwinkle
H
r
5-7
Turritellidae
Turritella communis
Lamarck
turrit shell
HD
g,m,s
20-45
Cerithium rupestre
Risso
horn shell
HD
m,w,s
20-35
Cerithium vulgatum
Brug.
horn shell
HD
m,w,s
20-65
Naticarius= Neverita josephina
Risso
moon snail
C
s,g,m
25-40
Lamarck
moon snail
C
s
30-45
Born
helmet shell
Curch
s
60-70
Cerithiidae
Naticidae
Cassididae
Naticarius millepunctata + others
Phalium = Cassis undulatum
NEOGASTROPODA (Order)
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
spotted pisania
C,SC
r
15-32
Muricidae
Hexaplex = Murex trunculus
–
–
C
–
–
Cyclope neritea1
L.
mud snail
C-SC
s,m
8-17
Renieri
mud snail
C-SC
r,s
8-15
L.
basket whelk
O
s,m
18-28
Brug.
cone shell
C
r,w
60-65
Nassariidae
Nassarius = Hinia costulata
Sphaeronassa mutabilis
Conidae
Conus mediterraneus
Shell ornaments from the Upper Paleolithic
305
Appendix 2 continued
Family
Genus species
Name source
Common name
Substrate
size (mm)
Diet
Adult
MESOGASTROPODA (Order)
Littorinidae
Littorina neritoides
L.
periwinkle
H
r
5-7
Turritellidae
Turritella communis
Lamarck
turrit shell
HD
g,m,s
20-45
Cerithium rupestre
Risso
horn shell
HD
m,w,s
20-35
Cerithium vulgatum
Brug.
horn shell
HD
m,w,s
20-65
Naticarius= Neverita josephina
Risso
moon snail
C
s,g,m
25-40
Lamarck
moon snail
C
s
30-45
Born
helmet shell
Curch
s
60-70
Cerithiidae
Naticidae
Cassididae
Naticarius millepunctata + others
Phalium = Cassis undulatum
NEOGASTROPODA (Order)
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
spotted pisania
C,SC
r
15-32
Muricidae
Hexaplex = Murex trunculus
–
–
C
–
–
Cyclope neritea1
L.
mud snail
C-SC
s,m
8-17
Renieri
mud snail
C-SC
r,s
8-15
L.
basket whelk
O
s,m
18-28
Brug.
cone shell
C
r,w
Conus mediterraneus
FRESH- AND BRACKISH WATER MOLLUSKS (lakes, ponds, slow moving rivers)
60-65
Nassariidae
Nassarius = Hinia costulata
Sphaeronassa mutabilis
Conidae
GASTROPODA (Class)
Neritidae
Theodoxus cf. jordani
Theodoxus cf. fluviatalis
Sowerby
river nerite
C
s,m
7-9
–
river nerite
C
s,m
7-10
BIVALVIA or PELECYPODA (Class)
FILIBRANCHIA (Order)
Glycymeridae
Pectinidae
Glycymeris = Pectunculus sp.
–
bittersweet
F
s,m,g
35-65
Pecten maximus
L.
giant scallop
F
r,s
100-150
F
s,m,g
25-90
EULAMELIBRANCHIA (Order)
Cardiidae
L.
Acanthocardia tuberculatum
cockle shell
1
L.
edible cockle
F
s,m,g
30-50
Cerastoderma edule
Notes: Pliocene fossil taxa are not included in this table. (¹) tolerates or prefers brackish water. Substrate codes: (r) rock and other
firm surfaces, (m) mud, (f) floating matter and bubbles, (s) sand, (v) varied, (w) weeds, (c) corals, (g) gravel or coarse sand, (sp)
sponges, (r/s) adheres to hard surfaces as juvenile but free swimming as adult. Nomenclature: (L.) Linnaeus; (Blainv.) Blainville;
(Monter.) Monterosato; (Payr.) Payraudeau; (Brug.) BruguiÀre. Molluskan diet codes: (H) herbivore, (O) omnivore, (D) detritivore, (SC) scavenger, (F) filter or suspension feeder, (C) carnivore.
Appendix 3: Relative frequencies of natural and human-caused damage to a sample of mollusk shells
found within well-defined hearths in Layer IV
Genus
(if > 5 MNI)
NISP
MNI
Shell com- wave-worn pre-dated burned perforated holebroken
examined examined pleteness
%
%
%
%
%
with cord with
wear
ochre
%
%
Homalopoma
15
14
0.93
93
20
40
100
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
59
30
0.93
37
0
25
100
34
0
4
124
116
0.93
38
3
29
100
28
6
Other genera
0
3
All taxa
Listings are by genera for common types, except G. albida, which is significantly larger than all other species of Gibbula in the ornament assemblages
306
M. C. Stiner
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